Handbook of Parenting

HANDBOOK OF PARENTING
This highly anticipated third edition of the Handbook of Parenting brings together an array of field-
leading experts who have worked in different ways toward understanding the many diverse aspects
of parenting. Contributors to the Handbook look to the most recent research and thinking to shed
light on topics every parent, professional, and policy maker wonders about. Parenting is a perennially
“hot” topic. After all, everyone who has ever lived has been parented, and the vast majority of people
become parents themselves. No wonder bookstores house shelves of “how-to” parenting books,
and magazine racks in pharmacies and airports overflow with periodicals that feature parenting
advice. However, almost none of these is evidence-based. The Handbook of Parenting is. Period.
Each chapter has been written to be read and absorbed in a single sitting, and includes historical
considerations of the topic, a discussion of central issues and theory, a review of classical and modern
research, and forecasts of future directions of theory and research. Together, the five volumes in the
Handbook cover Children and Parenting, the Biology and Ecology of Parenting, Being and Becoming
a Parent, Social Conditions and Applied Parenting, and the Practice of Parenting.
Volume 2, Biology and Ecology of Parenting, relates parenting to its biological roots and sets parenting
in its ecological framework. Some aspects of parenting are influenced by the organic makeup of human
beings, and the chapters in Part I, on the Biology of Parenting, examine the evolution of parenting, the
psychobiological determinants of parenting in nonhumans, and primate parenting, as well as the genetic,
prenatal, neuroendocrinological, and neurobiological bases of human parenting. A deep understanding of
what it means to parent also depends on the ecologies in which parenting takes place. Beyond the nuclear
family, parents are embedded in, influence, and are themselves affected by larger social systems. The chapters
in Part II, on the Ecology of Parenting, examine the ancient and modern histories of parenting as well as
epidemiology, neighborhoods, educational attainment, socioeconomic status, culture, and environment to
provide an overarching relational developmental contextual systems perspective on parenting.
Marc H. Bornstein holds a BA from Columbia College, MS and PhD degrees from Yale University,
and honorary doctorates from the University of Padua and University of Trento. Bornstein is President
of the Society for Research in Child Development and has held faculty positions at Princeton University
and New York University as well as academic appointments in Munich, London, Paris, New York,
Tokyo, Bamenda, Seoul, Trento, Santiago, Bristol, and Oxford. Bornstein is author of several children’s
books, videos, and puzzles in The Child’s World and Baby Explorer series, Editor Emeritus of Child
Development and founding Editor of Parenting: Science and Practice, and consultant for governments,
foundations, universities, publishers, scientific journals, the media, and UNICEF. He has published
widely in experimental, methodological, comparative, developmental, and cultural science as well as
neuroscience, pediatrics, and aesthetics.
HANDBOOK OF PARENTING
Volume 2: Biology and
Ecology of Parenting
Third Edition
Edited by Marc H. Bornstein

Third edition published 2019
by Routledge
52 Vanderbilt Avenue, New York, NY 10017
and by Routledge
2 Park Square, Milton Park, Abingdon, Oxon, OX14 4RN
Routledge is an imprint of the Taylor & Francis Group, an informa business
© 2019 Taylor & Francis
The right of Marc H. Bornstein to be identified as the author of the
editorial material, and of the authors for their individual chapters, has been
asserted in accordance with sections 77 and 78 of the Copyright, Designs
and Patents Act 1988.
All rights reserved. No part of this book may be reprinted or reproduced or
utilised in any form or by any electronic, mechanical, or other means, now
known or hereafter invented, including photocopying and recording, or in
any information storage or retrieval system, without permission in writing
from the publishers.
Trademark notice: Product or corporate names may be trademarks or
registered trademarks, and are used only for identification and explanation
without intent to infringe.
First edition published by Laurence Erlbaum Associates 1995
Second edition published by Taylor & Francis 2002
Library of Congress Cataloging-in-Publication Data
A catalog record for this book has been requested
ISBN: 978-1-138-22868-9 (hbk)
ISBN: 978-1-138-22869-6 (pbk)
ISBN: 978-0-429-40145-9 (ebk)
Typeset in Bembo
by Apex CoVantage, LLC
For Marian and Harold Sackrowitz
CONTENTS
Preface to the Third Edition ix

About the Editor xiv
About the Contributors xvi
PART I
Biology of Parenting 1
1 The Evolution of Parenting and Evolutionary Approaches to Childrearing 3

David F. Bjorklund and Alyson J. Myers
2 Psychobiology of Maternal Behavior in Nonhuman Mammals 30

Aya Dudin, Patrick O. McGowan, Ruiyong Wu, Alison S. Fleming,
and Ming Li
3 Parenting in Nonhuman Primates 78

Kim A. Bard
4 Genetics and Parenting 123

Amanda V. Broderick and Jenae M. Neiderhiser
5 Prenatal Parenting 166

David A. Coall, Anna C. Callan, Julie Sartori, and James S. Chisholm
6 The Social Neuroendocrinology of Human Parenting 220

Ruth Feldman
7 Neurobiology of Human Parenting 250

Eloise A. Stark, Alan Stein, Katherine S. Young, Christine E. Parsons,
and Morten L. Kringelbach
vii
Contents
PART II
Ecology of Parenting 285
8 Ancient History of Parenting 287

Valerie French
9 Modern History of Parenting 320

Peter N. Stearns
10 Epidemiology of Parenting 349

Rebecca M. Pearson, Miguel Cordero, and Priya Rajyaguru
11 Neighborhoods and Parenting 371

Elizabeth A. Shuey and Tama Leventhal
12 Parent Education Attainment and Parenting 400

Pamela E. Davis-Kean, Sandra Tang, and Nicholas E. Waters
13 Socioeconomic Status and Parenting 421

Erika Hoff and Brett Laursen
14 Culture and Parenting 448

Xinyin Chen, Rui Fu, and Wai Ying Vivien Yiu
15 Environment and Parenting 474

Robert H. Bradley
Index519
viii
PREFACE TO THE THIRD EDITION
Previous editions of the Handbook of Parenting have been called the “who’s who of the what’s what.”
This third edition of this Handbook appears at a time that is momentous in the history of parent-
ing. The family generally, and parenting specifically, are today in a greater state of flux, question, and
re-definition than perhaps ever before. We are witnessing the emergence of striking permutations on
the theme of parenting: blended families, lesbian and gay parents, teen versus fifties first-time moms
and dads, genetic versus social parents. One cannot but be awed on the biological front by technology
that now renders postmenopausal women capable of childbearing and with the possibility of parents
designing their babies. Similarly, on the sociological front, single parenthood is a modern-day fact of
life, adult child dependency is on the rise, and even in the face of rising institutional demands to take
increasing responsibility for their offspring parents are ever less certain of their roles and responsibili-
ties. The Handbook of Parenting is concerned with all these facets of parenting . . . and more.
Most people become parents, and everyone who ever lived has had parents, still parenting remains
a mystifying subject. Who is ultimately responsible for parenting? Does parenting come naturally,
or must parenting be learned? How do parents conceive of parenting? of childhood? What does it
mean to parent a preterm baby, twins, or a child on the autistic spectrum? to be an older parent, or
one who is divorced, disabled, or drug abusing? What do theories (psychoanalysis, personality theory,
attachment, and behavior genetics, for example) contribute to our understanding of parenting? What
are the goals parents have for themselves? for their children? What functions do parents’ cognitions
serve? What are the aims of parents’ practices? What accounts for parents believing or behaving in
similar ways? Why do so many attitudes and actions of parents differ so? How do children influence
their parents? How do personality, knowledge, and worldview affect parenting? How do social class,
culture, environment, and history shape parenthood? How can parents effectively relate to child care,
schools, and their children’s pediatricians?
These are many of the questions addressed in this third edition of the Handbook of Parenting . . .
for this is an evidenced-based volume set on how to parent as much as it is one on what being a parent
is all about.
Put succinctly, parents create people. They are entrusted with preparing their offspring for the
physical, psychosocial, and economic conditions in which their children eventually will fare and
hopefully will flourish. Amid the many influences on each next generation, parents are the “final
common pathway” to children’s development and stature, adjustment and success. Human social
inquiry—antedating even Athenian interest in Spartan childrearing practices—has always, as a matter
of course, included reports of parenting. Freud opined that childrearing is one of three “impossible
ix
Preface to the Third Edition
professions”—the other two being governing nations and psychoanalysis. One encounters as many
views as the number of people one asks about the relative merits of being an at-home or a working
mother, about what mix of day care, family care, or parent care is best for a child, about whether good
parenting reflects intuition or experience.
The Handbook of Parenting concerns itself with different types of parents—mothers and fathers,
single, adolescent, and adoptive parents; with basic characteristics of parenting—knowledge, beliefs,
and expectations about parenting—as well as the practice of parenting; with forces that shape
parenting—employment, social class, culture, environment, and history; with problems faced by
parents—handicap, marital difficulties, drug addiction; and with practical concerns of parenting—
how to promote children’s health, foster social adjustment and cognitive competence, and interact
with educational, legal, and religious institutions. Contributors to the Handbook of Parenting have
worked in different ways toward understanding all these diverse aspects of parenting, and all look to
the most recent research and thinking in the field to shed light on many topics every parent, profes-
sional, and policy maker wonders about.
Parenthood is a job whose primary object of attention and action is the child. But parenting
also has consequences for parents. Parenthood is giving and responsibility, and parenting has its own
intrinsic pleasures, privileges, and profits as well as frustrations, fears, and failures. Parenthood can
enhance psychological development, self-confidence, and sense of well-being, and parenthood also
affords opportunities to confront new challenges and to test and display diverse competencies. Par-
ents can derive considerable and continuing pleasure in their relationships and activities with their
children. But parenting is also fraught with small and large stresses and disappointments. The transi-
tion to parenthood is daunting, and the onrush of new stages of parenthood is relentless. In the final
analysis, however, parents receive a great deal “in kind” for the hard work of parenting—they can be
recipients of unconditional love, they can gain skills, and they can even pretend to immortality. This
third edition of the Handbook of Parenting reveals the many positives that accompany parenting and
offers resolutions for its many challenges.
The Handbook of Parenting encompasses the broad themes of who are parents, whom parents
parent, the scope of parenting and its many effects, the determinants of parenting, and the nature,
structure, and meaning of parenthood for parents. The third edition of the Handbook of Parenting is
divided into five volumes, each with two parts:
CHILDREN AND PARENTING is Volume 1 of the Handbook. Parenthood is, perhaps first
and foremost, a functional status in the life cycle: Parents issue as well as protect, nurture, and
teach their progeny even if human development is too subtle and dynamic to admit that paren-
tal caregiving alone determines the developmental course and outcome of ontogeny. Volume
1 of the Handbook of Parenting begins with chapters concerned with how children influence
parenting. Notable are their more obvious characteristics, like child age or developmental
stage; but more subtle ones, like child gender, physical state, temperament, mental ability, and
other individual differences factors, are also instrumental. The chapters in Part I, on Parenting
Across the Lifespan, discuss the unique rewards and special demands of parenting children of
different ages and stages—infants, toddlers, youngsters in middle childhood, and adolescents—
as well as the modern notion of parent-child relationships in emerging adulthood and adult-
hood and old age. The chapters in Part II, on Parenting Children of Varying Status, discuss
common issues associated with parenting children of different genders and temperaments as
well as unique situations of parenting adopted and foster children and children with a variety
of special needs, such as those with extreme talent, born preterm, who are socially withdrawn
or aggressive, or who fall on the autistic spectrum, manifest intellectual disabilities, or suffer a
chronic health condition.
x
BIOLOGY AND ECOLOGY OF PARENTING is Volume 2 of the Handbook. For parent-

ing to be understood as a whole, biological and ecological determinants of parenting need to
be brought into the picture. Volume 2 of the Handbook relates parenting to its biological roots
and sets parenting in its ecological framework. Some aspects of parenting are influenced by
the organic makeup of human beings, and the chapters in Part I, on the Biology of Parent-
ing, examine the evolution of parenting, the psychobiological determinants of parenting in
nonhumans, and primate parenting and then the genetic, prenatal, neuroendocrinological, and
neurobiological bases of human parenting. A deep understanding of what it means to par-
ent also depends on the ecologies in which parenting takes place. Beyond the nuclear family,
parents are embedded in, influence, and are themselves affected by larger social systems. The
chapters in Part II, on the Ecology of Parenting, examine the ancient and modern histories
of parenting as well as epidemiology, neighborhoods, educational attainment, socioeconomic
status, culture, and environment to provide an overarching relational developmental contextual
systems perspective on parenting.
BEING AND BECOMING A PARENT is Volume 3 of the Handbook. A large cast of
characters is responsible for parenting, each has her or his own customs and agenda, and the
psychological characteristics and social interests of those individuals are revealing of what
parenting is. Chapters in Part I, on The Parent, show just how rich and multifaceted is the
constellation of children’s caregivers. Considered first are family systems and then successively
mothers and fathers, coparenting and gatekeeping between parents, adolescent parenting,
grandparenting, and single parenthood, divorced and remarried parenting, lesbian and gay
parents, and finally sibling caregivers and nonparental caregiving. Parenting also draws on
transient and enduring physical, personality, and intellectual characteristics of the individual.
The chapters in Part II, on Becoming and Being a Parent, consider the intergenerational
transmission of parenting, parenting and contemporary reproductive technologies, the transi-
tion to parenthood, and stages of parental development, and then chapters turn to parents’
well-being, emotions, self-efficacy, cognitions, attributions, as well as socialization, personal-
ity in parenting, and psychoanalytic theory. These features of parents serve many functions:
They generate and shape parental practices, mediate the effectiveness of parenting, and help
to organize parenting.
SOCIAL CONDITIONS AND APPLIED PARENTING is Volume 4 of the Handbook.
Parenting is not uniform across communities, groups, or cultures; rather parenting is subject
to wide variation. Volume 4 of the Handbook describes socially defined groups of parents and
social conditions that promote variation in parenting. The chapters in Part I, on Social and
Cultural Conditions of Parenting, start with a relational developmental systems perspective
on parenting and move to considerations of ethnic and minority parenting among Latino
and Latin Americans, African Americans, Asians and Asian Americans, Indigenous parents,
and immigrant parents. The section concludes with the roles of employment and of poverty
on parenting. Parents are ordinarily the most consistent and caring people in children’s lives.
However, parenting does not always go right or well. Information, education, and support
programs can remedy potential ills. The chapters in Part II, on Applied Issues in Parenting,
begin with how parenting is measured and follow with examinations of maternal deprivation,
attachment, and acceptance/rejection in parenting. Serious challenges to parenting—some
common, such as stress, depression, and disability, and some less common, such as substance
abuse, psychopathology, maltreatment, incarceration—are addressed as are parenting interven-
tions intended to redress these trials.
THE PRACTICE OF PARENTING is Volume 5 of the Handbook. Parents meet the bio-
logical, physical, and health requirements of children. Parents interact with children socially.
xi
Parents stimulate children to engage and understand the environment and to enter the world
of learning. Parents provision, organize, and arrange their children’s home and local environ-
ments and the media to which children are exposed. Parents also manage child development
vis-à-vis childcare, school, the circles of medicine and law, as well as other social institutions
through their active citizenship. Volume 5 of the Handbook addresses the nuts-and-bolts of
parenting as well as the promotion of positive parenting practices. The chapters in Part I, on
Practical Parenting, review the ethics of parenting, parenting and the development of children’s
self-regulation, discipline, prosocial and moral development, and resilience as well as children’s
language, play, cognitive, and academic achievement and children’s peer relationships. Many
caregiving principles and practices have direct effects on children. Parents indirectly influence
children as well, for example, through relations they have with their local or larger commu-
nities. The chapters in Part II, on Parents and Social Institutions, explore parents and their
children’s childcare, activities, media, schools, and healthcare and examine relations between
parenthood and the law, public policy, and religion and spirituality.
Each chapter in the third edition of the Handbook of Parenting addresses a different but central
topic in parenting; each is rooted in current thinking and theory as well as classical and modern
research on a topic; each is written to be read and absorbed in a single sitting. Each chapter in this
new Handbook follows a standard organization, including an introduction to the chapter as a whole,
followed by historical considerations of the topic, a discussion of central issues and theory, a review
of classical and modern research, forecasts of future directions of theory and research, and a set of
evidence-based conclusions. Of course, each chapter considers contributors’ own convictions and
findings, but contributions to this third edition of the Handbook of Parenting attempt to present all
major points of view and central lines of inquiry and interpret them broadly. The Handbook of Parent-
ing is intended to be both comprehensive and state-of-the-art. To assert that parenting is complex is
to understate the obvious. As the expanded scope of this third edition of the Handbook of Parenting
also amply attests, parenting is naturally and intensely interdisciplinary.
The Handbook of Parenting is concerned principally with the nature and scope of parenting per
se and secondarily with child outcomes of parenting. Beyond an impressive range of information,
readers will find passim typologies of parenting (e.g., authoritarian-autocratic, indulgent-permissive,
indifferent-uninvolved, authoritative-reciprocal), theories of parenting (e.g., ecological, psychoana-
lytic, behavior genetic, ethological, behavioral, sociobiological), conditions of parenting (e.g., gender,
culture, content), recurrent themes in parenting studies (e.g., attachment, transaction, systems), and
even aphorisms (e.g., “A child should have strict discipline in order to develop a fine, strong charac-
ter,” “The child is father to the man”).
Each chapter in the Handbook of Parenting lays out the meanings and implications of a contribu-
tion and a perspective on parenting. Once upon a time, parenting was a seemingly simple thing:
Mothers mothered. Fathers fathered. Today, parenting has many motives, many meanings, and many
manifestations. Contemporary parenting is viewed as immensely time consuming and effortful. The
perfect mother or father or family is a figment of false cultural memory. Modern society recognizes
“subdivisions” of the call: genetic mother, gestational mother, biological mother, birth mother, social
mother. For some, the individual sacrifices that mark parenting arise for the sole and selfish purpose
of passing one’s genes on to succeeding generations. For others, a second child may be conceived to
save the life of a first child. A multitude of factors influences the unrelenting advance of events and
decisions that surround parenting—biopsychosocial, dyadic, contextual, historical. Recognizing this
complexity is important to informing people’s thinking about parenting, especially information-
hungry parents themselves. This third edition of the Handbook of Parenting explores all these motives,
meanings, and manifestations of parenting.
xii
Each day more than three-quarters of a million adults around the world experience the rewards
and challenges, as well as the joys and heartaches, of becoming parents. The human race succeeds
because of parenting. From the start, parenting is a “24/7” job. Parenting formally begins before
pregnancy and can continue throughout the life span: Practically speaking for most, once a parent,
always a parent. Parenting is a subject about which people hold strong opinions, and about which too
little solid information or considered reflection exists. Parenting has never come with a Handbook . . .
until now.
—Marc H. Bornstein
xiii
ABOUT THE EDITOR
Marc H. Bornstein holds a BA from Columbia College, MS and PhD degrees from Yale Uni-
versity, and honorary doctorates from the University of Padua and University of Trento. Bornstein
was a J. S. Guggenheim Foundation Fellow and he received a Research Career Development Award
from the National Institute of Child Health and Human Development. He also received the C. S.
Ford Cross-Cultural Research Award from the Human Relations Area Files, the B. R. McCandless
Young Scientist Award and the G. Stanley Hall Award from the American Psychological Associa-
tion, a U.S. PHS Superior Service Award and an Award of Merit from the National Institutes of
Health, two Japan Society for the Promotion of Science Fellowships, four Awards for Excellence
from the American Mensa Education & Research Foundation, the Arnold Gesell Prize from the
Theodor Hellbrügge Foundation, the Distinguished Scientist Award from the International Society
for the Study of Behavioral Development, and both the Distinguished International Contributions
to Child Development Award and the Distinguished Scientific Contributions to Child Develop-
ment Award from the Society for Research in Child Development. Bornstein is President of the
Society for Research in Child Development and a past member of the SRCD Governing Council
and Executive Committee of the International Congress of Infancy Studies.
Bornstein has held faculty positions at Princeton University and New York University as well as
academic appointments as Visiting Scientist at the Max-Planck-Institut für Psychiatrie in Munich;
Visiting Fellow at University College London; Professeur Invité at the Laboratoire de Psychologie
Expérimentale in the Université René Descartes in Paris; Child Clinical Fellow at the Institute for
Behavior Therapy in New York; Visiting Professor at the University of Tokyo; Professeur Invité at
the Laboratoire de Psychologie du Développement et de l’Éducation de l’Enfant in the Sorbonne
in Paris; Visiting Fellow of the British Psychological Society; Visiting Scientist at the Human Devel-
opment Resource Centre in Bamenda, Cameroon; Visiting Scholar at the Institute of Psychology
in Seoul National University in Seoul, South Korea; Visiting Professor at the Faculty of Cognitive
Science in the University of Trento, Italy; Profesor Visitante at the Pontificia Universidad Católica
de Chile in Santiago, Chile; Institute for Advanced Studies Benjamin Meaker Visiting Professor,
University of Bristol; Jacobs Foundation Scholar-in-Residence, Marbach, Germany; Honorary Fel-
low, Department of Psychiatry, Oxford University; Adjunct Academic Member of the Council of the
Department of Cognitive Sciences, University of Trento, Italy; and International Research Fellow at
the Institute for Fiscal Studies, London.
xiv
About the Editor
Bornstein is coauthor of The Architecture of the Child Mind: g, Fs, and the Hierarchical Model of Intelligence,
Gender in Low- and Middle-Income Countries, Development in Infancy (5 editions), Development: Infancy
through Adolescence, Lifespan Development, Genitorialità: Fattori Biologici E Culturali Dell’essere Genitori,
and Perceiving Similarity and Comprehending Metaphor. He is general editor of The Crosscurrents in
Contemporary Psychology Series, including Psychological Development from Infancy, Comparative Methods in
Psychology, Psychology and Its Allied Disciplines (Vols. I–III), Sensitive Periods in Development, Interaction
in Human Development, Cultural Approaches to Parenting, Child Development and Behavioral Pediatrics,
and Well-Being: Positive Development Across the Life Course, and general editor of the Monographs in
Parenting series, including his own Socioeconomic Status, Parenting, and Child Development and Accultura-
tion and Parent-Child Relationships. He edited Maternal Responsiveness: Characteristics and Consequences,
the Handbook of Parenting (Vols. I–V, 3 editions), and the Handbook of Cultural Developmental Science
(Parts 1 and 2), and is Editor-in-Chief of the SAGE Encyclopedia of Lifespan Human Development.
He also coedited Developmental Science: An Advanced Textbook (7 editions), Stability and Continuity
in Mental Development, Contemporary Constructions of the Child, Early Child Development in the French
Tradition, The Role of Play in the Development of Thought, Acculturation and Parent-Child Relationships,
Immigrant Families in Contemporary Society, The Developing Infant Mind: Origins of the Social Brain, and
Ecological Settings and Processes in Developmental Systems (Volume 4 of the Handbook of Child Psychology
and Developmental Science). He is author of several children’s books, videos, and puzzles in The Child’s
World and Baby Explorer series. Bornstein is Editor Emeritus of Child Development and founding Edi-
tor of Parenting: Science and Practice. He has administered both Federal and Foundation grants, sits on
the editorial boards of several professional journals, is a member of scholarly societies in a variety
of disciplines, and consults for governments, foundations, universities, publishers, scientific journals,
the media, and UNICEF. He has published widely in experimental, methodological, comparative,
developmental, and cultural science as well as neuroscience, pediatrics, and aesthetics. Bornstein was
named to the Top 20 Authors for Productivity in Developmental Science by the American Educa-
tional Research Association.
xv
ABOUT THE CONTRIBUTORS
Kim A. Bard is a Professor of Comparative Developmental Psychology in the Department of

Psychology, University of Portsmouth, UK. She was educated at Wheaton College, Massachusetts
(BA with Honors) and Georgia State University (MA and PhD in Comparative/Developmental
Psychology). She was employed previously at Emory University as a Research Scientist at the Yerkes
Regional Primate Research Center and at the Clinical Developmental and Applied Research Pro-
gram of the Human Genetics Laboratory, and as a Research Fellow in the Department of Psychology.
Bard is an Associate Fellow of the British Psychological Society and Past President of the Primate
Society of Great Britain and European Federation for Primatology. Currently, she sits on the Advi-
sory Board of Primates, is an Associate Editor for Animal Cognition, and sits on the Editorial Boards of
American Journal of Primatology, Child Development Perspectives, and Emotion Review. Bard is the author
of Responsive Care: Behavioral Intervention for Nursery-Reared Chimpanzees and a co-editor of The Cul-
tural Nature of Attachment: Contextualizing Relationships and Development.
David F. Bjorklund is Professor of Psychology at Florida Atlantic University. He received a BA

in Psychology from the University of Massachusetts, an MA in Psychology from the University of
Dayton, and a PhD in Developmental Psychology from the University of North Carolina at Chapel
Hill. He is the author of Children’s Thinking: Cognitive Development and Individual Differences, Why
Youth Is Not Wasted on the Young, co-author of Looking at Children: An Introduction to Child Develop-
ment, Parents Book of Discipline, Applied Child Study and The Origins of Human Nature: Evolutionary
Developmental Psychology, Child and Adolescent Development: An Integrative Approach, and the General
Psychology; he is also editor of several books. He served as Associate Editor of Child Development and
the Journal of Experimental Child Psychology and is Editor of the Journal of Experimental Child Psychol-
ogy. Bjorklund is a contributing editor to Parents Magazine. His research interests include children’s
cognitive development and evolutionary developmental psychology.
Robert H. Bradley is Professor of Psychology and Director of the Center for Child and Family
Success at Arizona State University. Bradley was a faculty member at the University of Arkansas at
Little Rock. He served as associate editor for Child Development and Early Childhood Research Quar-
terly and chaired the Biobehavioral and Behavioral Research subcommittee at NIH. Bradley is one of
the developers of the HOME Inventory. He was an investigator in the NICHD Study of Early Child
Care and Youth Development and the Early Head Start Research and Evaluation Study.
xvi
About the Contributors
Amanda V. Broderick is a PhD student studying Child Clinical Psychology at The Pennsylvania
State University. Broderick earned her BA in psychology with a minor in applied statistics at the
University of Michigan and her MA at The Pennsylvania State University. Broderick uses com-
plementary research designs to understand interparental and parent-child relationships as conduits
through which risk for maladaptive child development is mitigated or conferred.
Anna C. Callan is Senior Lecturer in the School of Medical and Health Sciences at Edith Cowan
University, Perth, Western Australia. Callan was awarded her PhD from the University of Manches-
ter, where she was subsequently employed as a Research Associate, before working at the Telethon
Kids Institute and Edith Cowan University. Callan has expertise in the field of environmental epi-
demiology, with a particular interest in the health impacts of prenatal and early life exposures to
substances including persistent chemicals, plasticizers, and perfluoroalkyl acids.
Xinyin Chen is Professor of Psychology at the Graduate School of Education, the University
of Pennsylvania. He holds a BA from East China Normal University and MA and PhD from the
University of Waterloo. He received a Scholars Award from the William T. Grant Foundation and
several other awards for his scientific work. He served as the President of the International Society for
the Study of Behavioral Development. His research interest is mainly in children’s and adolescents’
socioemotional functioning, social relationships, and family socialization processes with a focus on
cross-cultural issues in Brazil, Canada, China, Italy, and the United States. He has edited or co-edited
Peer Relationships in Cultural Context, Social Change and Human Development, Socioemotional Develop-
ment in Cultural Context, and Values, Religion, and Culture in Adolescent Development.
James S. Chisholm is Emeritus Professor in the School of Anatomy, Physiology, and Human Biol-
ogy at the University of Western Australia in Perth. He received his BA from Wesleyan University
and MPhil and PhD from Rutgers University. He has conducted fieldwork among the Navajo and
Aboriginal people in Arnhem Land and taught at the University of New Mexico, the University
of California, Davis, and the University of Western Australia. He is the author of Navajo Infancy:
An Ethological Study of Child Development and Death, Hope and Sex: Steps to an Evolutionary Ecology of
Mind and Morality.
David A. Coall is a Senior Lecturer in the School of Medical and Health Sciences at Edith Cowan
University and Adjunct Research Fellow in the School of Medicine, The University of West-
ern Australia. He completed his PhD at The University of Western Australia and a Postdoctoral
Research Fellowship in the Institute of Psychology, University of Basel, Switzerland. His research
focuses on intergenerational influences on health, such as the influence the mother’s environment
has on placental development and pregnancy outcomes and the influence grandparents have in the
lives of grandchildren. Coall has authored publications in evolutionary anthropology, pediatrics, cog-
nitive science, mental health, and maternal and child health.
Miguel Cordero is a child clinical psychologist with a MA in Behavioural Neurosciences at the

University of Chile. He was a consultant in perinatal and pediatric mental health at Dr. Sotero del
Rio Hospital in Santiago, Chile in charge of the implementation of Chile Crece Contigo at the
Ministry of Health, a large-scale policy to reduce early childhood development inequalities. He
was also responsible for implementing a large multi-center randomized trial to assess the impacts
of implementing “Nobody’s Perfect Parenting Program” in primary health clinics from Chile.
Currently, he is a PhD student at the Population Health Sciences Institute at the Bristol Medical
School.
xvii
Pamela E. Davis-Kean is a Professor of Psychology and Research Professor of the Institute for
Social Research at the University of Michigan where she is the Program Director of the Population,
Neurodevelopment, and Genetics program. She received her BA from Florida State University and
her MA and PhD from Vanderbilt University. Davis-Kean is a Fellow of the Association of Psycho-
logical Science. She is an Associate Editor of Advances in Methods and Practices in Psychological Science.
Davis-Kean’s primary research focus is on parental educational attainment and how it can influence
the development of the home environment throughout childhood, adolescence, and the transition to
adulthood. Davis-Kean is co-editor of Socializing Children through Language.
Aya Dudin is a PhD student in the McMaster Integrative Neuroscience Discovery and Study pro-
gram. Dudin completed an HBSc in Behavior Genetics and Neurobiology from the University of
Toronto. She is interested in understanding the underlying neurobiology of maternal behavior in
mammals. Aya has co-authored work in Social Neuroscience as well as Depression and Anxiety.
Ruth Feldman is the Simms-Mann Professor and Director of the Center for Developmental and
Social Neuroscience at the Interdisciplinary Center, Herzlia, with a joint appointment at Yale Uni-
versity Child Study Center. Her research focuses on the neurobiology of affiliation, biobehavioral
synchrony, longitudinal follow-up of high-risk infants, and the effects of touch-based interventions.
She studies the role of oxytocin in human bonding, the parental brain, kangaroo contact, the neu-
robiology of conflict resolution, the effects of maternal depression, and the development of targeted
interventions.
Alison S. Fleming is a Professor Emerita of Psychology at the University of Toronto. Fleming

received her PhD from The Institute of Animal Behavior, Rutgers University. Fleming focused her
research efforts on understanding what makes a mother want to mother and what influences how she
does so. This work has been taken to multiple levels of explanation in both rodent and humans, rang-
ing from the role of hormones, odors, and other sensory cues from the young, past and present his-
tory and experiences, the brain, and the underlying genetic and neurochemical mechanisms. Fleming
is the recipient of the University of Toronto Excellence in Research Award and was inducted into
the Royal Society of Canada. She was awarded a Canada Research Chair in Neurobiology, and
she received the Daniel S. Lehrman Lifetime Achievement Award from the Society of Behavioral
Neuroendocrinology.
Valerie French was Associate Professor of History at the American University. She was active in
professional organizations on women’s and minority issues. She published extensively on ancient
childhood, women in antiquity, Alexander the Great, and Greek historiography. French coauthored
Historians and the Living Past: The Theory and Practice of Historical Study.
Rui Fu is a PhD candidate in the program of Human Development and Quantitative Methods at
the University of Pennsylvania. Her research interests focus on children’s and adolescents’ academic,
social, and psychological functioning from a cross-cultural perspective. She also serves as an Early
Career Scholar Representative of the SRCD Asian Caucus.
Erika Hoff is Professor of Psychology at Florida Atlantic University. Hoff earned an AB Ed from
the University of Michigan, an MS from Rutgers–The State University of New Jersey, and a PhD
from the University of Michigan. Her research addresses the relations among properties of children’s
early environments, their language experience, and their language development. She has studied
effects of maternal education and effects of dual language exposure on children’s language growth.
xviii
She is a member of the U.S. Bridging the Word Gap Research network, which focuses on inter-
ventions to remedy SES-related disparities in children’s early experience and language skills. She
has been associate editor of the Journal of Child Language and Child Development. She is currently
associate editor of the Journal of Experimental Child Psychology. She is editor of multiple books on
early language development, including Research Methods in Child Language and Childhood Bilingualism:
Research on Infancy Through School Age.
Morten L. Kringelbach is a Professor of Neuroscience at the University of Oxford, UK, and Uni-
versity of Aarhus, Denmark. His research uses neuroimaging and whole-brain computational models
of hedonia (pleasure) linked to infants, taste, and music to discover how to increase eudaimonia (the
life well-lived). He has published 14 books. He is a fellow of The Queen’s College, Oxford, of the
Association for Psychological Science, is on the advisory board of Scientific American, and is a board
member of the world’s first Empathy Museum.
Brett Laursen is Professor of Psychology and Director of Graduate Training at Florida Atlantic
University. He is also Docent Professor of Social Developmental Psychology at the University of
Jyväskylä, Finland. Laursen received his PhD and MA from the Institute of Child Development at
the University of Minnesota, and his BA from Nebraska Wesleyan University. He is a Fellow of the
American Psychology Association (Division 7, Developmental), a Fellow and Charter Member of
the Association for Psychological Sciences, and the recipient of an Honorary Doctorate from Örebro
University, Sweden. He is currently the Editor-in-Chief of the International Journal of Behavioral Devel-
opment. Laursen’s research focuses on parent-child and peer relationships during childhood and ado-
lescence and their influence on individual social and academic adjustment. His edited books include
Handbook of Developmental Research Methods and Handbook of Peer Interactions, Relationships and Groups.
Tama Leventhal is Professor in the Eliot-Pearson Department of Child Study and Human Devel-
opment at Tufts University. She received her PhD from Teachers College, Columbia University and
was previously affiliated with Johns Hopkins University. Her primary research focus is the role of
neighborhood contexts in the lives of children, youth, and families. Leventhal was formerly a U.S.
Department of Housing and Urban Development Postdoctoral Urban Scholar, a William T. Grant
Scholar, and a Foundation for Child Development Changing Faces of America’s Children Young
Scholar. She was Co-director of the MacArthur Network on Housing and Families with Children
and is currently Co-director of the Housing and Children’s Healthy Development study. She is Asso-
ciate Editor of the Journal of Research on Adolescence and Applied Developmental Science, Co-editor of
the Handbook of Child Psychology and Developmental Science: Volume 4: Ecological Settings and Processes in
Developmental Systems, Steering Committee Chair of the University-Based Child and Family Policy
Consortium, and Board of Directors of the Council on Contemporary Families.
Ming Li is Professor of Psychology at the University of Nebraska-Lincoln. Li was educated at the

Peking University and University of Toronto. He is a Fellow of Divisions 6 and 28 of the American
Psychological Association. He is a co‑editor of The Neuropsychopathology of Schizophrenia: Molecules,
Brain Systems, Motivation, and Cognition.
Patrick O. McGowan is Associate Professor of Biological Sciences at the University of Toronto

Scarborough, where he is Principal Investigator of the Laboratory of Environmental Epigenetics and
Development. He also holds faculty appointments in the departments of Cell and Systems Biology,
Psychology and Physiology at the University of Toronto. He received his PhD at Duke University
and was Sackler Postdoctoral fellow in Psychiatry at McGill University. He is an associate editor of
xix
Frontiers in Epigenomics and Epigenetics. McGowan’s research focuses on biological mechanisms linking
exposures to stressors, particularly early in life, with the programming of health and disease.
Alyson J. Myers is a PhD candidate in Developmental Psychology at Florida Atlantic University.

Myers received her MA at Florida Atlantic University and is affiliated with Denise Herzing and the
Wild Dolphin Project. She studies wild Atlantic spotted dolphin (Stenella frontalis) behavior, specifi-
cally synchrony and its development in juveniles. Her interests include ethology, evolutionary devel-
opmental psychology, and cognitive development.
Jenae M. Neiderhiser is a Distinguished Professor of Psychology and Human Development

and Family Studies at Pennsylvania State University. She is co-director of the Gene-Environment
Research Initiative and is a faculty affiliate of the Social Sciences Research Institute and the Child
Study Center at Pennsylvania State. She received her BS in Psychology from the University of
Pittsburgh and her PhD in Human Development and Family Studies from Pennsylvania State
University and was previously on the faculty of the Center for Family Research, in the Depart-
ment of Psychiatry and Behavioral Science at The George Washington University. Neiderhiser’s
work focuses on how genes and environments work together throughout the lifespan and the role
of the prenatal environment. She has been Associate Editor of the Journal of Research on Adolescence,
is on the editorial board of several developmental psychology journals, is currently Associate Edi-
tor for Frontiers in Behavioral and Psychiatric Genetics, and is on the Scientific Advisory Board of the
TwinLife project.
Christine E. Parsons is an Associate Professor at the Interacting Minds Centre, Department of

Clinical Medicine, Aarhus University. She completed her PhD in Psychology at Maynooth Univer-
sity, Ireland and did her postdoctoral training in the Department of Psychiatry, University of Oxford.
She has held positions as college lecturer in Psychology at Harris Manchester College, University of
Oxford, and as Assistant Professor at the Center of Functionally Integrative Neuroscience, Aarhus
University. Her research focuses on understanding the human parental brain using different brain
imaging and behavioral methods.
Rebecca M. Pearson is Lecturer Psychiatric Epidemiology, Based at the Centre for Academic
Mental Health, University of Bristol. Pearson received a first degree in Applied Psychology from
Cardiff University and a PhD in Psychiatric Epidemiology from the University of Bristol. Pearson
is currently leading an EU-funded research program investigating intergenerational transmission of
mental health problems through parenting.
Priya Rajyaguru is an Academic Clinical Fellow in Psychiatry working clinically in psychiatric

medicine and academically at the University of Bristol. Rajyaguru graduated from Cardiff University
with a MBBCh in Medicine and BSc in Psychological Medicine.
Julie Sartori is a PhD candidate with a background in nursing, as a Research Assistant with the
Placenta Project, RAINE Study, and the Australian Fathers Study. She completed her BSc, taught
Medical Science units, and worked in the public and private health sectors. Her research framework
examines maternal, paternal, fetal, and placental factors that affect the growth and development of
the placenta with a focus on morning sickness.
Elizabeth A. Shuey is a Policy Analyst at the Organisation for Economic Co-operation and Devel-
opment where she works on issues related to early childhood, well-being of Indigenous students,
and development of social and emotional skills across international contexts. She was formerly a
xx
Society for Research in Child Development Policy Fellow in the Office of Planning, Research and
Evaluation in the Administration for Children and Families within the U.S. Department of Health
and Human Services as well as a Doris Duke Fellow for the Promotion of Child Well-Being. Shuey
received her PhD from the Eliot-Pearson Department of Child Study and Human Development
at Tufts University and was recognized with the Society for Child and Family Policy and Practice
Dissertation Award.
Eloise A. Stark is currently reading for a DPhil in Psychiatry at Somerville College, University
of Oxford. Her research concerns the neural, attentional, and behavioral mechanisms underlying
parent-infant interaction. She completed her undergraduate degree in Experimental Psychology at
Harris Manchester College, University of Oxford. Stark is a member of the British Psychological
Society and has written for The Conversation and The Psychologist magazine.
Peter N. Stearns is Professor of History and Provost Emeritus at George Mason University. Stearns
has written widely on world and emotions history. His books include A History of Shame, The Indus-
trial Turn in World History, Guiding the American University: Challenges and Choices, Doing Emotions
History, Gender in World History, Satisfaction Not Guaranteed: Dilemmas of Progress in Modern Society,
Childhood in World History, and American Fear: The Causes and Consequences of High Anxiety. He
also edited the Encyclopedia of World History. Stearns taught at the University of Chicago, Rutgers
University, and Carnegie Mellon University. He served as vice president of the American Historical
Association, Teaching Division, and he founded and edited the Journal of Social History.
Alan Stein is Professor of Child and Adolescent Psychiatry at the University of Oxford. He also
holds an Honorary Professorship in the School of Public Health, University of the Witwatersrand,
South Africa, and is an Honorary Fellow of the Child, Youth, Family and Social Development
Programme of the HSRC, South Africa. He received his medical training at the University of the
Witwatersrand, Johannesburg, South Africa. He was previously affiliated with the University of
Cambridge, the Royal Free and University College London Medical School, and the Tavistock
Centre. His main area of research concerns the development of young children in the face of adver-
sity; the ultimate aim of this work is to develop interventions to enhance children’s early develop-
ment and support their families. He is an authority on early child development, especially perinatal
mental health and its effect on parenting and later child development and health. He jointly led the
Lancet perinatal mental health series.
Sandra Tang is a Research Investigator at the Institute for Social Research at the University of
Michigan. She earned a PhD in Applied Developmental and Educational Psychology from the
Lynch School of Education at Boston College. Her research program focuses on the role of the
family in shaping children’s educational success, especially for those children who grow up in risky
contexts. She is co-editor of Socializing Children through Language.
Nicholas E. Waters is a PhD candidate in Developmental Psychology at the University of Michi-

gan and an NICHD Fellow in Interdisciplinary Science. He received his BA from the University
of Michigan in Psychology. His research focuses on children’s cognitive and academic skill develop-
ments and the neurobiological and contextual factors that contribute to different developmental
trajectories.
Ruiyong Wu is an associate professor at the Yangzhou University, China. He received his MS and
PhD degrees from College of Life Science at Shaanxi Normal University. His research focuses on the
behavioral and neurochemical mechanisms underlying maternal behavior.
xxi
Wai Ying Vivien Yiu received her BA from the University of California, Los Angeles, and is
currently a PhD student at the Graduate School of Education, the University of Pennsylvania. Her
research interests include the influence of culture, social change, parenting, and children’s socioemo-
tional development.
Katherine S. Young is a Postdoctoral Researcher at the Anxiety and Depression Research Center,
Department of Psychology, University of California, Los Angeles. She completed a DPhil in Psy-
chiatry at the University of Oxford investigating the neurobiological basis of adults’ responsiveness
to infant vocalizations and how this might be disrupted in depression. Her current research focuses
on investigating responsiveness to social cues among young adults at risk for anxiety and depression
as well as studying the effects of treatment for social anxiety disorder on activity and functional con-
nectivity in the brain.
xxii
PART I
Biology of Parenting
1
THE EVOLUTION OF PARENTING
AND EVOLUTIONARY
APPROACHES TO
CHILDREARING
Introduction
Child and developmental psychologists, sociologists, educators, and policy makers have long viewed
parenting and the family as the most significant influence on the developing child. As such, parent-
ing has traditionally been viewed as an important source of “environmental” variability in the long
debated (and still controversial) nature/nurture dichotomy. At one level, of course, parenting is a
potent part of a child’s environment. An infant’s very survival depends on parents. There is nothing
in the external world so critical to a child’s success in life as her or his parents. Yet, parenting also
straddles the nature side of the traditional continuum. Parenting is not only important to humans,
but it is central to the survival of many species of animals, including all mammals and many birds
(Rosenblatt, 2002). Evolutionary biologists have long recognized this fact, arguing that, in order for
individuals to get their genes into the next generation, they must make investments in mating and,
following conception, parenting (Hamilton, 1964; Trivers, 1972). How much is invested in mating
versus parenting will vary among species and between females and males within a species, depending
on characteristics of the developing offspring and ecological conditions. But parenting—the care and
nurturing of offspring between conception and independence—is universal among mammals and,
depending on the species-typical pattern of such investment, influences how offspring are reared.
Homo sapiens, however, have taken parenting to new heights, not simply because of our use of
language, advanced cognition, cultural transmission of knowledge, or societal institutions, but also
because of the extended period of immaturity of our young. As all mammals, human children are
conceived within their mothers’ bodies, fed after birth with mother-produced milk, and eventually
mature to be able to fend for themselves. But the period of immaturity and dependency is extended
in humans relative to other primates. This prolonged period of youth is seemingly necessitated by the
intellectual demands of human society; children cannot learn enough in a decade of life to function
effectively in any human group. This intellectual immaturity is accompanied by physical immaturity
of offspring that puts extraordinary demands on human parents, surpassing those of any other land
mammal. Such pressures have shaped how parents around the world treat children, the structure of
the human family, and relationships between women and men.
In this chapter, we provide an evolutionary view of human parenting. In a first section, we review
briefly the basic tenets of evolution by natural selection and some of the major ideas of the field
of evolutionary psychology, particularly evolutionary developmental psychology. We then review a
more specific evolutionary theory, Trivers’s (1972) parental investment theory, which accounts for the
3
amount of investment females and males put into parenting (all actions related to rearing an offspring
to reproductive age) versus mating (including the seeking, attaining, and maintaining of a mate). We
next examine some of the selection pressures that produced our species, and how those pressures led
to patterns of parenting and the structure of the family that characterize our species today. We then
take a closer look at some of the factors influencing the decisions parents and other people make for
investing in children. The final major section of the chapter examines some of the evolved psycho-
logical mechanisms—genetically coded “messages” that, following epigenetic rules, interact with the
environment over time to produce behavior—associated with infant-mother attachment, evolved
mechanisms underlying neglect, abuse, and infanticide, and how an evolutionary perspective ques-
tions the very concept of “parenting” as conventionally defined. In all, we argue that an evolutionary
perspective not only tells us where patterns of childrearing came from, but where they may head in
the future as ecological conditions change, and how many problems of contemporary parenting can
be understood, and perhaps solved.
Principles of Evolution and Evolutionary Developmental Science
Evolution by Natural Selection

The basic ideas behind Charles Darwin’s (1859) great theory of “descent with modification” are
surprisingly simple yet frequently misunderstood, particularly when applied to human behavior. The
core of evolutionary theory is the concept of natural selection, which, simply stated, refers to individu-
als who are well suited to their environment leaving more progeny than less well-suited (or less fit)
individuals. Natural selection works because there is variation among members of a generation; that is,
there are different combinations of physical and behavioral traits among individuals within a species.
Critically, these traits, as well as individual differences in these traits, are heritable. Characteristics that
result in an individual surviving and reproducing are passed down from one generation to the next,
whereas characteristics that are associated with early death or low levels of reproduction decrease
in frequency in the population. Characteristics of the individual interact with features of the local
ecology, and it is this interaction that is responsible for increases and decreases in characteristics over
time. This is the process of selection, and through this process, adaptive changes in individuals, and
eventually species, are brought about.
Natural selection is a highly interactive process, involving an active organism’s response to a some-
times changing environment. Evolutionary theorists often use phrases such as “the trait was selected
by the environment” as a shorthand to refer to this complex interaction among an organism, herita-
ble traits of that organism, and the environment. However, the term “selection” does not imply some
deliberate or foresighted process (e.g., selecting for more “advanced” individuals). Natural selection,
and thus evolution, is blind to the future; individuals who fit well with a current environment survive,
and those who fit less well die. Nevertheless, the process, although blind, is an active one, reflecting
the bidirectional relation between an organism with heritable traits and the environment.
Darwin used the term reproductive fitness to refer to the likelihood that an individual will become
a parent and a grandparent. Contemporary evolutionary theorists, taking advantage of the scien-
tific advances in genetics that have occurred since Darwin’s time, use the concept of inclusive fitness
(Hamilton, 1964) to take into consideration the influence that an individual may have in getting
additional copies of her or his genes into subsequent generations. For example, a child possesses 50%
of a parent’s genes. Thus, it is in the parent’s best genetic interest to see that an offspring survives so
that copies of the parent’s genes are passed on to grandchildren (each of whom will possess 25% of
a grandparent’s genes). A person can further benefit the transmission of her or his genes by helping
relatives, who share a smaller percentage of genes. For example, by helping to rear a sister’s four chil-
dren, each of whom shares, on average, 25% of her genes, a woman can further increase her genetic
4
The Evolution of Parenting
contribution to the next generation, thereby increasing her inclusive fitness. Of course, none of this
happens intentionally or consciously. After all, people do not walk around calculating exactly how
related they are to one another before deciding to act altruistically. Rather, the underlying mecha-
nisms are in terms of unconscious evolved “strategies.” Moreover, such patterns are observed in non-
human mammals, birds, and social insects, indicating that self-awareness is not ordinarily involved.
Principles of Evolutionary Developmental Science

Although the principles of evolution should be the same for physical, behavioral, or cognitive char-
acteristics, psychologists investigating the evolution of behavior or cognition, particularly human
behavior or cognition, have made explicit some of these principles. Moreover, developmentalists have
added to or modified slightly some of these principles to achieve a better understanding of the role
of evolution in contemporary human behavior (e.g., Bjorklund and Ellis, 2014; Bjorklund, Hernán-
dez Blasi, and Ellis, 2016; Bjorklund and Pellegrini, 2000, 2002; Geary and Bjorklund, 2000), and we
examine briefly some of these principles here.
First, an evolutionary account of behavioral or cognitive characteristics does not imply genetic
determinism. Certainly, evolutionary change implies change in the frequency of genes within a
population; but evolutionary psychologists argue that behavioral change occurs as a result of a trans-
actional relation between an organism and its environment and that the eventual behavioral pheno-
type of an organism is not predetermined by its genes. From this perspective, development involves
the expression of evolved, epigenetic programs, from conception through old age, as described by the
developmental systems approach (e.g., Bjorklund, Ellis, and Rosenberg, 2007; Gottlieb, 2007; Gottlieb,
Wahlsten, and Lickliter, 1998). Development occurs as a result of the bidirectional relations between
all levels of biological and experiential factors, from the genetic through the cultural. “Experience,”
from this perspective, involves not only exogenous events but also self-produced activity, as reflected
by the firing of a nerve cell in response to solely endogenous factors. Functioning at one level
(e.g., the genetic) influences functioning at adjacent levels (e.g., neuronal) with constant feedback
between levels.
Because the experiences of each individual are unique, there should be substantial plasticity in
development. Yet, there is much that is universal about humans (or any species), and this seem-
ing discrepancy is resolved when one recognizes that infants of a species, beginning at conception,
inherit not only a species-typical genome but also a species-typical environment. To the extent
that individuals grow up in environments similar to those of their ancestors, development should
follow a species-typical pattern. From the developmental systems perspective, there are no simple
cases of either genetic or environmental determinism. Infants are not born as blank slates; evolution
has prepared them to “expect” certain types of environments and to process some types of informa-
tion more readily than others (Bjorklund, 2015). Yet, it is the constant and bidirectional interaction
between various levels of organization, which changes over the course of development, that produces
behavior. For example, differences in the quality and quantity of parental investment affect children’s
development and influence their subsequent reproductive and childcare strategies (e.g., Belsky, Stein-
berg, and Draper, 1991; Ellis et al., 2012, see discussion to follow).
Second, there is a need for an extended childhood to learn the complexities of human com-
munities. Homo sapiens spend a disproportionate amount of time as prereproductives. From an evo-
lutionary perspective, the benefits associated with an extended period of immaturity must have
outweighed the costs. We believe that the most important and difficult things children need to learn
are related to the social complexity of human groups (e.g., Alexander, 1989; Bjorklund and Pel-
legrini, 2002; Dunbar, 2010), although the time to master tool use and food acquisition techniques
(e.g., Kaplan and Gangestad, 2005; Kaplan, Hill, Lancaster, and Hurtado, 2000) would also require an
extended juvenile period.
5
Third, many aspects of childhood serve as preparations for adulthood and were selected over the
course of evolution. Many sex differences in social and cognitive abilities are good examples (Geary,
2010). Evolutionary psychologists have often focused on sex differences (Hyde, 2014), proposing
that women and men have different self-interests and thus have evolved different psychologies. This
focus is reflected especially in sex differences with regard to mating, childrearing, and intra-sex
competition. However, these behaviors, dispositions, and cognitions do not appear with the first
blast of pubertal hormones or on hearing the cries of one’s newborn infant, but have developmental
histories, with children adapting their gender-specific behavior to local norms, based on evolved
predispositions. Such sex differences should not be viewed as a form of biological determinism,
destining women and men to narrow and unchanging roles. Rather, girls and boys are biased toward
different environments and experiences via evolved epigenetic rules, and, to the extent that one’s
environment supports those biases, children will develop in a species-typical fashion. Although these
epigenetic rules may be necessary, they are not sufficient to produce a particular developmental pat-
tern (Bjorklund, 2015; Bjorklund et al., 2007). Human behavior is highly flexible, and although some
outcomes are more likely than others, all require environmental support to be realized.
Fourth, different selection pressures operate on organisms at different times in ontogeny. Although
some aspects of infancy and childhood can be seen as preparations for adulthood, other features have
been selected in evolution to serve an adaptive function at that time in development only and not
to prepare the child for later life (Bjorklund, 1997, 2007). For example, some aspects of infancy may
serve to foster the attachment between an infant and mother to increase the chances of survival at
that time in ontogeny, and not only to prepare the child for later adult relationships. Evolution, we
propose, has endowed children (and the juveniles of other species) with many characteristics that
adapt them well to their immediate environments and not solely to prepare them for a future one.
Fifth, simply because some social, behavioral, or cognitive tendency was adaptive for our ancestors,
does not mean that it continues to be adaptive for modern humans. Similarly, just because some ten-
dencies (such as violence among young adult males) are “natural” based on evolutionary examination,
does not mean that they are morally “good,” excusable, or inevitable. For example, humans’ penchant
for sweet and fatty foods can be seen as a formerly adaptive disposition that, in modern environments
with grocery stores and Ben and Jerry’s Rocky Road Ice Cream, produces increased risk of obe-
sity and heart attacks. Similarly, formal schooling represents a situation in which many of children’s
evolved tendencies do not fit well with the demands of modern society. From the perspective of
evolutionary psychology, much of what we teach children in school is “unnatural,” in that teaching
involves tasks never encountered by our ancestors (see papers in Geary and Berch, 2016). Many other
aspects of social and childrearing behavior, perhaps adaptive for small groups of hunters and gatherers
living on the brink of survival, may not be adaptive for modern people living in nation-state societies.
Although not all developmentalists have embraced the evolutionary principles described here
(e.g., Witherington and Lickliter, 2016), evolutionary thinking has increasingly permeated theoriz-
ing in developmental science and is well on its way to becoming a metatheory—a common set of
broad, overarching assumptions and principles—for developmental psychology (Bjorklund, 2016,
2018). Evolutionary theory has been particularly influential for researchers studying cognitive and
social-cognitive development (e.g., Geary, 2005; Tomasello, 2016). However, the greatest impact of
evolutionary theory as it relates to developmental science has likely been to parenting, dating back
to the theorizing of John Bowlby (1969) on attachment and, especially, to Robert Trivers’s (1972)
parental investment theory, and we describe Trivers’s influential theory in the following section.
Parental Investment Theory

Human parents, particularly mothers, devote substantial time, resources, and energy to rearing their
children. Given humans’ extended period of youth, there is likely no other species that devotes as
6
much time and energy to their offspring from conception to adulthood as Homo sapiens. To try to
make sense of exactly why parents are so involved in a child’s life, it is important to understand the
evolutionary reasons why selection would act to produce parents who invest so much in their chil-
dren. First, children are a parent’s most direct route to genetic immortality. Although a person can
serve her or his inclusive fitness by helping rear nieces, nephews, and younger siblings, reproductive
fitness is most directly served by having children who grow up to become reproductive members of
the community. From this perspective, evolution should operate to select parents who provide the
means by which their offspring attain maturity, and later, carry on the parents’ reproductive lineage.
Parents should provide not only the physical means necessary for their children’s survival (e.g., food,
shelter), but also the means by which children develop competencies in the social groups in which
humans live.
At first glance it may seem as if both females and males should be equally likely to invest in their
children, but this is not the case. For most species, including humans, females invest more heavily in
their offspring than males. This observation and the theory developed around it known as parental
investment theory, was first postulated by Trivers (1972). Trivers based his ideas on Darwin’s (1871)
theory of sexual selection. Sexual selection refers to members of one sex displaying preferences for
certain characteristics of individuals of the other sex, for example peahens showing a preference for
the elaborate tails of peacocks. Darwin believed that sexual selection would occur for two reasons:
First, there would be competition within one sex for access to the other and, second, there would
be differential choice of mate selection by members of one sex for members of the other. Generally,
sexual selection takes the form of males competing with one another for access to females, whereas
females choose among males, often based upon signs of a male’s genetic fitness, successful domination
over competing males, and the likelihood of his providing resources to her and her offspring.
But why is this pattern, with males competing and females choosing, found so clearly both cross-
culturally and across species? The answer, Trivers theorized, lies in the amount of parental investment
each parent typically contributes to an offspring. Differential amounts of parental investment actually
occur before the child is even conceived. For mammals, this is due to the fact that females produce a
finite number of eggs that are large and immobile. Males, in contrast, produce an unlimited number
of small, mobile sperm throughout their lifetimes. This difference in sex-cell size causes females’ eggs
to be more costly metabolically, and thus a limited resource, relative to sperm. Furthermore, the fact
that her eggs are immobile means that conception will happen inside the female’s body, and she will
then carry the child through the gestational period, and usually, be primarily responsible for lactation
and care of the infant after birth. Male investment can theoretically end following copulation. As
such, males have higher potential reproductive rates, in that, following insemination of a female, they
can seek additional mating opportunities; in contrast, once conception has occurred, females’ mating
opportunities end (at least temporarily) and their parenting efforts begin. The end result is that male
mammals typically invest more in mating than parenting, whereas the reverse pattern is found for
females. This greater initial investment by females, Trivers argued, is what sets in motion two differ-
ing strategies as to how to go about finding and maintaining a mate and rearing subsequent offspring.
Depending on the requirements of the young, there are substantial species differences in the
amount of post-copulatory investment males provide to their offspring. Males of some species con-
tribute literally no support to their progeny or the mother, whereas others may spend consider-
able time and energy garnering resources for their offspring, and even spend time in “childcare.”
However, in greater than 95% of mammals, males provide little or no postnatal investment to their
offspring (Clutton-Brock, 1991).
Human males are an exception to the typical mammalian pattern. But despite the well-known
role of fathers as providers (“bringing home the bacon”), and to a lesser extent as caregivers, women
in all cultures provide more support and engage their children more frequently than men (Barnard
and Solchany, 2002; Parke, 2002). This pattern is observed in traditional cultures (Lancy, 2015), in
7
industrialized societies (Whiting and Whiting, 1975), and persists in modern societies in which
women work outside the home (Hetherington, Henderson, and Reiss, 1999). There were changes
in Western cultures over the 20th century, with many fathers spending significant time with their
children, sometimes approaching the time investment made by mothers. However, in these same
societies, the number of children living in homes headed by females increased fourfold since 1960
(see Cabrera, Tamis-LeMonda, Bradley, Hofferth, and Lamb, 2000). Thus, social forces in today’s
world influence degree of paternal investment, but the overall pattern is still women devoting more
of their time in childcare than men, even in the most enlightened families.
The consequences of such differential investment in offspring have important implications for
sex differences in behavior. For women, sexual intercourse brings with it possible conception and
pregnancy. Until recently, infants required breast milk to survive, and this could be provided only by
the mother or other lactating females. Fathers in generations past could not take the 3 a.m. feeding;
the responsibility for feeding infants fell solely to mothers, making postnatal parental investment for
women obligatory (as it is for females of other mammalian species). Men’s minimum required invest-
ment is today, and surely was in the ancient past, substantially less.
As a direct result of this differential minimum investment between women and men, women
tend to be more cautious in assenting to sex than men (Oliver and Hyde, 1993). Women must not
only evaluate the physical qualities of a potential mate (is he healthy, strong, fertile, and so forth), but
they also must evaluate his access to resources (is he wealthy, of high status, or otherwise capable of
supporting a family) and the likelihood of his sharing them with her and her offspring. In contrast,
men are less concerned with the resources of a future mate or her likelihood of sharing. His greater
interest lies with her genetic fitness (is she healthy?) and her ability to conceive, give birth, and care
for a child. These are not necessarily conscious concerns of either sex, for they are reflected in the
behavior of nonhuman animals as well (Clutton-Brock, 1991; Trivers, 1972, 1985).
Members of the less investing sex compete with one another for access to the more investing
sex. In many mammals, the result of such competition is a physically larger male. Increased size and
strength afford males a competitive edge with other males and are associated, in many species, with
higher social status and greater access to females (Geary, 2000, 2010). (High-status or otherwise suc-
cessful males do not simply “take” females as mates; rather, by being successful in competition with
other males, they possess traits that females, over evolutionary time, have come to prefer.) Females,
of course, also compete with one another over males (Smuts, 1995), but female-female competition
is rarely as physically fierce as that between males and is much less apt to result in injury or death
(Campbell, 1999, 2013). Moreover, most females will eventually find a mate, even if an undesirable
one; in contrast, some males will be have no access to females, “shut out” of the Darwinian game
altogether.
Finally, whereas maternity is always certain, paternity never is. It is within the women’s body that
the child is conceived and carried to term, making maternity a sure thing. Males, in contrast, have
no such assurance. A man could spend time, energy, and other resources investing in another man’s
biological child, which would not be adaptive from an evolutionary (reproductive fitness) perspec-
tive. As a result, men are apt to question the paternity of their children and may be less likely to invest
in a child when that child’s paternity is in question. In general, males may be more likely to invest
minimally in their offspring because they know that females will continue to invest in their child,
even if the male invests little, or even deserts her completely.
In sum, evolutionary theory predicts that mothers will be more likely than fathers to invest heav-
ily in their offspring. This phenomenon is seen both cross-culturally and in many species of mam-
mals (as well as most sexually reproducing species). When fathers do invest, they are most likely to
do so when they are sure that the child is genetically their own, and they are sure that the child is
healthy enough to reach reproductive age. This pattern is prevalent in humans today, but it is widely
assumed that it is an old one that has evolved in our species over the past 5 million years.
8
The Environment of Evolutionary Adaptedness

Depending on how one defines modern humans, animals identifiable as Homo sapiens appear in the
fossil record as long ago as 300,000 years before present in the form of Archaic Homo sapiens, or as late
as 35,000 years ago, when the first unambiguous evidence of artistic expression is seen. Anatomically
modern humans are found in fossil records dating back about 100,000 years (see Tattersall, 2013).
But human-like creatures, termed collectively hominins to refer to bipedal (upward walking) apes
including humans and our ancient ancestors, date back 5 to 7 million years ago (mya). Hominins
include members of the Homo genus, but also members of the Australopithecus and Ardipithecus genera.
Although determining with certainty the species to which any fossil belongs is difficult, it is none-
theless certain that many physically and presumably behaviorally different species of hominins have
existed over the past 7 million years, with several different species of hominins living at the same time.
Homo sapiens are the only living member of this group, all others becoming extinct.
Homo sapiens is a relatively young species that has not changed much over the past 100,000 years,
at least physically, and certainly little at all over the past 35,000 years or so. But humans’ physical
conservatism belies a behavioral and cognitive flexibility that has resulted in a radical change in how
we live as a species. The advent of agriculture and a sedentary lifestyle beginning about 12,000 years
ago changed drastically how most human beings lived. For most of the history of Homo sapiens and
its immediate forbears, individuals lived in small, nomadic groups, living off the land, gathering
fruits and vegetables (mostly the work of women), scavenging from the kills made by other animals,
and hunting (mostly the work of men). In one form or another, it was in such hunter/gathering/
scavenging environments in which the modern human mind evolved. Although life has changed
substantially for most members of our species since the advent of agriculture and sedentary lifestyles,
there has not been sufficient time for our brains, and the evolved psychological mechanisms within
them, to evolve. Basically, modern humans possess brains and minds adapted for life in a very different
environment than they find themselves living today. This ancient environment is often referred to
as the environment of evolutionary adaptedness. What was this environment like, how did our ancestors
behave, and what pressures were there that resulted in the modern human mind, and, importantly for
this chapter, how did these pressures lead to the human way of rearing children?
It is impossible to specify exactly what the environment of evolutionary adaptedness was like, in
part because it is impossible to define precisely what time period this term represents. On the one
hand, humans share an evolutionary history with all extant primates and mammals. Thus, histori-
cal environments in which these ancestral mammals and, later primates, evolved are also relevant to
modern humans. If we take as our starting point, however, the period in which the genetic line that
would eventually lead to Homo sapiens separated from the line that would lead to modern chimpan-
zees, we find a period of about 5 to 7 million years, beginning in the forests and savannas of Africa.
Because of the dearth of fossil and archeological evidence for periods much before 2 mya, it is dif-
ficult to say anything with confidence about the lifestyles of the various species of australopithecines.
However, based on what fossil evidence we do have and on the way in which chimpanzees live today,
it is highly likely that hominins were always social species. Based again on limited fossil evidence, the
organization of chimpanzee and bonobo troops, and the lifestyles of contemporary hunter-gatherers,
it is likely that the size of most social groups during the Pleistocene was relatively small (probably
between 30 and 60 people), consisting of both closely related and unrelated individuals, who inter-
acted on a regular basis.
As in all societies today and for the vast majority of mammals, mothers were the primary caregiv-
ers to their children. Fathers likely provided protection to their mates and offspring and support in
the form of food and other tangible resources (Kaplan et al., 2000), but likely spent relatively little
time in direct childcare. Some males surely had several mates, meaning that some females shared the
resources and attention of a single male and that some males had no access to reproductive females.
9
Females probably reached puberty relatively late (late teens, early twenties), and gave birth every 3 to
5 years, with pregnancy often following the cessation of nursing a previous child (Kaplan et al., 2000;
Konner, 2010). Infant mortality was surely high, and, even for those who did survive to adulthood,
life was relatively brief by contemporary standards, with few people living past 40 years (Austad,
1997). However, if contemporary hunter-gatherer societies are any indication, it is likely that there
were always some “old” people (i.e., beyond 60 or 70 years) in every group (Hill and Hurtado, 1991;
Kaplan et al., 2000).
Although hominin groups were usually small, social relationships, especially among large-brained
members of the Homo genus, were surely complex. Humans in all societies around the world coop-
erate and compete with one another and with people from outside groups. Trade among different
social groups is universal to humans, as is warfare. We are aware of no other mammal that engages in
trade, and only the chimpanzee displays anything similar to war parties, attacking and killing mem-
bers of another group of their own species (Goodall, 1986).
All species, as all individuals, have histories, and what happened in the past influences what hap-
pens now and what will happen in the future. Although humans do not have imprinted in their
brains in any simple way the experiences of their ancestors, the environments of ancient Homo sapiens
shaped our ancestors’ social and cognitive evolution, with modern children and adults inheriting
essentially the same brains as our hunter-gatherer forebears, designed by natural selection to solve
recurrent problems of survival. In the next section, we examine some of the selection pressures that
led to the evolution of human intelligence and the human family.
What Were the Selection Pressures that Led to the Modern

Human Mind and the Human Family?
There have been many hypotheses about the “causes” of human evolution. Selection pressures related
to hunting, tool use, navigating large environments, coping with variable environments, diet, and
dealing with conspecifics have all been suggested (among others) as the principal “cause” of human
evolution. There is, of course, no single cause for the evolution of any species, including humans.
Rather, evolution surely proceeded as the result of a confluence of interacting factors, with no single
one being identified as a simple “cause” or “consequence” of another. This does not mean, however,
that some hypotheses of human evolution are not better than others, and the one we prefer, which,
we believe accounts well for humans’ unique cognitive abilities and style of childrearing, focuses
on three interrelated factors: an enlarged brain and the accompanying cognitive abilities, increased
social organization and the need to better cooperate and compete with conspecifics, and an extended
juvenile period (Bjorklund and Bering, 2003; Bjorklund and Pellegrini, 2002). Each of these fac-
tors in concert with the others contributed to changes in what it took for infants to survive and to
grow up to become reproductive members of their group. We describe briefly here the role of social
intelligence and “big brains” in the human evolution story, and focus on how these factors may have
contributed to a prolonged juvenile period, necessitating increased parental investment.
The Significance of Social Intelligence

A number of theorists have proposed that the single most important selection pressure in the evo-
lution of human intelligence was sociality—interacting, competing, and cooperating with other
members of the species (e.g., Alexander, 1989; Bjorklund and Pellegrini, 2002; Byrne and Whiten,
1988; Dunbar, 2010; Hare, 2011; Humphrey, 1976; Tomasello, 2014). As hominin groups became
more complex, a greater social intelligence was required to maneuver the often stormy waters within
small groups of long-lived conspecifics. Individuals who could reflect on their own knowledge,
10
intentions, and desires, and, importantly, the knowledge, intentions, and desires of others (theory of
mind), would have been at an advantage in cooperating and competing with others both within and
outside their immediate group.
As social cohesion became more important in primate and hominin groups, the need to control
sexual and aggressive responses also increased in importance (Bjorklund and Harnishfeger, 1995).
This need to control sexual and aggressive urges may have been particularly true for early humans,
attributed, in part, to changes in females’ receptivity to sex. In many primates, there is considerable
competition between males for access to estrus females. The receptivity of females to sexual advances
varies across species, with female chimpanzees, bonobos, and some monkeys being receptive for a
period of time beyond the period of estrus, resulting in extended competition among males. In con-
trast to the other great apes, human females do not show any (obvious) outward signs of ovulation,
and, unlike other mammals, they present permanently swollen mammaries, whether nursing or not,
which have become constant sexual signals for males, despite their unreliability in predicting sexual
receptivity or ovulation. Thus, sexual receptivity in both human females and males cannot be deter-
mined by physical body signs, such as swollen genitals in apes. Moreover, both females and males are,
in theory, continually receptive sexually, with their willingness to copulate being primarily under the
control of social and not hormonal factors.
These changes in female sexual behavior (potentially receptive even when not ovulating) and
appearances (no outward sign of sexual receptivity or ovulation) may have contributed significantly,
along with other factors, to human pair bonding. For a male hominin, whose investment is required
if his long-dependent offspring is to survive, it is critical that he be confident that his resources are
going to his genetic progeny and not to the offspring of another male. But being certain of paternity
can be problematic in a species in which ovulation, and thus fertility, cannot be known by the male,
and when females, as well as males, are potentially sexually receptive at all times. To counteract this
dilemma, males may resort to some form of mate guarding, in which they hover near their mates
during her fertile time, preventing her access to other males. But males cannot guard their mates all
of the time. And, although it may seem to be to the female’s advantage to have as many options in
terms of potential mates as she can, mating with a large number of males would do her little good
if none of them contributed significantly to the support of her offspring, which seems to have been,
if not necessary at least highly desirable, in hominins (Geary, 2010). One solution to these problems
may have been the “invention” of neurochemical systems (opioids and oxytocin) that fostered strong
emotional bonds between a female and male, producing marginally (and temporary) monogamous
behavior in the pair, long enough so that children can reach an age so that they can care for them-
selves (Fisher, 1992).
As we have suggested, one condition necessary for substantial paternal investment to evolve
would be a high degree of paternity certainty. This seems to have been achieved in contemporary
humans. Studies from a broad range of countries have estimated the degree of paternity discrepancy
(in which the domestic father is not the genetic father) to be between 7% and 15% (see Bellis and
Baker, 1990; Lerner and von Eye, 1992), although more recent studies put the rate at between 1%
and 2% (Larmuseau et al., 2013). Thus, although women clearly engage in extra-mate copulations
(surely enough for men to have evolved mechanisms to guard against cuckoldry), they apparently do
not frequently make cuckolds of their mates. The result is a male who can be relatively confident of
the paternity of his offspring, a female who obtains resources for herself and her offspring from her
mate, and an offspring who survives past infancy.
Inhibitory abilities necessary for increasing the control of sexual and aggressive behaviors would
require increased neural capacity, and they may have been part of the selective pressures that led to
enhanced brain size, particularly of the neocortex, in the hominin line. Alternately, other factors may
have been primarily responsible for the increase in brain size seen in hominins over the past 5 million
11
years, with greater inhibitory abilities being a by-product of this increase, co-opting neural circuits
that had been selected for other purposes. Nonetheless, once inhibitory abilities did increase, the
behaviors they produced were subject to natural selection. And whether they were primarily a cause
or a consequence of increased brain power, what is undeniable is that brain size did increase, and, for
better or worse, Homo sapiens’ large brain and resulting cognitive processes define us, more than any
other feature, as a species.
Large Brains
Humans have disproportionately large brains relative to their body size ( Jerison, 1973, 2002). Brains
are very expensive in terms of the calories they consume, so that having “more brain” than needed
to control the body must have substantial benefits for survival. Homo sapiens’ large cranium did not
materialize out of thin air, however. Primates in general have larger brains than expected by their
body size (represented by the encephalization quotient, or EQ, which reflects brain size relative to
the expected brain size for an animal of a specified body size; Jerison, 1973); humans merely reflect
an extension of a pattern already observed in primates. Figure 1.1 shows the EQ for chimpanzees
and for several hominin species. (An EQ of 1.0 is the “expected” value, with EQs greater than
1.0 reflecting “more brain” than predicted for an animal of a specified size.) As can be seen, the
encephalization quotient for Australopithecus afarensis was only slightly greater than that of modern
chimpanzees. From this point on in evolution, brain weight relative to body weight increased at a
rapid rate. One set of factors responsible for this change was related to the increased social com-
plexity of hominin groups, although changes in diet, technology, and responses to modifications in
climate all likely played interacting and contributory roles. But regardless of the reasons (i.e., selective
pressures) for increased brain size, there must be some mechanisms within the organism for achiev-
ing this change. One important mechanism, we believe, can be found in alterations of patterns of
development, which, in turn, would provide additional changes that must pass through the sieve of
natural selection.
7
Encephaliztion Quotient
1
P. troglodytes A. afarensis H. habilis H. erectus H. sapiens
Figure 1.1 Encephalization quotients for chimpanzees (Pan troglodytes) and four hominid species (data for
chimpanzees from Jerison, 1973; data for hominids from Tobias, 1987). An EQ of 1.0 is the
“expected” value, with EQs greater than 1.0 reflecting “more brain” than predicted for an animal
of a specified size.
12
The Consequences of Delaying Development

Although humans’ brains are bigger than those of their ancestors, somewhat ironically, one mecha-
nism by which brains increased in size was the process of delayed development. Some evolutionary
changes can be brought about by changing patterns of development. Genetic-based differences in
developmental rate have been referred to as heterochrony (de Beer, 1958; Gould, 1977). For simplic-
ity sake, we will talk about only two general forms of heterochrony, acceleration, in which the rate of
development (ontogeny) in an individual is accelerated or extended relative to one’s ancestors, and
retardation, in which development is slowed down or delayed in comparison to ancestral patterns.
In one sense, big brains are a good example of accelerated development. The development of the
brain of Homo sapiens is clearly extended beyond that of its progenitors. Yet, to achieve that exten-
sion required the delaying of a pattern of growth rate typical of the prenatal period to postnatal life.
The primate brain develops rapidly in comparison to the overall size of the body (Bonner, 1988).
For chimpanzees, macaque monkeys, and other primates, brain growth slows quickly after birth; this
is much less so for humans. Rather, the rate of prenatal brain growth for humans continues over the
first 2 years of life (Gould, 1977). By 2 years of age, the human brain has attained 50% of its eventual
adult weight; in contrast, total body weight is only about 20% of what it will eventually be (Tan-
ner, 1978). Increasing the time the brain grows increases the number of neurons that are produced
(Finlay, Darlington, and Nicastro, 2001) and also results in the extension of dendritic and synaptic
growth, so that the human brain has more neurons and more interconnections among neurons than
the brains of other primates (Finlay et al., 2001; Gibson, 1991). Although most parts of the brain have
undergone enlargement in human evolution, the effects are most pronounced on the neocortex, the
so-called “thinking” portion of the brain.
The extension of embryonic growth rates for the brain into the second year of life was neces-
sitated by some physical limitations of human females. Big brains require big skulls, and if a human
newborn’s skull were as large as “expected” given the eventual adult size (and given the standard
primate rate of pre- and postnatal brain development), the infant’s head would be too large to fit
through the birth canal. The size of women’s hips (which determine the size of the birth canal) are
limited by the need for bipedality. A woman with hips large enough to give birth to an infant having
the cranium the size of a contemporary 2-year-old child would not be able to walk. Thus, evolution-
ary pressures that resulted in an enlarged brain required that pregnancy be extended only to the point
where the infant skull would fit through the birth canal. The result is a physically immature infant,
motorically and perceptually far behind the sophistication of other primate infants (see Antinucci,
1989; Gibson, 1991).
Along these lines, Piantadosi and Kidd (2016) proposed the evolving relation between brain size
and parental care was the driver for human evolution. Humans are physically immature at birth
(altricial), being born early enough to accommodate for their large brain size. However, due to this
early birth, newborns are rather helpless and require extensive caregiving to survive. This extensive
caregiving requires more intelligence, which, according to Piantadosi and Kidd, led to larger brains.
Piantadosi and Kidd proposed a model illustrating that remarkably large brains and high intelligence
are due to runaway selection associated with caring for physically immature and helpless infants.
This runaway selection is similar to the runaway dynamics that are seen in sexual selection, where,
for example, female preference may cause selection to run out of control and result in males hav-
ing traits that are not beneficial for survival. If a longer tail helps a bird to fly more quickly, aiding
survival, females will prefer the longer-tailed males. However, this selection may run out of control,
as females choose even longer tails over time, resulting in plumage that is large and detrimental to
avoiding predators, as seen in peacocks. The current model examines birth age, adult head/brain
size, and an individual’s intelligence, all of which are subject to selection pressures. Piantadosi and
Kidd’s results suggest that there were strong effects of infant altriciality on primate intelligence.
13
The researchers suggested that humans’ advanced means of reading social intentions may have
evolved to care for helpless neonates and that these demands may also be associated with the rise of
social systems of cooperative breeding, in which mothers receive assistance from mainly female kin
in rearing offspring (Hrdy, 2009). As a result, human brain and cognitive development soon became
accelerated relative to their primate cousins, due in large part to the retention of fetal brain-growth
rate (Langer, 1998; Parker and McKinney, 1999).
Rate of brain development is not the only aspect of ontogeny that is delayed. As a species, humans
spend a disproportionate amount of time as prereproductives. Worldwide today, the average age
of menarche is between 12.5 and 13.5 years. However, for both girls and boys, there is typically
a period of low fertility, extending the nonreproductive years even further (Bogin, 1999; Tanner,
1978). Based on historical data and data from traditional cultures (Hill and Hurtado, 1996; Kaplan
et al., 2000), it is likely that our ancient ancestors were closer to 18 to 20 years of age before being
fully reproductive. Such delayed reproduction is all the more impressive when one considers that the
likely life expectancy of our hominin forebears was substantially less than ours today, meaning that
many children would die before reaching reproductive age, and that many others would have only a
limited number of reproductive years (Volk and Atkinson, 2013). Many women, for example, surely
died in childbirth. When looked upon in hindsight, our delayed maturation had substantial risks.
Given these risks, the selective pressures for this delayed maturation must have been derived from
strong compensatory advantages of the immature state, most notably increased flexibility of learning.
Human development is different from that of other primates not only in quantitative terms (i.e.,
being slow and extended), but also in qualitative terms. For example, Bogin (1999) proposed five
stages of development for Homo sapiens: infancy, childhood, juvenility, adolescence, and adulthood,
two of which (childhood and adolescence) are not observed in any other species. Infancy ends with
the cessation of nursing and is followed in other mammals by the juvenile period, in which the
young animal is no longer dependent on its parents but is not yet sexually mature. In contrast, wean-
ing in humans occurs between 2 and 5 years of age, but it is another several years before children can
eat an adult diet and otherwise fend for themselves. The juvenile period in humans (often referred to
as middle childhood) is followed by adolescence, with its characteristic growth spurt, and continues
until sexual maturity, typically in the late teen years. No other species displays this rapid growth spurt
before adulthood, although chimpanzees and bonobos also apparently have a post-menarche period
of infertility (Bogin, 1999).
Based on fossil evidence, Bogin (1999) estimated that the life stages of our australopithecine
ancestors were similar to that of chimpanzees (Pan troglodytes), consisting of a period of infancy
lasting 5 or 6 years, followed by a juvenile period with adulthood beginning about 12 years of age.
According to Bogin, it is only with the beginning of the Homo line that a period of childhood is
seen, and only in modern Homo sapiens is there evidence for a period of adolescence. In addition
to the emergence of childhood and adolescence, the length of juvenility and adulthood is longer in
humans than in other primates and is almost certainly longer than for our hominin ancestors. There
is also evidence from fossil dental and cranial development that brain development in Neanderthals
was much faster than in modern humans (e.g., Akazawa, Muhesen, Dodo, Kondo, and Mizouguchi,
1995; Dean, Stringer, and Bromage, 1986; Zollikofer, Ponce de León, Martin, and Stucki, 1995).
Mithen (1996) used this evidence to suggest that the modern human mind, with the ability to com-
municate between different cognitive modules, required an extended juvenile period, and Nielsen
(2012) proposed that the emergence of fantasy play and imitation during childhood were necessary
components for the evolution of human intelligence.
There are many possible reasons for the extension of developmental periods in humans (Bogin,
1999), but the very fact of this developmental extension indicates than ancient members of the Homo
line were able to keep children alive long enough to reach an age at which they could reproduce
themselves. Note also that the extended developmental period is associated with an enlarged brain.
14
We, and others (Bogin, 1999; Dunbar, 2010), believe that the extended period of youth and an
enlarged brain was necessitated to master the increasing complexity of the social environment. In
fact, research showing relations among large brains, an extended immaturity, and social complexity
was reported by Joffe (1997), who compared aspects of brain size with length of the prereproductive
period and aspects of social complexity for 27 primates, including humans. Joffe reported that the
proportion of the lifespan spent as a juvenile was positively correlated with group size and the relative
size of the nonvisual neocortex. This is the part of the primate brain that is associated with complex
problem solving, including memory. Joffe argued that social complexity exerted selection pressures
for increased nonvisual neocortex in primates and an extension of the juvenile period.
Extended childhoods would also be useful for mastering other important skills in addition to
social intelligence. For example, Kaplan et al. (2000) proposed that it was ancient humans’ shift
to a higher-quality diet that necessitated greater cognitive skills and thus an extended childhood
to learn. Chimpanzees, for example, rely primarily on a diet of easily extracted fruit and plants with
low nutrition density. Such foods, when available, can be obtained relatively easily even by juveniles.
Chimpanzees obtain only a small portion of their diet via hunting, which provides foods of high
nutrition density. Hunting, however, is engaged in mainly by adults, and takes considerable time to
learn. Kaplan et al. (2000) examined food-gathering procedures in contemporary hunter-gatherer
societies and noted that, similar to chimpanzees, children often forage for low-density, easily acces-
sible foods, such as ripe fruit, at young ages and become relatively adept at the task. Extracting foods
of higher nutrition density, such as roots and tubers or vertebrate meat through hunting, is performed
effectively only by older individuals and require many years to master.
What does all this have to do with the formation of the human family and patterns of parenting?
First, the enlargement of the Homo brain required that much of brain growth continues postnatally,
due to restrictions of the female anatomy. This extended physical dependency was coupled with an
extended childhood due to the need to learn the complexity of one’s social environment (or pos-
sibly, in addition, to learn the mechanisms for processing high-quality food), which further extended
the time children spend as prereproductives. Humans’ extended period of immaturity necessitated
that adults, particularly mothers, be disposed to care for their long-dependent offspring. Although
natural selection may have increased the probability that parents are disposed to care for their chil-
dren, ancestral parents could not invest indiscriminately in offspring. Rather, parents and other adults
responsible for the care of children, must evaluate how best to devote their time, effort, and material
resources to children, a topic we turn to in the following section.
Investing in Children
If children are to grow to become sexually mature and economically productive members of their
community, they require substantial support from their parents. Parents allocate effort and resources
to their offspring that could otherwise be devoted to mating effort or spent on their own physical
development and acquisition of resources. Yet, allocating resources to an infant not only limits one’s
own ontogeny and mating efforts, but also compromises opportunities to invest in other offspring,
both those born and unborn (Keller, 2000). Although it may seem obvious that parents, particularly
mothers, will do anything to enhance the survival of their children, there are factors, in both contem-
porary and ancient environments, that affect how much mothers are willing to invest. These factors
include the health of a child, the conditions of the local economy/ecology, the presence of additional
children, the age and reproductive status of the parents (particularly the mother), and the amount of
social support available to help rear a child, among other factors (Bornstein, 2015).
We focus here on the issue of social support. Human mothers likely have never reared a child
“alone.” Because of the extended dependency of their offspring, human mothers must spend more
time caring for their offspring than mothers of other mammals, leaving less time for activities that
15
would be important for their own growth and that of their other offspring. This extended depend-
ency of offspring made it necessary for a mother to receive assistance from others, including resources
and some childcare from the father, but also support from members in the community. In this section,
we discuss briefly the two most likely sources of support for a mother and her offspring: fathers and
children’s grandparents.
The Importance of Paternal Investment

The long period of offspring dependency meant that a male’s genetic success could not be measured
just by how many children he sired. His inclusive fitness would depend on how many of his offspring
reached sexual maturity, assuring him of becoming (at least) a grandfather. To increase the odds of
this happening, his help in rearing his children would be needed. Human males devote more time to
“parenting” than the vast majority of mammals (Clutton-Brock, 1991) and, in most contemporary
hunter-gatherer societies, provide the majority of calories consumed by both their offspring and
their mates (Kaplan et al., 2000). Increased paternal investment permitted human females to rear
multiple dependent offspring and to cut the childhood mortality rate in half in comparison to other
primates and group-hunting carnivores (Lancaster and Lancaster, 1987).
The significance to survival and success of paternal investment is not just speculative, but is sup-
ported by evidence from modern societies, contemporary hunter-gatherers, and historical records
(see Geary, 2000, 2010, for reviews). For all types of data sets, children’s mortality rates are higher
and their social status is lower when fathers are absent. Moreover, in contemporary U.S. America,
the quality of a father’s active and supportive involvement in his children’s lives is positively associ-
ated with emotional regulation, academic achievement, and social competence (Cabrera et al., 2000;
Lamb, 1997).
Grandparental Support
Although fathers may be the most important source of support to a mother and her children, in all
societies, support also comes from related kin, most often from the child’s grandparents. The condi-
tions under which grandparents are apt to provide support are similar to the conditions under which
fathers are likely to make investments—when genetic relatedness is high. Maternal grandparents, like
the mothers themselves, can be quite confident that the baby is related to them, whereas paternal
grandparents, as fathers, can never be 100% certain of paternity. As such, evolutionary theory predicts
that, on average, maternal grandparents will invest more in their grandchildren than will paternal
grandparents, and the research literature has consistently confirmed this relation (Smith and Drew,
2002). Studies from a variety of countries have shown that maternal grandparents have more contact
with and show greater solicitude toward their grandchildren than paternal grandparents, even after
controlling for the distance the grandparents live from their grandchildren (Smith and Drew, 2002).
Moreover, maternal grandfathers are viewed as devoting more care to their grandchildren than
paternal grandmothers, despite the greater childcare role that women play in all cultures (e.g., Dan-
ielsbacka and Tanskanen, 2012; Euler and Weitzel, 1996). A similar pattern of investment has been
found for maternal versus paternal aunts and uncles (Gaulin, McBurney, and Brakeman-Wartell,
1997; Pashos and McBurney, 2008).
Consistent with these findings, child survival is increased by the presence of a maternal grand-
mother. For example, based on 150 years of German birth/death records, Beise and Voland (2002)
reported that children with living maternal grandmothers were more likely to survive than chil-
dren without living maternal grandmothers, and, in rural Ethiopia, help provided by a mother’s
mother was associated with lower child mortality (Gibson and Mace, 2005). Other research has
shown that the presence of a maternal grandmother was associated with higher fertility and
16
survival rates for Canadian and Finnish farm families (Lahdenperä, Lummaa, Helle, Tremblay, and
Russell, 2004).
Grandparents who contribute to the success of their adult offspring and their grandoffspring can
serve to decrease infant morbidity and mortality rates, which increases their inclusive fitness. Some
have even speculated that such grandparental investment has contributed significantly to Homo sapi-
ens’ longevity (e.g., Hawkes, O’Connell, and Blurton Jones, 1997; O’Connell, Hawkes, and Blurton
Jones, 1999). Characteristics associated with longevity can be selected for if older people continue
to reproduce. That is, long-lived individuals can pass these characteristics directly to their offspring.
However, most human females in traditional societies have their last children 20 or 30 years before
they die. Thus, both women who live long lives and those who live shorter lives are likely to have
reproduced before natural selection will have had an effect on genes associated with longevity. But
genes for longevity can be selected for if long-lived (but nonreproducing) individuals foster their
grandchildren’s survival.
There is evidence from a variety of species for the “grandmother hypothesis” (because it is pri-
marily grandmothers and not grandfathers who provide support to their grandchildren). Vervet
monkeys, baboons, lions, and humans in traditional societies all benefit from the presence of a grand-
mother (Hawkes, 2004; Hawkes et al., 1997; Packer, Tatar, and Collins, 1998). For example, research
with the Hadza, a small group of foragers living in Africa’s Rift Valley, found that older women’s
foraging was particularly important for the nutrition of young children who had been weaned but
who were not yet prepared to eat adult food (O’Connell et al., 1999). Hawkes et al. (1997) reported
that, in families in which mothers were nursing, the nutritional status of weaned children was related
to the foraging efforts of their grandmothers rather than their mothers. If this pattern reflects ancestral
populations, the result would have been to increase fertility by permitting mothers to wean a child
earlier and become pregnant again sooner. Without grandmother support for weaned children, nurs-
ing would likely continue for several more years, reducing the total number of children a female
could expect to have.
Although most mammal mothers have sole responsibility for the care of their infants, this is
not the case for humans. Unlike the vast majority of male mammals, human fathers in all societies
provide some childcare and/or resources to their children, with additional care being provided by a
host of alloparents, people other than the genetic parents who help care for a child (Hrdy, 2009). In
many cultures, maternal grandmothers are the most frequent alloparents, contributing to the health,
survival, and perhaps even longevity of their grandoffspring. Yet, natural selection not only provides
clues to parents and others about whom to invest in (e.g., genetically related children), but also pro-
vides psychological mechanisms to increase the chances that investment will indeed be made, a topic
we turn to in the next section.
Evolutionary Perspectives on Attachment and Parenting

Evolutionary theory can provide some insights into how patterns of human parenting came to be,
but does an evolutionary approach to parenting provide anything more than an interesting historical
perspective? We argue that it does, that looking at parenting through the lens of evolutionary theory
can be useful for understanding important aspects of childrearing relevant to people in contemporary
societies (Bjorklund and Ellis, 2014; Bjorklund and Pellegrini, 2002; Keller, 2000). Perhaps the most
empirical research related to parenting from an evolutionary perspective concerns attachment and
the consequences that styles of attachment have for subsequent development. This is due, in part,
to the fact that Bowlby (1969), the founder of modern attachment theory, saw attachment from
an ethological (as well as psychoanalytic) perspective, believing that attachment served an adaptive
function to infants in the environment of evolutionary adaptedness. In the following sections, after a
brief discussion of evolutionary developmental models of attachment, we look at how evolutionary
17
explanations can be helpful in understanding the possible psychological mechanisms in play when
parenting “goes wrong,” resulting in the neglect, abuse, or even the death of children at the hands of
their parents.
Evolutionary Adaptations that Promote Attachment

It is one thing to say that adults are biased toward caring for their helpless offspring, but it is quite
another to propose mechanisms for such behavior. We have argued previously that infants (and their
parents) have evolved low-level perceptual biases that, in interaction with species-typical experiences,
result in species-typical behavior (Bjorklund, 2015; Bjorklund and Ellis, 2014; Bjorklund et al., 2007).
For example, human infants are biased to attend to displays of biological motion (e.g., Bardi, Regolin,
and Simion, 2011, 2014) and to face-like stimuli (e.g., Easterbrook, Kisilevsky, Hains, and Muir, 1999;
Mondloch et al., 1999), and to match certain facial gestures of adult models, such as tongue protru-
sion (neonatal imitation; Meltzoff and Moore, 1977; but see Oostenbroek et al., 2016), each of which
serves to foster social relationships (i.e., attachment) with their caregivers.
Adults also possess evolved mechanisms, increasing the likelihood that they will form attachments
to their helpless wards. In fact, Bowlby (1969) proposed that infants evolved certain signals and abili-
ties to promote social relationships with their parents (see also Goetz, Keltner, and Simon-Thomas,
2010). Such cues can be physical (e.g., baby-schema features described by Lorenz, 1943, including a
rounded and large head relative to body size, fat cheeks, a flat nose, big eyes located below the mid-
dle of the face profile, and short and broad extremities), behavioral (e.g., smiling, clumsy movements;
Bowlby, 1969), or vocal (e.g., infants’ crying as cues to health; DeVries, 1984; Kringelbach, Stark,
Alexander, Bornstein, and Stein, 2016; Soltis, 2004).
In recent years, research has focused on the effects that baby-like faces, as described by Lorenz’s
baby-schema, have on adults’ attitudes and behavior toward infants. Research has consistently reported
that adults rate baby-faced infants as cuter, more attractive, friendlier, and more helpless and powerless
than infants and older children with more mature faces (e.g., Alley, 1981; Leibenluft, Gobbini, Har-
rison, and Haxby, 2004; Senese et al., 2013); such immature physical features are also associated with
enhanced motivation for caregiving, and even a higher likelihood of making a favorable adoption
decision (e.g., Glocker et al., 2009; Waller, Volk, and Quinsey, 2004). Adults continue to view young
faces positively until about 4.5 years of age, after which adults’ ratings of likeability and attractive-
ness of infant faces are similar to ratings of adult faces (Luo, Li, and Lee, 2011). Although faces may
no longer serve as powerful cues promoting caregiving for young children, adults respond positively
to some aspects of preschool-age children’s verbal expressions of cognitive immaturity (e.g., “The
sun’s not out because it’s mad”), with children expressing such statements being rated more highly
on items reflecting positive affect (e.g., likeable, cute) and helplessness, and lower on items reflecting
negative affect (e.g., sneaky, feel more irritated with) than children expressing more mature cognition
(e.g., “The sun’s not out because a cloud moved in front of it”; Bjorklund, Hernández Blasi, and Per-
iss, 2010; Hernández Blasi, Bjorklund, and Ruiz Soler, 2015). Some research has shown that this posi-
tive bias for immature faces (Fullard and Reiling, 1976; Gross, 1997; but see Borgi, Cogliati-Dezza,
Brelsford, Meints, and Cirulli, 2014) and immature cognition (Hernández Blasi and Bjorklund, 2018;
Periss, Hernández Blasi, and Bjorklund, 2012) is not observed until later adolescence, beginning
about 15 years of age, suggesting that the bias may be related to the possible onset of parenthood.
These biases of infants to attend to social cues and of adults to be positively disposed to immature
features of infants and children serve to promote the survival of helpless, dependent offspring, pri-
marily through the development of attachment. Infant-mother attachment is common throughout
the animal world, particularly in mammals and birds. According to Del Giudice (2009), attachment
is a biologically based motivational system that evolved to protect children from danger while moti-
vating caregivers to provide care. Bowlby (1969) believed that, although all but the most deprived
18
of infants become attached to their mothers or mother figures, there were measurable differences
in the quality of attachment, with some forms of attachment (notably secure) being associated with
better psychological outcomes than others (notably insecure). Moreover, it was behaviors of mothers
that served to establish and maintain style of attachment. Research by Ainsworth and her colleagues
(e.g., Ainsworth, Blehar, Waters, and Wall, 1978; Ainsworth and Wittig, 1969) over the past 50 years
has generally supported Bowlby’s contention. For example, securely attached infants are likely to
have mothers (or other caregivers) who respond to them contingently and who are responsive to
their signals of physical and social need (e.g., Harrist and Waugh, 2002; Isabella and Belsky, 1991);
and longitudinal research has demonstrated that children and adolescents who were classified as
securely attached as infants and toddlers display better social and cognitive functioning than those
who had been classified as insecurely attached (Thompson, 2006). These relatively robust patterns
led many to the conclusion, consistent with Bowlby’s original proclamation, that secure attachment
represents the most adaptive style, with aspects of insecure attachment being predictive of poor
adjustment and psychopathology (see Thompson, 2006).
Other theorists, however, speculated that attachment systems are flexible and evolved to adapt
individuals to subsequent environments (e.g., Belsky et al., 1991). From this perspective, different
patterns of attachment should develop as a function of the ecological conditions of a child’s local
environment (including amount of parental investment). Moreover, attachment classifications should
reflect adjustments to contemporary environments and should not necessarily be stable over time
when ecological conditions vary (Lewis, Feiring, and Rosenthal, 2000).
The evolutionary-based theory that has generated the most research and controversy in this area
is that of Belsky et al. (1991). Belsky et al. (1991) formulated psychosocial acceleration theory that links
childhood experience, psychological development, somatic development, and reproductive strategies.
They proposed that aspects of children’s environments affect their attachment style and also impor-
tant aspects of later reproductive strategies. According to Belsky et al. (1991, p. 650),
a principal evolutionary function of early experience—the first 5 to 7 years—is to induce

in the child an understanding of the availability and predictability of resources (broadly
defined) in the environment, of the trustworthiness of others, and of the enduringness of
close interpersonal relationships, all of which will affect how the developing person appor-
tions reproductive effort.
Rather than viewing secure attachment as being the “best strategy” for a child to follow, they pro-
posed that humans have evolved mechanisms that are sensitive to features of the early childhood
environment that induce rate of pubertal maturation (especially in girls) and influence reproductive
strategies. Specifically, they suggested that children from homes characterized by high stress, insecure
attachment, and father absence, attain physical maturity early, are sexually promiscuous, and form
unstable pair bonds. This pattern contrasts to children from low-stress, secure attachment, and father-
present homes, who reach puberty later, delay sexual activity, and form more stable pair bonds. The
former strategy may be adaptive for children growing up in harsh and unpredictable environments
with little expectation of social support. In such cases, both females and males invest relatively more
in mating than parenting, taking a “quantity over quality” perspective. In the latter case, in which
children receive social support in a low-stress, adequately resourced environment, they invest rela-
tively more in parenting than mating, taking a “quality over quantity” perspective. In other words,
Belsky and his colleagues proposed that children follow alternate reproductive strategies, depending
on the availability of resources in their rearing environment, which results in differential investment
in the next generation.
Although it is beyond the scope of this chapter to review the literature that has accumulated on
this issue, the hypothesis has generally been supported by the research literature (see Belsky et al.,
19
2007; Ellis, 2004; Ellis, Figueredo, Brumbach, and Schlomer, 2009). Children from high-stress, father-
absent homes who experience harsh and unpredictable early environments participate in more risky
and aggressive behavior, have earlier pubertal maturation (especially females) and sexual debut, form
unstable pair bonds, become pregnant earlier, and will provide less investment to their offspring than
children who experience less harsh and more predictable environments (e.g., Ellis, 2004; Ellis et al.,
2009; Nettle, 2010; Nettle and Cockerill, 2010; Placek and Quinlan, 2012). Similar effects have been
observed in some nonhuman mammals. For example, male rats reared in heightened environmental
stress conditions showed more adolescent play-fighting behavior (Parent and Meaney, 2008) and
adult aggressive behavior (Menard and Hakvoort, 2007), whereas female rats shifted to an earlier
reproductive strategy (Meaney, 2007). This sex difference of enhanced effects for females makes sense,
given the differential investment in offspring by females and males. Because females’ investment in
any conception is greater than that of males, they should be more sensitive to environmental factors
that may affect the rearing of offspring (such as malnutrition, stress, lack of resources) than males.
Evolved Mechanisms Underlying Neglect, Abuse, and Infanticide

It seems a given that all parents want the best for their children—that children are parents’ route to
immortality. From a Darwinian perspective, reproduction is the sine qua non of success, making situ-
ations in which parents do not act in the best interests of their children paradoxical. These situations
become a bit easier to understand, however, when one considers that any given child is only one of
potentially many offspring, some of whom may be better candidates for continuing a parent’s genetic
heritage than others.
Differential parental investment. Parents often choose to invest differentially in their offspring, invest-
ing the most in those who have the greatest chance of reaching reproductive age and thus carrying
forth the parents’ genes. Parents must balance costs associated with care of a specific child against
resources that can be used for other children, both those born and yet born, and for the parents
themselves. Differential investment in offspring is most apparent in the behaviors of mothers. In ages
past, it seems likely that mothers who were skilled at identifying cues to a child’s future reproductive
success could invest more time, energy, and resources in those children, influencing substantially the
likely survival of her various offspring. Mothers who were less proficient at making these discrimi-
nations or less reluctant to act on perceived differences, were likely to squander scarce resources on
a child who may not make it to adulthood, no matter the degree of investment made. From this
perspective, evolution has selected mothers who are skillful at identifying which children, as well
as which circumstances, are best suited to rearing a child to reproductive years (Hrdy, 1999, 2009).
Reduced maternal investment can take many forms. Children may be neglected, receiving less
attention, medical care, and food than they might need; they may be abused, wet-nursed, fostered out
with relatives or even strangers, or left in the custody of a religious institution. Infants and children in
some cultures have been sold into slavery, or at the extreme, put to death (Hrdy, 1999).
Under what conditions would parents, particularly mothers, decide to reduce investment in chil-
dren? One set of salient cues comes directly from infants themselves. Sickly babies may be a bad
investment, particularly if caring for a sickly child means devoting fewer resources to healthier chil-
dren or postponing becoming pregnant again with the chance of having a healthy baby who is more
likely to survive and thrive. Although our society places a high value on the life of even the most
sickly infants, this is not universal. For example, anthropological data indicate that the killing of a
deformed or seriously ill infant was sanctioned in about one-third of the traditional cultures studied
(Daly and Wilson, 1984). In our own society, children with intellectual impairment or those who
have other congenital defects, such as Down syndrome, spina bifida, cystic fibrosis, or cleft palate, are
abused at rates 2 to 10 times higher than unaffected children (see Daly and Wilson, 1981 for review);
20
and when these children are institutionalized, parental interest rapidly decreases, and many are not
visited ever (Daly and Wilson, 1988).
Differential investment in sickly infants is often less severe, as exemplified by Mann (1992), who
examined the interaction between mothers and their premature and extremely low-birth weight
twins. Although there were few differences in the interaction patterns between mothers and each of
their twins at 4 months of age, by 8 months all mothers in the study showed more positive behav-
ior toward the healthier of the two twins. That is, maternal preferences were clearly linked to the
baby’s health, mediated, quite surely, by the differential behavior and appearance of the two siblings
(Sameroff and Suomi, 1996).
Other factors that influence maternal investment include the child’s age, such that older children
(who, by living as long as they have, demonstrate viability) often receive more investment than
younger children, particularly in times of high stress and low resources (Daly and Wilson, 1988);
mother’s reproductive status, with younger mothers being more likely to neglect, abuse, or kill their
infants than older mothers (Daly and Wilson, 1988; Lee and George, 1999), presumably because
younger mothers have greater opportunity for having more children than older mothers; and social
support, with mothers who have little social support being more likely to abandon an infant than
mothers with greater social support (Daly and Wilson, 1988; Lancaster, 1989).
We have provided evidence of neglect, abuse, and infanticide to illustrate the extremes that parents
sometimes go in making decisions about parental investment. From a broader perspective, human
parents are generally supportive of their children, with abuse and infanticide being relatively rare
phenomena. However, the circumstances under which humans abandon their infants are similar to
those seen for many other species. Hrdy (1999) suggested that mothers may kill their own infants
when other means of birth control are unavailable and they were unwilling or unable to commit
themselves to further care of the infant. However, with notable exceptions, mothers rarely plan to kill
their babies. To quote Hrdy (1999, p. 297):
Rather, abandonment is at one extreme of a continuum that ranges between termination of

investment and the total commitment of a mother carrying her baby everywhere and nurs-
ing on demand. Abandonment is, you might say, the default mode for a mother terminating
investment. Infanticide occurs when circumstances (including fear of discovery) prevent a
mother from abandoning it. Although legally and morally there is a difference, biologically
the two phenomena are inseparable.
Stepparent investment. Incidence of neglect, abuse, and even death, although still rare, are more
likely to occur at the hands of stepparents than biological parents. From a strictly inclusive fitness
perspective, any resource a stepparent provides to stepchildren will not benefit that parent’s fit-
ness. The stepchild possesses none of the stepparent’s genes, and presumably the adult’s resources
could be better spent supporting her or his own genetic offspring. Yet, stepparenting is widespread
throughout the world and through recorded history, and despite the myths and realities of the fate of
children at the hands of stepparents, the vast majority of stepparents love and care for their children
(Hetherington and Stanley-Hagan, 2002). Perhaps the first question we should ask is, “Why should
a stepparent provide any resources to a stepchild?”
Most evolutionary psychologists have suggested that parental investment from a stepparent is
actually investment in mating, not investment in parenting (e.g., Anderson, Kaplan, and Lancaster,
1999). For example, a stepfather provides support to his wife’s children from a previous mating to
maintain her companionship and for the children he will father with her. Women with children
from a previous male select men who will not only provide support for themselves and their future
offspring, but also for their children from previous matings.
21
But stepparents rarely provide the same level of support to their stepchildren as they do to their
natural children. Research from a wide range of cultures indicates that the amount of financial
resources parents provide, and the amount of time spent interacting with stepchildren, is significantly
less than for natural children (e.g., Flinn, Leone, and Quinlan, 1999; Marlowe, 1999). For example,
Anderson and his colleagues (1999) found that the amount of financial resources children in the
United States are likely to receive for their college education was considerably less for families
that consisted of a stepfather and a biological mother than for families consisting of two biological
parents. In an observational study of the Hazda, biological fathers communicated, played with, and
nurtured (held, fed, pacified, cleaned) more their natural children than their stepchildren (Marlowe,
1999), despite adults’ claims of equal feelings and care for natural and stepchildren.
Not providing as much money for college for a stepchild as for a biological child, or playing more
with a natural child than with a stepchild, reflects pancultural differences in the amount of parental
investment made to biological versus nonbiological offspring, consistent with the tenets of parental
investment theory. But having a smaller college fund than a biological offspring is far from stepchild
abuse. Yet, all cultures appear to have their own versions of Cinderella. Such folklore, unfortunately,
has a basis in reality.
Child abuse and homicide are both more likely when a child lives with a stepparent than with
two biological parents (Daly and Wilson, 1988, 1996). In a Canadian study, Daly and Wilson (1985)
reported that children were 40 times more likely to be abused if they lived with a stepparent than
with two natural parents. Differences remained substantial even after potentially confounding fac-
tors such as poverty, mother’s age, and family size were statistically controlled. Perhaps even more
disturbing are findings for child homicide, a crime that, unlike child abuse, is almost always reported
to authorities. Daly and Wilson (1988) examined the results of several surveys of crime data from
around the world and reported a similar pattern independent of country: Children were more likely
to be killed by a stepparent than by a genetic parent, with this difference being particularly large for
children under 2 years of age. Rates of child homicide were sometimes more than 100 times larger
for stepchildren than for biological children. Even the risk of unintentional death (e.g., drowning) is
greater in stepfamilies than in biologically intact families. For example, in one Australian study pre-
school children’s unintentional deaths were significantly higher for children from stepfamilies than
for children from intact or single-parent families (Tooley, Karakis, Stokes, and Ozanne-Smith, 2006).
Murder of one’s stepchildren, of course, is not sanctioned by modern societies. (Although in some
contemporary hunter-gatherer societies, when a man marries a woman with children, it is acceptable
for her young children be put to death; see Daly and Wilson, 1988). Men who kill their stepchildren
are inevitably convicted and incarcerated, so there is no adaptive value to killing a stepchild. Yet, the
fact that abuse and murder of stepchildren are enormously greater than for natural children suggests
that the restraints against acting violently toward nonrelated children are much less than the restraints
involved with one’s genetic children. The love and affection that parents “naturally” feel toward their
biological children must be nurtured, often with substantial effort, to be felt for stepchildren. We are
not suggesting that killing stepchildren, or unrelated children in general, was once adaptive in our
evolutionary past, and that the higher rates of abuse and homicide observed for stepchildren than
for natural children represent the activation of these atavistic mechanisms. Rather, we argue that in
high-stress situations in which violence is apt to occur, the evolved tendencies that inhibit aggressing
against one’s biological children are not as easily activated for one’s stepchildren.
Is “Parenting” the Right Concept to Describe Human Childrearing?

From what we have presented to this point, as well as from a common-sense perspective, there should
be no debate that parents play a key role in their children’s development. However, the “parent-
ing” model that most U.S. Americans seem to adopt—with parents (and teachers) actively teaching
22
their children the skills and values of their culture—may be at odds with how our forechildren
were treated and developed. For example, anthropologists have reported that parents in traditional
societies, including hunter–gatherer societies, rarely explicitly teach their children; rather, children
acquire the skills and norms of their culture through observation and during play with mixed-age
peers (e.g., Konner, 2010; Lancy, 2015). A similar point was made by Scarr (1992, 1993) more than
25 years ago (quite controversial at the time, see Baumrind, 1993; Jackson, 1993), who proposed that
“ordinary differences between families have little effect on children’s development, unless the fam-
ily is outside of a normal, developmental range. Good enough, ordinary parents probably have the
same effects on their children’s development as culturally defined super-parents” (Scarr, 1992, p. 15).
In other words, “superparenting” is not required to rear a successful adult; rather, children adapt to
variations in childrearing, which, claimed Scarr, is a product of natural selection. A species such as
Homo sapiens, that lives in varied environments and under a broad array of cultural traditions, must be
flexible to the vagaries of “ordinary” parenting if the species is to continue.
More recently, Gopnik (2016) has made a similar argument, proposing that the popular term “par-
enting” implies shaping children’s lives with the goal of creating successful adults. Gopnik likens this
method to being a carpenter, where one shapes the material into a product of your choice. According
to Gopnik, the stress for parents trying to shape their children into successful adults is high and the
expectations set by parents can cause undue stress in their children. Gopnik (2016) suggests a differ-
ent metaphor for parenting, that of the gardener. Instead of trying to shape children into successful
adults, parents should provide a caring environment in which children can grow and develop, just as
gardeners do with their plants. She argues children have evolved to develop in this way; it was the
way our ancestors developed, and it is still relevant for our modern world. As previously discussed,
humans are unique in that we have a prolonged juvenile period that includes childhood and adoles-
cence. This prolonged period of immaturity allows for children to explore their surroundings, and
they are intrinsically motivated to do so. Children learn about their environment and how the world
works through their exploration, skills that will help them to successfully navigate adulthood. Play
is also an essential aspect of childhood; children engage in pretend or fantasy play, which helps them
learn about interacting with others and provides practice for dealing with hypothetical situations and
symbols. Gopnik argues that, consistent with the way children evolved to learn as reflected by the
childhoods of hunter-gatherers (Konner, 2010; Lancy, 2015), parents should not tell their children
how to play or shape them into what they desire or consider successful, but instead to provide an
environment that is safe, nurturing, and stable to allow them to explore the world’s possibilities.
Conclusions
Ever since Darwin, there has been the recognition of continuity in cognitive and social function-
ing among different species. Homo sapiens share a heritage with other primates and mammals, and
evolutionary theory provides the means of assessing that heritage. In many ways, when it comes to
parenting, humans are just another mammal. They invest substantially in their offspring, with females
investing disproportionately more than males; they consider the availability of resource and the likeli-
hood of a “payoff ” when devoting resources to their children; males’ and grandparents’ investment
is based partly on degree of genetic certainty; and the sex differential in parental investment dictates,
to a substantial degree, the ways females and males relate to one another. Yet, in other ways, human
parenting is different from that of other mammals, and such differences are also predicted from evo-
lutionary theory. Because of the confluence of a number of factors, including a big brain and the
cognitive ability that accompanies it, increased social complexity, and, most critically we believe, an
extended period of youth, human children require greater investment to reach maturity than the
young of other primates. This period of prolonged youth means that fathers must contribute more
to their offspring if they are to be successful than is the case for the vast majority of males from other
23
mammalian species. This has led directly to the formation of the human family, which, although tak-
ing many specific forms, is universal in our species. Evolutionary theory provides the “big picture”
for how the human family and our particular way of parenting has come about. It is a fascinating
story, we believe, but it is more than just history; it also provides a perspective that helps us understand
important issues of parenting in contemporary societies.
Many people unfamiliar with evolutionary theory assume that it is concerned only with species
universals—traits that characterize all normal members of the species (or all members of one sex).
Individual differences, the argument goes, are ignored or handled poorly by evolutionary theory. As
the examples provided earlier in the application of parental investment theory to the phenomenon
of child abuse and to quality of attachment indicate, this depiction of evolutionary theory is inac-
curate. Evolutionary psychological approaches consider how evolved mechanisms become expressed
over development as a function of local ecological conditions. Although evolutionary psychology
certainly proposes there are universal mechanisms characterizing members of a species, this is not
equivalent to proposing hard-wired “instincts,” impervious to environmental variations. Just as an
individual inherits a species-typical genome, she or he also inherits a species-typical environment.
Both genome and environment are constrained, in that there can be only so much variation to still
produce a viable organism. But that variation can be substantial, and evolutionary theory can be use-
ful in predicting and explaining individual differences in important social, emotional, and cognitive
realms, and possibly suggesting means to deal with persistent societal problems.
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2
PSYCHOBIOLOGY OF MATERNAL
BEHAVIOR IN NONHUMAN
MAMMALS
Aya Dudin, Patrick O. McGowan, Ruiyong Wu,
Alison S. Fleming, and Ming Li
Introduction
Mammalian mothers of different species may differ in the extent to which physiological and psy-
chological factors contribute to the postpartum expression of their nurturant behavior. In all species
that have been studied, however, the physiological determinants are only realized in individuals that
have had certain developmental histories and that are psychologically “prepared” by their physical
and psychological environments (Rosenblatt and Snowdon, 1996). In many mammalian species the
hormonal changes associated with late pregnancy and parturition predispose the newly parturient
female to be nurturant with her offspring, to nurse, clean, and protect them. However, whether
these nurturant behaviors in fact occur at the appropriate time and in the appropriate way depends
on a host of psychological and environmental factors. Enhanced morbidity or mortality of young
or reduced responsiveness by mammalian mothers occurs if mothers are stressed during pregnancy
or parturition, severely food deprived, or are placed in low-resourced environments (Lyons, Kim,
Schatzberg, and Levine, 1998), if ambient temperature precipitously rises, if pups are sickly, or if
the nesting area is inadequate (Herskin, Jensen, and Thodberg, 1998; Kinsley, 1990; Leon, Cooper-
smith, Beasley, and Sullivan, 1990). However, mothers are also robust; to eliminate maternal behavior
entirely, environmental restrictions or debilitations experienced by mothers have to be extreme and
depend on the animal’s background and genetics (Aubert, Goodall, Dantzer, and Gheusi, 1997; Jen-
kins, McGowan, and Knafo-Noam, 2016; McGuire, Pachon, Butler, and Rasmussen, 1995; Mileva-
Seitz, Bakermans-Kranenburg, and van IJzendoorn, 2016; Stolzenberg and Champagne, 2016b). In
nonhuman primates, examples of the importance of interactive influences of early experiences and
hormones on mothering are still more pronounced. Harlow’s “motherless” monkeys raised in social
isolation and on wire mothers, or monkeys who are young and inexperienced, neglected or bat-
tered their own (usually first) offspring, despite having apparently normal pregnancies and child-
births (Brett, Humphreys, Fleming, Kraemer, and Drury, 2015; Coe, 1990; Harlow, 1963; Ruppenthal,
Arling, Harlow, Sackett, and Suomi, 1976). In human beings, a host of background and psychological
factors increases risk of mothering disorders, including poverty, low education, social isolation, lack of
supports, and immaturity, being themselves victims of abuse (Agrati and Lonstein, 2016b; Barrett and
Fleming, 2011; Britton, 2008; Brummelte and Galea, 2016; Correia and Linhares, 2007; Daly, 1990;
Dennis, Heaman, and Vigod, 2012; Di Florio et al., 2013; Eisenberg, 1990; Lancaster, Gold, Flynn,
and Yoo, 2010; Lomanowska, Boivin, Hertzman, and Fleming, 2017; Milgrom et al., 2008; O’Hara
30
Maternal Behavior in Nonhuman Mammals
and Swain, 1996; van Bussel, Spitz, and Demyttenaere, 2009; Vesga-López et al., 2008; Viguera et al.,
2011) and their genetics (Mileva-Seitz et al., 2016).
Even in “lower” species and situations where hormones exert clear and powerful influences on
maternal behavior, the behaviors will not occur or will be masked by competing responses given dys-
functional past or present experiences. Conversely, in primates and human beings, in particular, the
clear importance of these background and situational factors may seem to mask the role of biological
factors in early mothering; however, a variety of approaches help to unmask contributions of these
biological influences (see Fleming, Lonstein, and Lévy, 2016).
This chapter provides a summary of the interaction between psychological and physiological
influences in the expression of maternal behavior in nonhuman mammalian mothers. We first
briefly discuss sensory, experiential, and other psychological factors that regulate maternal behavior
in selected nonprimate mammals (including altricial species with litters: primarily rats and mice,
and also precocial species with one or two young: e.g., sheep). We then summarize neural bases,
neurochemical mechanisms, and genetics of maternal behavior, focusing on research on the roles of
dopamine and serotonin in the limbic and hypothalamic regions and in their related genes.
The specific approach adopted here assumes that maternal behavior is not regulated in a unitary
fashion, but depends for its expression on activation of a variety of behavioral systems, mediated by
multiple neurochemical and neuranatomical substrates. It assumes further that hormones and asso-
ciated neurochemicals do not automatically trigger behavior, but instead act on substrates whose
activation is influenced by the animal’s social/psychological and physical environments, both past and
present. The interaction between newborn and the mother alters the basic mechanism of behavioral
expression in both.
In this chapter, the maternal behavior of rodents receives considerably more attention than
maternal behavior in other nonprimate mammals. This orientation is based on a number of con-
siderations. First, comparisons among rat and other nonprimate mammals (where appropriate data
exist) show many similarities in physiological regulation of parenting (Lonstein, Levy, and Fleming,
2015). In addition, most research on mechanisms of maternal behavior focuses on Rattus norvegicus
mothers, and as a result, understanding of this species is more complete. Finally, despite large differ-
ences in behavioral organization between the rat and human maternal behavior, the rat has proven
to be a productive model for the analysis of human maternal behavior and has provided insights
into possible mechanisms at work in human beings (Corter and Fleming, 1990; Fleming, Ruble,
Krieger, and Wong, 1997; Fleming, Steiner, and Corter, 1997; Lonstein et al., 2015; Tombeau
Cost et al., 2016). However, because of the extensiveness of research on rodent maternal behavior,
this review does not attempt to be complete or exhaustive (Bridges, 2016b; Numan and Young,
2016; Olazábal et al., 2013a, 2013b). Its approach reflects most strongly the views and work of the
authors.
The chapter is divided into a number of sections. The first section describes the basic techniques
used in the study of maternal behavior in rodents. The second section discusses the effects of the
maternal hormones on maternal behavior and on other behaviors that undergo change when a
female gives birth. This section also describes the transition that occurs in the regulation of maternal
behavior after the initial period of hormonal priming and emphasizes factors regulating the long-
term maintenance of maternal behavior. In this section the roles of learning, memory, and reinforce-
ment are considered. The third section of the chapter considers the role of sensory factors in the
onset and maintenance of maternal behavior, with particular attention to olfactory and somatosen-
sory input during mother-litter interactions. The fourth and fifth sections of the chapter describe
in considerable detail what is known about the neurochemistry and neuroanatomy of maternal
behavior and of other behaviors that co-occur with maternal behavior in the postpartum animal.
The sixth section briefly describes work on the genetic and epigenetic bases of maternal behavior.
31
Aya Dudin et al.
The final section of the chapter summarizes some of the main themes raised in the chapter and issues
yet to be explored.
Methodological Issues in the Study of Rodent Maternal Behavior

The popularity of the rodent (mouse and rat) as the animal of choice in the analysis of the physiol-
ogy of parenting is based on both practical and scientific considerations. The laboratory rodent is an
easy animal to breed, care for, and test. Its use permits the application of experimental manipulations
providing greater control over physiological or psychobiological variables that is not possible with
nonlaboratory animals, primate models, or human beings. For instance, to understand the role of
hormones in the regulation of parenting we can analyze the behavior before and after the removal of
the gland that produces the suspected hormone or before and after the administration (by injection
or capsule) of the suspected hormone. Using a similar “extirpation and replacement” paradigm, we
can evaluate the involvement of different sensory systems or different neural circuits in behavioral
regulation by observing behavior before and after the destruction of specific neuronal cell groups
within the brain by a variety of lesioning techniques (Afonso, Sison, Lovic, and Fleming, 2007; Li and
Fleming, 2003a, 2003b) or by the use of psychoactive drugs (Li, 2015) that either block or enhance
the functioning of specific neurotransmitters in the brain. Conversely, we can attempt to augment
or facilitate the expression of the behavior in initially nonmaternal animals by the application of
electrical (Morgan, Watchus, Milgram, and Fleming, 1999b), hormonal (Fleming, Cheung, Myhal,
and Kessler, 1989a), or drug stimulation (Kinsley et al., 1994; Wu, Gao, Chou, Davis, and Li, 2016)
that mimics the action of naturally occurring neurotransmitters. In some cases, when a longitudinal
design is impractical, different groups sustaining different experimental and control conditions are
compared. Thus, for instance, we might test the behavior of a group of animals which have been
injected with a particular hormone or chemical that we suspect is normally released when animals
become maternal at parturition and compare their maternal behavior with the behavior shown by
a control group of animals which have received injections containing a related but biologically inert
substance (i.e., placebo).
Convergent with these invasive experimental approaches, we can also undertake correlational,
rather than experimental, analyses and relate changes in physiology with changes in ongoing behav-
ior; thus we can explore electrical events or hormonal or neurochemical changes that occur when
an animal expresses maternal behavior. For instance, we can use microdialysis plus high performance
liquid chromatography (HPLC) techniques to measure neurotransmitters that are released during
ongoing behavior (Afonso, Shams, Jin, and Fleming, 2013). Alternatively, by means of immunocy-
tochemical staining techniques we can determine whether particular proteins, neuropeptides, or
other brain chemicals are produced in the mother’s brain in response to interactions with offspring
(Numan and Numan, 1994; Zhao and Li, 2012). The role of early experiences in the development
of the behavior can be studied by comparing the adult behavior and neurobiology of animals that
have been raised under different early environmental conditions, with or without a mother or sib-
lings or with and without certain nest-related cues (Rees, Akbari, Steiner, and Fleming, 2008; Rees,
Steiner, and Fleming, 2006). Effects of adverse early experiences prenatally as well as postnatally can
be assessed by evaluating the adult behavior of offspring of mothers who are stressed, malnourished,
or who are administered agents or toxins, like antipsychotics, cocaine, or alcohol (Francis and Kuhar,
2008; Li, 2015). The effects of genotype and these maternal effects can then be partialled out by
comparing animals that have been raised by their own mothers and those raised by foster mothers.
Least well formulated are the technologies associated with establishing which genes or gene
complexes regulate adult maternal behavior (Akbari et al., 2013). Strategies that have been employed
to study heredity and genetic factors in the regulation of maternal behavior include comparison
of different strains, cross-fostering within and between strains (Gomez-Serrano, Tonelli, Listwak,
32
Sternberg, and Riley, 2001), the analysis of transgenic mice mutants that lack specific genes (so called
“knockout” mice) that underlie the production of specific proteins and receptors in brain that are
involved in the expression of maternal behavior (Caldwell, Aulino, Freeman, Miller, and Witchey,
2016), and use of molecular techniques that assess the animal’s genetic profile and activation of par-
ticular genes during ongoing behavior (Ribeiro, Agmo, Musatov, and Pfaff, 2016). As well, the use
of functional magnetic resonance imaging techniques in awake rats allows a delineation of the func-
tional neural circuitry and exploration of roles of neuropeptides, such as oxytocin and vasopressin, in
the maternal brain (Febo, 2011; Febo, Felix-Ortiz, and Johnson, 2010; Febo and Ferris, 2014). Finally,
newer techniques that have not yet been adequately utilized in the study of maternal behavior
include optogenetics, which would permit the specific manipulation of individual neurons within
the maternal neural circuit (Wu, Autry, Bergan, Watabe-Uchida, and Dulac, 2014). In the discussion
that follows, many of the techniques, strategies, or approaches described above have been adopted
to augment our understanding of the physiology of parenting. To facilitate navigation through the
somewhat more technical portions of this chapter we provide a brief description of the relevant ter-
minologies and techniques at the beginning of some of the more technical sections.
Description of Maternal Behavior at Parturition

Although the study of maternal behavior in rodents has generated a smaller literature than has the
study of many other species-typical behaviors, it is by no means a new area of interest. In fact, some
of the most detailed and informative descriptions of the rat mother-litter interactions were provided
by Wiesner and Sheard in their seminal book Maternal Behavior in the Rat, published in 1933.
Based on this long history of research, we have a relatively complete picture of the phenomenol-
ogy of rat maternal behavior. The new mother rat is maternally responsive to newborn pups as soon
as they emerge from the birth canal (Hudson, Cruz, Lucio, Ninomiya, and Martinez-Gomez, 1999;
Rosenblatt, Lehrman, and Rheingold, 1963). At parturition, she pulls off the amniotic sac, eats the
placentas, and cleans off the pups (Hudson et al., 1999; Kristal, Thompson, Heller, and Komisaruk,
1986). Within the first 30 minutes after parturition, she gathers all the pups together, retrieves them
to a nest site, mouthes and licks them, and adopts a nursing posture over them; she does all this with-
out prior experience interacting with pups (Fleming and Rosenblatt, 1974a).
When a female interacts with her litter after the birth, she engages in many proximal interactions.
After the nest has been constructed and the pups have been retrieved into it, the dam spends a con-
siderable proportion of her time mouthing and licking the pups, especially their anogenital regions
(Moore, 1990), a behavior that functions to promote urination and elimination by the offspring and
to maintain the dam’s fluid balance (Friedman, Bruno, and Alberts, 1981). The dam also gathers the
young underneath her ventrum, permitting pups to gain access to her teats and suckle (Stern, 1989).
Once the pups begin to suckle, the dam usually adopts a high arch crouch posture over them and
becomes immobile for a period (Stern, 1989, 1990; Stern, Dix, Bellomo, and Thramann, 1992). The
high motivational state of the new mother is illustrated by observations that, if new mothers are
prevented from actively exhibiting these proactive maternal behaviors (by application of a muzzle
over their snouts), they nevertheless spend considerable time nudging and pushing at the pups and
manipulating them with their forepaws (Stern and Keer, 1999). In addition, they will overcome their
anxiety to retrieve pups from the open arms of an elevated plus maze into the closed arms (Yang
et al., 2015). The virgin animal, by contrast, is not maternally responsive when first presented with
newborn foster pups (Rosenblatt, 1967; Wiesner and Sheard, 1933), and she does not retrieve pups
on the elevated plus maze (Yang et al., 2015). In fact, initially she moves away from pups and actively
avoids them (Fleming and Luebke, 1981b; Terkel and Rosenblatt, 1971). However, within 1 to 2
days of continuous pup stimulation, the virgin becomes habituated to pups and is willing to lie down
in close proximity to them (Fleming and Luebke, 1981b; Fleming and Rosenblatt, 1974a; Terkel
33
Aya Dudin et al.
and Rosenblatt, 1971); after 5 to 10 days of continuous contact with foster pups, the virgin begins
to respond maternally (Rosenblatt, 1967), showing a pattern of behavior that resembles that of the
new mother, but also showing some differences (Lonstein and De Vries, 1999). This experimental
procedure has come to be known as “pup-induction” or “pup-sensitization”, and is useful to study
the influences of parturitional hormones on maternal behavior.
Associated with changes in actual responses to pups, new mothers undergo changes in multiple
psychological states which contribute to these responses. These changes are in the realms of moth-
ers’ cognition, memory, executive functions, impulsivity, emotion regulation, and anxiety (Lonstein
et al., 2015). Changes in these functions are reflected, either immediately after parturition or later in
the postpartum period, in the pattern of interactions with the young. They are, in turn, regulated by
neural and neurochemical systems that intersect with mechanisms more directly implicated in moth-
ering per se. Similar psychological changes occur in species ranging from rats to humans (Barrett
and Fleming, 2011; Lomanowska et al., 2017; Lomanowska and Melo, 2016; Lonstein et al., 2015).
Hormonal Effects on the Onset of Maternal Behaviors

Although early endocrine studies (Beach and Wilson, 1963; Lott and Fuchs, 1962; Riddle, Lahr, and
Bates, 1935) did not provide conclusive evidence for endocrine involvement in the regulation of
maternal behavior, those studies provided an approach to the analysis of the hormonal control of
behavior, using extirpation and replacement strategies. There is now substantial evidence that the
hormones associated with late pregnancy and the parturitional period acting on brain receptors
(Bridges, 2016b) account for the rapid activation of maternal responsiveness seen at parturition (Ter-
kel and Rosenblatt, 1972). These include the steroid hormones, estradiol and progesterone, which are
synthesized by the ovaries and released into the circulatory system and the protein hormones, vaso-
pressin, prolactin, and oxytocin, which are released within the brain and from cells or nerve terminals
within the “master” endocrine organ, the pituitary gland (Beery, McEwen, MacIsaac, Francis, and
Kobor, 2016b; Bridges, 1990; Bridges, 2016a; Feldman, 2016; Insel, 1990; Numan, 1994; Numan and
Young, 2016; Olazábal and Alsina-Llanes, 2016; Rosenblatt, 1990; Rosenblatt and Snowdon, 1996;
Stolzenberg and Champagne, 2016a). Other important neurotransmitters in regulation of mother-
ing include dopamine, norepinephrine, serotonin, GABA, and endorphins (Afonso, King, Novakov,
Burton, and Fleming, 2011; Afonso, Grella, Chatterjee, and Fleming, 2008; Afonso, King, Chatter-
jee, and Fleming, 2009; Afonso et al., 2013; Blass, 1996; Chen et al., 2014; Gao, Wu, Davis, and Li,
submitted; Hansen, Bergvall, and Nyiredi, 1993; Kendrick, Keverne, Chapman, and Baldwin, 1988;
Keverne, 1988; Numan and Young, 2016; Smith, Piasecki, Weera, Olszewicz, and Lonstein, 2013;
Wu, Gao, Chou, Davis, and Li, 2016). These hormones and neurochemicals serve multiple functions.
They prepare the prospective mother physiologically, by acting on mammary tissue prior to the
initiation of lactation (Tucker, 1994) and by acting on the uteri, first to maintain the integrity of the
implanted conceptus, and then to promote uterine contractions and parturition as well as analgesia
during the birth process (Hodgen and Itskovitz, 1988; Kristal et al., 1986). These hormones also
contribute to elevated maternal responsiveness and reduced anxiety shown by the newly parturient
mother (Bridges, 1990; Ferreira, Pereira, Agrati, Uriarte, and Fernandez-Guasti, 2002; Insel, 1990;
Lonstein, 2007a; Pereira and Ferreira, 2016; Rosenblatt, 1990). A regimen of hormones designed to
simulate these pregnancy and parturitional changes administered to maternally inexperienced virgins
can induce a very rapid onset of retrieval, crouching, and licking in response to foster pups (Bridges,
1990; Insel, 1990; Rosenblatt, 1990).
Although the “parturitional” hormones might seem to activate maternal behavior in a unitary
fashion, in fact the different hormones and neurochemicals probably exert somewhat different
behavioral effects, and any one hormone or neurochemical probably exerts multiple effects. Moreo-
ver, their varied effects probably result from their action on a variety of different neural pathways.
34
Figure 2.1 Functional neuroanatomy mediating maternal and related psychological processes in mammals.
Neuroanatomical structures include olfactory bulbs, amygdala, nucleus accumbens, bed nucleus of
stria terminalis (BNST), medial preoptic area (MPOA), ventromedial hypothalamus (VMH), ante-
rior hypothalamic nucleus (AHN), ventral pallidum (VP), prefrontal cortex (PFC), ventral tegmental
area (VTA), doral and median raphe (DR and MR), and parietal cortex. Relevant neurochemistry
include catecholamines, serotonin (5-HT), dopamine (DA), the neuropeptides, and opioids.
Source: Adapted from Fleming, O’Day, and Kraemer (1999).
For instance, the hormones progesterone and estradiol might facilitate the expression of maternal
behavior by altering a number of behavioral propensities, and these alterations provide the behavio-
ral environment in which maternal responses can be most easily expressed (Fleming, 1987; Fleming
and Corter, 1988). Specifically, as schematized in Figure 2.1, these hormones promote changes in
the female’s attraction to the odors of pups, reduce her fearfulness in their presence, and facilitate the
ease with which she learns about their characteristics, possibly by augmenting the pups’ reinforc-
ing value. Together, these hormonal effects are seen to indirectly augment maternal responsiveness
during the periparturitional period by reducing the competing effects of alternative nonmaternal
behaviors and by insuring that dams will continue to respond to pups when the period of hormonal
priming ends (Fleming et al., 2016; Lonstein et al., 2015). Thus, as shown in the following discus-
sion, postpartum animals differ from virgins on a number of psychological dimensions because of
the action of hormones.
Emotional and Anxiety Changes in the New Mother

Naturally parturient females are less avoidant when presented with pups than are virgin animals.
More generally, they are less neophobic, being more willing to approach an unfamiliar intruder and
to enter and explore a novel environment (Fleming and Luebke, 1981a; Lonstein, 2007a; Lonstein
et al., 2015; Pawluski, Lonstein, and Fleming, 2017).
In postpartum laboratory rats, these emotional changes can be found using many paradigms
within 24 hours after parturition, last for about one week, and require recent physical contact with
the litter although suckling per se is unnecessary (Lonstein, 2005, 2007; Ragan and Lonstein, 2014).
In fact, some reduction in neophobia, fear, and anxiety even accompanies the maternal state in
35
Aya Dudin et al.
sensitized nulliparous female rats (Agrati, Zuluaga, Fernandez-Guasti, Meikle, and Ferreira, 2008 Fer-
reira et al., 2002 Pereira, Uriarte, Agrati, Zuluaga, and Ferreira, 2005). However, the relation between
anxiety and mothering is not linear. It has also been suggested that in genetically “unselected” ani-
mals, a moderate level of anxiety that is neither too high (rendering dams over-reactive to threat) nor
too low (rendering them naively under-reactive) is optimal for maternal ability to focus attention on
the needs of the pups despite threats in the environment (Ragan and Lonstein, 2014). Anxiety effects
also depend on the context. Studies of the relation between natural variation in anxiety and mother-
ing in laboratory rodents find no significant relation when tested under relatively benign conditions
(Curley, Jensen, Franks, and Champagne, 2012), but mother rats genetically selected for high anxiety
are more effective mothers under challenging conditions compared to low-anxiety dams (Neumann,
Krömer, and Bosch, 2005). Thus, maternal anxiety state becomes most influential for rats in risky
environments that could disrupt maternal motivation to remain with the pups and provide them
care. However the relation is complex, as work by Bosch (2011) shows that high-anxiety mother rats
spend more time in the nest, retrieve pups faster, and show enhanced maternal aggression in com-
parison to low-anxiety rats. This seems to suggest that under some circumstances elevated anxiety is
positively correlated with enhanced maternal performance (Bosch, 2011).
These emotionality differences between postpartum and virgin females appear to be hormonally
and/or neurochemically mediated (Agrati and Lonstein, 2016a; Fleming, 1986; Fleming et al., 1989a;
Lonstein, 2005; Lonstein, 2007b; Pereira and Ferreira, 2016). The regimen of progesterone and estra-
diol that facilitates the onset of maternal behavior in the virgin rat also reduces pup-avoidance and
measures of timidity in an open-field apparatus (Fleming, Cheung, Myhal, and Kessler, 1989b). At
a brain level, the postpartum reduction in postpartum anxiety is likely maintained by neurochemi-
cals released in the brain when mothers physically interact with the litter; removing the ovaries, or
adrenal or pituitary glands, does not affect the anxiety-related behavior of postpartum rats, whereas
preventing the physical contact with pups before testing results in increases the dams’ anxiety-like
behaviors (Agrati and Lonstein, 2016b). Among the neuropeptides implicated in postpartum anxiety
are vasopressin, oxytocin, and GABA (Lonstein et al., 2015; Pawluski et al., 2017).
Moreover, lactating animals show marked hyporesponsiveness of the stress system and differences
between virgin and lactating females in their hypothalamic-pituitary-adrenal responses to stressors; in
comparison to mothers, virgins show enhanced stressor-induced release of hypothalamic-pituitary-
adrenal (HPA) related hormones (corticosterone, adrenocorticotropic hormone [ACTH]), as well
as enhanced baseline corticotrophin-releasing factor (CRF) in the paraventricular nucleus (PVN)
region of the hypothalamus (Neumann et al., 1998; Windle et al., 1997). These HPA differences are
associated with differences on a variety of emotionality tasks (Hard and Hansen, 1985; Neumann,
2001; Neumann, 2003; Neumann et al., 1998; Neumann, Torner, and Wigger, 2000; Silva, Bernardi,
Nasello, and Felicio, 1997b).
Finally, the assumption that reduced timidity contributes to elevated maternal responsiveness is
supported by findings that manipulations that reduce the animals’ timidity or anxiety, like benzodi-
azepines (Hansen, Ferreira, and Selart, 1985) or early handling (Mayer, 1983), also facilitate maternal
responding. Panesar and Fleming (Panesar, Rees, and Fleming, 2000) found that high concentrations
of glucocorticoids injected into an adrenalectomized virgin animal inhibits the expression of many
components of pup-induced maternal behavior, whereas the same high concentration facilitates
maternal behavior in the postpartum animal.
Sensory Changes in the New Mother

In addition to their effects on dams’ affective state, parturitional hormones also alter their responsive-
ness to pup-related cues. In the following discussion we first describe hormonal effects on olfactory-
mediated responses and then on their somatosensory processing.
36
Odor Cues
The study of the sensory control of maternal behavior was pioneered by Beach and Jaynes in 1956;
it involved observing maternal behavior in experienced mother rats after the systematic removal of
the different sensory systems, either singly or in combination (Beach and Jaynes, 1956a, 1956b). This
early study suggested that no single sensory system is essential for the expression of the behavior but
that their combined removal produces additive deficits, leading Beach and Jaynes to conclude that
maternal behavior is under multisensory control. Although these results seem to apply quite well
to animals that have had maternal experience, we now know that single denervations of a number
of sensory systems can have profound effects on the expression of maternal behavior in the mater-
nally inexperienced animal (Fleming and Rosenblatt, 1974a; Stern, 1989) and that specific sensory
cues from the pups play important roles both in motivating responsiveness and in guiding ongoing
behavior.
In comparison to virgins, new mothers without direct experience with pups prefer nest material
taken from the nest of a new mother and her pups to material taken from a virgin’s nest or clean
material. Virgins show no such preference (Bauer, 1983; Fleming et al., 1989b). Moreover, virgins
treated with a regimen of hormones designed to mimic the parturitional changes in progesterone
and estradiol also exhibit a preference for pup-related odors (Fleming et al., 1989b); the additional
observation that injections of morphine can induce an aversion to pup odors (Bridges, 1990; Kinsley
and Bridges, 1988; Kinsley, Morse, Zoumas, Corl, and Billack, 1995) suggests that, at the time of par-
turition, low concentrations of this neurochemical in relevant parts of the brain heightens attraction
to pup-related odors. Experience with these odors can facilitate maternal responding; adult virgin
rats show more rapid maternal inductions if they have been preexposed to pup odors and vocaliza-
tions during their early development (Gray and Chesley, 1984; Moretto, Paclik, and Fleming, 1986).
Although difficult to demonstrate, it seems that preexposure to pup cues at a distance (primarily
odors and vocalizations) in adulthood also facilitates maternal responding, at least among females
whose responsiveness is high to begin with. Thus, a higher proportion of animals exhibits immediate
maternal behavior during maternal tests if they have been preexposed to pup odors than if they have
not been (Orpen and Fleming, 1987). Finally, if virgins are rendered unable to smell pups by olfac-
tory bulb removal, which mediates the sense of smell, they are not avoidant with pups, but instead
exhibit a very rapid onset of maternal behavior, as though the pups’ odors in the context of other
pup cues are aversive (Fleming and Rosenblatt, 1974b, 1974c; Fleming, Vaccarino, Tambosso, and
Chee, 1979).
New mothers develop an attraction to pup-related odors and are guided by them. However, in
rats neither the main (airborne) nor the accessory (where activating molecules are transmitted nor-
mally by means of an aqueous medium) olfactory systems are essential for mothering behavior to
be expressed. Following destruction of the olfactory mucosa, a normal latency to onset of maternal
behavior occurs in primiparous females (Benuck and Rowe, 1975; Jirik-Babb, Manaker, Tucker, and
Hofer, 1984; Kolunie and Stern, 1995), even though retrieval may be delayed because dams who
cannot detect odors (anosmic) take longer to locate pups at a distance (Benuck and Rowe, 1975;
Kolunie and Stern, 1995). Similarly, no deficit in maternal behavior is observed after removing the
vomeronasal organ (VNO) or cutting the vomeronasal nerves, which eliminate input from the acces-
sory olfactory system (Fleming, Gavarth, and Sarker, 1992; Jirik-Babb et al., 1984; Kolunie and Stern,
1995). Latency to retrieve is unaffected in these VNO-operated females, suggesting that the acces-
sory olfactory system is not critical for even locating pups. Inconsistent results on maternal behaviors
have been reported using bilateral removal of the olfactory bulbs (bulbectomy) which eliminates
both main and accessory olfactory input. These effects range from profound disturbances and/or
cannibalism (Benuck and Rowe, 1975; Fleming and Rosenblatt, 1974b; Kolunie and Stern, 1995;
Schwartz and Rowe, 1976) to no reported deficits at all (Beach and Jaynes, 1956b; Fleming, Kuchera,
37
Aya Dudin et al.
Lee, and Winocur, 1994); however, peripherally induced anosmia by zinc sulfate reduces licking by
new mother rats (Fleming and Rosenblatt, 1974c).
In contrast to rats, mice mothers depend on olfaction for the regulation of their maternal behav-
ior. In the laboratory mouse, removal of the olfactory bulb (olfactory bulbectomy) prevents nest
building, reduces nursing, and induces cannibalism (Gandelman, Zarrow, and Denenberg, 1971; Gan-
delman, Zarrow, Denenberg, and Myers, 1971; Sato, Nakagawasai, Tan-No, Onogi, Niijima, and
Tadano, 2010b; Vandenbergh, 1973). Moreover, the deletion of genes involved in olfactory signal
transduction (SCN9A or Cnga2) in the main olfactory epithelium results in deficits in pup retrieval
(Fraser and Shah, 2014; Weiss et al., 2011). However, removing accessory olfactory functioning or
deleting Trpc2 (gene coding for ion channels in the vomeronasal organ) does not interfere with the
expression of maternal behavior in mice (Fraser and Shah, 2014; Lepri, Wysocki, and Vandenbergh,
1985). The importance of main olfactory cues for mouse mothering is diminished in experienced
mothers, who can apparently compensate for a loss of olfactory function by using other sensory
information (Dickinson and Keverne, 1988; Seegal and Denenberg, 1974).
In contrast to rats also, maternally discriminating species, like some ungulates that recognize their
young and allow them to suckle while rejecting others, olfaction is key (Lévy et al., 1995; Pitcher,
Harcourt, and Charrier, 2010; Romeyer et al., 1993). Inducing anosmia in sheep mothers (ewes)
before parturition prevents recognition of their own lambs and so any young is accepted to suckle
(Baldwin and Shillito, 1974; Bouissou, 1968; Lévy et al., 1995; Poindron, 1976; Romeyer, Poindron,
and Orgeur, 1994). In this case, the main, but not the accessory olfactory system is involved (Lévy
et al., 1995). Primiparous ewes rendered anosmic before parturition show reduced maternal behavior.
By contrast, females with lesions of the accessory olfactory system show little disturbance in mater-
nal care (Lévy et al., 1995). The olfactory cues that attract ewes to any newborn lamb are linked to
amniotic fluid. Removing amniotic fluid from the neonate’s coat reduces maternal licking and, in
primiparous ewes, prevents acceptance behavior while increasing aggression (Lévy and Poindron,
1987). In experienced mothers, coating lambs in amniotic fluid alone is sufficient to induce maternal
acceptance (Basiouni and Gonyou, 1988; Lévy and Poindron, 1984). Thus, olfactory cues provided
by amniotic fluid ensure appropriate maternal behavior at parturition in sheep, especially in inexpe-
rienced mothers (Corona and Lévy, 2015).
Olfactory Cues Have Inhibitory Effects

In contrast to postpartum rats, in virgin rats the induction of anosmia disinhibits the expression of
maternal behavior. That is, olfactory cues associated with birth and pups prevent nulliparous female
rats from being maternal. Virgin or nonpregnant female rats are simply repelled by placenta, amni-
otic fluid, and pup odor (Kristal, 1980), but anosmia eliminates these aversive properties and a rapid
onset of maternal behavior results (Carretero, Segovia, Gomez, and Del Cerro, 2003; Fleming and
Rosenblatt, 1974b; Fleming et al., 1979). These inhibitory effects seem to be mediated by both the
main and accessory systems and in a number of different species (rat: Fleming et al., 1979; rabbit:
González-Mariscal, Chirino, Beyer, and Rosenblatt, 2004; Chirino, Beyer, and González-Mariscal,
2007; hamsters: Marques, 1979), although of course this not the case in many strains of mice, which
may require olfaction for dam maternal behavior.
In rats, we have no idea which specific pup or pup-related odors influence the dam’s attraction to
pups, although evidence suggests that pup anogenital licking by the mother is facilitated by secretions
from the pup preputial glands (glands around the anal region; Brouette-Lahlou, Vernet-Maury, and
Chanel, 1991). Moreover, the relevant component in the secretion seems to be a pheromone-like
compound called dodecyl propionate (Brouette-Lahlou, Amouroux, et al., 1991). It appears likely
that, because rats do not become attached to individual offspring or even individual litters—although
they respond differentially to pups based on sex (Cavigelli, Ragan, Barrett, and Michael, 2010; Moore
38
and Morelli, 1979)—the odor of individual pups or litters is less relevant than is the odor that charac-
terizes the developmental age of the pups and/or the mother’s postpartum stage. These odors could
derive from many sources in addition to preputial glands, including uterine fluids, mother’s milk,
maternal diet, and maternal excretory products. Mother rats will consistently lick some pups over
others in a litter; however, whether actual “attachment” occurs and/or the reasons for this differential
behavior is unknown (Beery, McEwen, MacIsaac, Francis, and Kobor, 2016a; Pan, Fleming, Lawson,
Jenkins, and McGowan, 2014; Ragan, Harding, and Lonstein, 2016a). In mice, for instance, Doane
and Porter (1978) found that dams could discriminate pups being nursed by females fed the same
diet as themselves from those nursed by mothers fed a different diet. The existence of nest-specific
odors is also suggested by the work of Bauer (1983), Kinsley and Bridges (1988), Kinsley (1990), and
Leon (1978).
In sheep, there is considerable evidence that at parturition the ewe develops an attraction to
amniotic fluids, which had been aversive prior to birth, and this attraction begins to fade by 2 hours
postpartum (Lévy, Poindron, and Le Neindre, 1983; Poindron and Lévy, 1990). This attraction is
apparently induced by the synergistic action of prepartum estrogen, genital stimulation, and oxytocin
release associated with the parturition (Poindron and Lévy, 1990) and functions to enhance the ewe’s
maternal licking and grooming and willingness to accept the neonatal lamb (Lévy and Poindron,
1984, 1987). Finally, olfactory input clearly constitutes the initial basis of the ewe’s selective bond
with her lamb. As indicated above, ewes that are unable to smell their lambs exhibit enhanced mater-
nal behavior to all lambs and do not develop a selective bond with any one lamb (Corona and Lévy,
2015; Keller, Meurisse, and Lévy, 2004).
Touch Cues
Hormones exert their effects on multiple sensory systems. We now discuss effects on somatosensory
function. Rat mothering involves physical interactions with pups which activate the mothers’ soma-
tosensory systems. When mothers mouthe, lick, and retrieve pups they receive tactile input to the
very sensitive mouth (perioral) region. Work by Kenyon, Cronin, and Keeble (1983) and by Stern
(Stern, 1990; Stern and Johnson, 1989; Stern and Kolunie, 1989) indicates that this stimulation of the
mouth region is essential to the complete expression of maternal behavior during the early postpar-
tum period. Stern and her colleagues (Stern and Johnson, 1989; Stern and Kolunie, 1989) found that,
if the mouth region is desensitized through use of a muzzle, anesthesia injected into the cheek region,
or transection of the nerves in this region, mother rats will not exhibit normal crouching behavior.
If anesthetized and transected, they also do not retrieve or lick pups. In fact, tactile stimulation of
the mouth area seems to be necessary for activation of the pronounced nursing posture (ventroflex-
ion) necessary for successful suckling by young (Stern, 1990, 1996). Given the clear importance of
tactile sensation of the mouth and cheek regions, it is interesting that estradiol also enlarges the area
of responsiveness of this facial area (Bereiter and Barker, 1975, 1980; Bereiter, Stanford, and Barker,
1980), presumably heightening maternal sensitivity to pup-generated touch cues.
The ventral trunk region with its teats is obviously another important contributor to somatostim-
ulation. When dams crouch over and nurse pups, they receive tactile input through touch receptors
in the ventral skin surface and teat stimulation by suckling pups. The importance of suckling stimula-
tion for the release of prolactin, glucocorticoids, and oxytocin (the hormones of lactation) and activa-
tion of the milk-ejection or “letdown” reflex is well known (Wakerly, Clarke, and Sumerlee, 1988).
However, the fact that thelectomized females (with teats removed), or females whose individual teats
have been anesthetized, engage in motivated maternal behavior suggests that teat stimulation by
suckling young is not necessary for the retrieval, licking, or hovering over pups, although teat stimu-
lation is clearly necessary for the occurrence of the high crouch involved in nursing behavior (Stern
et al., 1992; Stern, Dix, Pointek, and Thramann, 1990). Similarly, insufficient ventral stimulation due
39
Aya Dudin et al.
to the presence of too few pups, chilled pups, or pups prevented from suckling fails to elicit the high
arch crouch in dams (Stern and Johnson, 1989). Although teat stimulation may not be necessary for
the expression of most maternal behaviors, the ventral surface surrounding the nipples may be. If a
dam is given a local anaesthetic that desensitizes the ventrum (Stern and Johnson, 1989) or wears a
specially devised spandex jacket covering the ventrum (Morgan, Fleming, and Stern, 1992), pups do
not gather under the mother’s ventrum and attach to the teats but instead gravitate to her exposed
neck region, where the fur presumably has the right tactile, temperature, and odor characteristics
(Magnusson and Fleming, 1995; Morgan et al., 1992). Under these conditions, licking and a variety
of other maternal behaviors are considerably distorted.
Although such systematic experiments have not been performed in other mammals, studies in
sheep indicate that preventing nursing but not licking, by placing the newborn lamb in a wire mesh
cage with its lid open for either 4 or 12 hours, has little consequence on maternal behavior even in
primiparae (Otal et al., 2009; Poindron and Le Neindre, 1980). Moreover, there is no indication that
preventing only nursing impairs the recognition of one’s own lamb in primiparous or multiparous
mothers (Otal et al., 2009). By contrast, deprivation of all physical contact with the newborn lamb
has a drastic impact on the development of maternal responsiveness and selectivity in both sheep
(Otal et al., 2009) and goats (Bordi et al., 1994; Romeyer et al., 1993). Whether these effects are
caused by the lack of perioral stimulation, or by the absence of ingesting amniotic fluids and the
impairment of some perception of olfactory cues from the young, is unknown. The latter possibility
is likely, given the important role of olfaction in maternal behavior of these ungulates.
Auditory and Visual Cues

Although somatosensory (touch), thermal (temperature), and olfactory (odor) cues are probably most
important for the regulation of maternal behavior, other cues may also contribute to the proximal
control of behavior and may be influenced by hormones. For instance, ultrasonic calls, above the
range of human hearing, emitted by pups when they are in distress, cold, or out of the nest result in
the mother’s locating them from a distance and retrieving them back into the nest (Allin and Banks,
1972; Brewster and Leon, 1980); such directional orientation to pup ultrasounds is facilitated by
associated pup odor cues (Farrell and Alberts, 2002; Smotherman, Bell, Starzec, Elias, and Zachman,
1974). In fact, Brouette-Lahlou, Godinot, and Vernet-Maury (1999) reported that pup ultrasounds
stimulate the initiation of maternal anogenital licking of pups, which is then facilitated or patterned
by pup preputial secretions acting on the maternal vomeronasal system (Brouette-Lahlou et al., 1999;
Brouette-Lahlou, Vernet-Maury, Godinot, and Chanel, 1992). That ultrasonic calls may acquire
motivational properties is suggested by the observation that virgin animals do not awaken in response
to these calls, whereas postpartum mothers do. The brain neuropeptide oxytocin plays an important
role in the mediation of pup retrieval behavior in female mice acting by enhancing auditory cortical
pup ultrasonic distress calls (Marlin, Mitre, D’Amour, Chao, and Froemke, 2015). They found that
pup retrieval behavior was accelerated by oxytocin in the left auditory cortex, and oxytocin receptors
were preferentially expressed in the left auditory cortex.
Visual cues may also contribute to maternal responsiveness, although their role must be restricted
to proximal interactions and distances over which the dam can see (newborn pups are more effective
at eliciting retrieval in the newly parturient dam than are older pups; Peters and Kristal, 1983; Stern,
1985). That said, neither sight nor hearing is necessary for the expression of maternal behavior. In
the absence of both, maternally experienced dams show normal interactions with pups (Beach and
Jaynes, 1956a, 1956b; Herrenkohl and Rosenberg, 1972).
Taken together, hormones reduce pup-avoidance, augment the dam’s attraction to pup-related
odorants and ultrasonic vocalizations, and sensitize the mother to tactile cues that promote a rapid
onset of maternal behavior at parturition.
40
Experiential Effects on the Maintenance and Retention

of Maternal Behaviors
Processes regulating the long-term maintenance of maternal behavior differ from those involved
in its onset. The female first undergoes a transition period during which hormones interact with
environmental and experiential processes in the regulation of behavior. However, once this transition
period is over, by the end of the first postpartum week, behavior is maintained primarily by sensory
influences and processes of learning and reinforcement. Hormonal effects on maternal behavior at
this time are minimal (Lonstein et al., 2015).
Behavioral Changes From Birth to Weaning

Mothers respond to their offspring for a considerable period after parturition, although the quality
of responsiveness changes as the young grow and mature. In general, young are weaned at about 20
to 25 days, and although mothers continue nursing young over this period, nursing bouts become
shorter, inter-bout intervals become longer, and dams spend increasing time away from their young
(Leon, Croskerry, and Smith, 1978). In fact, once the young are mobile, by 12 to 15 days of age,
the mother increasingly distances herself from the pups; she rarely retrieves them, her nest becomes
matted, and she terminates nursing bouts before the infants have had their fill (Fleming and Blass,
1994). By 15 to 20 days of age, pups begin to supplement their diet with solid food, which they first
encounter in the mother’s milk, then as particles of food in the mother’s saliva or on the mothers
body (mouth, head, and fur), and then when they follow their mother to the food source (Alberts
and Cramer, 1988; Galef, 1989; Galef and Beck, 1990).
At a more proximal level, after the first few postpartum days, mothers and litter develop a rhythm of
interaction in which the dam alternates between being in the nest and nursing the young and leaving
the nest, out of litter contact. Leon and his colleagues (Leon et al., 1990) showed that the duration of
the long nursing nest bouts is regulated by an interaction of hormonal factors and the thermal charac-
teristics of the nest, the huddle of pups, and the mother. Dams experience a rise in body temperature
when nursing and get off the pups when they experience acute hyperthermia (Leon et al., 1978). In
comparison to nonlactating animals, dams have a chronically elevated core temperature, making them
more vulnerable to hyperthermia ( Jans and Leon, 1983a, 1983b). Moreover, endocrine changes associ-
ated with suckling and lactation contribute to the elevated core body temperature. The developing pups
induce their mothers to release both prolactin and ACTH, which provoke the release of progesterone
and corticosterone, respectively. Progesterone then elevates the maternal thermal set point, and corticos-
terone increases maternal heat production and possibly heat retention. The resulting chronic increase in
maternal heat load makes mothers vulnerable to a further acute increase in their heat load, eventually
driving up maternal brain temperature and forcing the interruption of pup contact (Leon et al., 1990).
Although many hormones are associated with lactation, the primary ones are the peptides, pro-
lactin, adrenocortical hormone, and oxytocin as well as adrenal steroids but not the “parturitional”
hormones known to be associated with the onset of responsiveness. Although both prolactin and
oxytocin have been implicated in the onset of maternal responsiveness (Bridges, 1990; Insel, 1990),
there is no evidence that these “lactational” hormones contribute directly to the dam’s motivation to
continue to respond nurturantly during the lactational period or are directly involved in the long-
term effects of experience seen in differences between first time mothers and multiparous mothers.
Parity and Effects of Postpartum Experiences

The importance of postpartum experience is reflected both in the short term, during the first post-
partum week, and in the long-term maintenance of behavior of the primiparous female and across
41
Aya Dudin et al.
subsequent births. The latter, expressed as the parity effect, is extensively and thoroughly described
by Bridges (2016b) who describes the multiple behavioral and physiological differences between
primiparous and multiparous mothers. Bridges (2016b) reported for instance that in the rat with a
regular estrous cycle, levels of prolactin, estrogen, and corticosterone are lower in multiparous than
in primiparous females and some of these differences persist also as a function of prior parity in the
pregnant animal (Bridges, 2016b). Hence, with experience, lower levels of hormones are necessary to
produce the same effects; that is, multiparous animals are more sensitive to the effects on brain and/or
behavior of hormones than are primiparous animals. Consistent with this idea, multiparous animals
are more responsive to pup stimulation than primiparous animals in terms of their dopamine release
into the nucleus accumbens (NA) with presentation of pups (Afonso et al., 2008). Parity-induced
differences in the sensitivity of receptor activation have also been reported—some consistent with
the idea that multiparous mothers are more sensitive, and some counter to this conceptualization
(Bridges, 2016b).
In terms of the nature of the relevant experience associated with parity effects, considerable work
has focused on the initial few postpartum days when new mothers first encounter their young and
interact with them. This interaction is laid down in memory and becomes rapidly consolidated
(Bridges, 2016b; Lonstein et al., 2015).
Once animals initiate responsiveness to young at parturition, whether as first time mothers or
multiparous mothers, the continued expression of maternal behavior after 4 to 5 days post-parturition
seems no longer to be based on hormones but is, instead, based both on experiences acquired by the
mother when she interacts with pups under the influence of hormones and on experiences acquired
during the lactational period (Bridges, 2016b; Fleming, Morgan, and Walsh, 1996). Thus, processes
of learning and memory sustain the behavior beyond the period of hormonal priming and into the
next parity (Li and Fleming, 2003a).
If pups are removed from newly parturient (or cesearan-delivered) females before dams have
had the opportunity to interact with the pups, maternal responsiveness declines over the next 3 to
5 days and reaches low virgin levels by day 10, by which time animals have usually reinitiated their
estrous cycles (Orpen, Furman, Wong, and Fleming, 1987). However, if females give birth (or are
C-sectioned) and within 24 to 36 hours are permitted to interact with pups for as little as a half
hour before separation, dams continue to be maternal in tests undertaken 10 days later (Orpen and
Fleming, 1987). Not surprisingly, a longer interactive exposure period results in a longer retention
of responsiveness (Bridges, 1975, 1977; Cohen and Bridges, 1981; Fleming and Sarker, 1990). This
long-term change in behavior as a result of experience interacting with pups has come to be known
as the “maternal experience effect” or “maternal memory” and has now been demonstrated in other
species (e.g., rabbit; González-Mariscal et al., 1998).
Parturitional hormones influence the robustness of maternal learning and memory. Animals
which acquire maternal experience under the influence of the parturitional hormones (whether
postpartum animals or virgins treated with hormones) exhibit better retention of maternal behavior
30 days later than do animals who are not being stimulated with hormones at the time of the mater-
nal experience (virgins or nonparous but experienced animals). Moreover, the optimal condition
for the expression of maternal behavior occurs when both the initial experience and the test occur
during a period of hormonal priming (Fleming and Sarker, 1990). There is considerable evidence
that multiparous animals (which have experienced a previous pregnancy, parturition, and period of
pup-rearing) are less disturbed than primiparous mothers by a variety of experimental manipulations
including C-section (Moltz, Robbins, and Parks, 1966), endocrine manipulations (Moltz, Levin,
and Leon, 1969; Moltz and Wiener, 1966), morphine administration (Bridges, 1990; Kinsley and
Bridges, 1988), and brain lesions (Fleming and Rosenblatt, 1974b; Franz, Leo, Steuer, and Kristal,
1986; Numan, 1994; Schlein, Zarrow, Cohen, Denenberg, and Johnson, 1972) that could disrupt
maternal behavior. In addition, pup cues, which are initially ineffective in eliciting maternal behavior
42
in first-time mothers, come to be effective in multiparous animals (Noirot, 1972). Finally, among
ewes, exogenous hormones are most effective in enhancing responsiveness in experienced animals
(Poindron and Le Neindre, 1980).
Although most work in this area has focused on mother’s learning about her offspring, this
enhanced learning ability seems not to be specific to the maternal context; in comparison to virgins
or nonmothers, new mothers during the postpartum period also show enhanced learning in other
contexts, involving other forms of social learning (Fleming, Kuchera, et al., 1994), as well as spatial
learning and hunting (Bodensteiner, Cain, Ray, and Hamula, 2006; Gatewood et al., 2005; Kinsley
et al., 2014; Kinsley and Lambert, 2008; Kinsley et al., 1999; Lemaire et al., 2006; Love et al., 2005;
Pawluski, Vanderbyl, Ragan, and Galea, 2006; Pawluski, Walker, and Galea, 2006; for opposite results
see Bodensteiner et al., 2006; Darnaudéry et al., 2007).
Early postpartum learning also occurs in sheep, and this learning maintains their maternal respon-
siveness beyond the peri-parturitional period (Lévy and Keller, 2008). However, maternal respon-
siveness fades rapidly. A 36- to 72-hour separation period that follows 4 hours of contact after
parturition induces rejection of the familiar lamb (Keller, Meurisse, and Lévy, 2005; Lévy et al., 1991).
This decline in maternal responsiveness cannot be compensated for by increasing initial mother-
young because the rejection is also observed when the separation is performed after a week of
postpartum interaction (Keller et al., 2005). Maternal selectivity can, however, be strengthened over
time. Although selective mothers exposed to the lamb for 4 hours just after birth are not able to
retain selectivity after 24 or 36 hours of separation, memory of the lamb is maintained if ewes and
their lambs have been in contact a week (Keller et al., 2005; Lévy et al., 1991). Hence, offspring
recognition memory is labile and has a short duration during the initial postpartum period, whereas
maternal selectivity strengthens over time, suggesting the involvement of consolidation processes for
emergence of the latter.
Taken together, these studies indicate that experiences acquired under hormones are also more
easily activated by a combination of hormones and exposure to relevant pup stimuli in the absence
of hormones. There are a number of ways hormones could act to promote these robust experience
effects. They could increase the salience of associative cues, most likely unconditioned pup cues
(e.g., proximal tactile or olfactory) during the learning phase; they could act to facilitate or strengthen
the association between the conditioned and unconditioned pup-associated cues; finally, they could
produce internal cues that themselves act as conditioned stimuli, a mechanism that could explain the
apparent state dependency of the maternal-hormone interactions described above. Research has not
yet identified which of these hormone mechanisms are most important.
Sensory Mechanisms Involved in Maternal Experience

Although it is clear that interactive experience is important for the long-term retention of behavior,
which aspect of the experience is important is not known. In this section, we consider different
sensory experiences the animal acquires while interacting with pups. As will become apparent, both
somatosensory and chemosensory inputs are important for the long-term retention of maternal
behavior.
During early interactions with pups, the dam is multiply stimulated by distal visual and auditory
cues and by more proximal tactual, chemosensory and, possibly, thermal cues, and these may well be
important aspects of the maternal experience (Stern, 1989). The significance of somatosensory and
chemosensory stimulation for the maintenance of maternal responding is well established; if the mother
is prevented from crouching over her young during the postpartum period, but receives other distal
inputs, her responsiveness declines more rapidly with earlier weaning ( Jakubowski and Terkel, 1986;
Stern, 1983). Moreover, Orpen and Fleming (1987) found that, if mothers were separated from their
litters by a wire mesh floor during the 1-hour postpartum exposure phase, so that they could see, hear,
43
Aya Dudin et al.
and distally smell pups, but received no tactile or proximal chemosensory input, on tests 10 days later
they showed no long-term benefit of maternal experience, but instead responded to pups as virgins do.
These data indicate that ventral stimulation is probably an essential feature of the maternal experience.
The additional findings that dams need to receive somatosensory perioral input from the mouth region
to exhibit normal maternal licking and crouching (Stern and Johnson, 1989; Stern and Kolunie, 1989),
and that licking during exposure is correlated with responsiveness during test (Morgan et al., 1992),
point also to the importance of chemosensory and somatosensory stimulation for the maternal expe-
rience effect (Morgan et al., 1992). However, olfactory learning in this context is clearly paramount.
During interactions with pups, dams learn about specific olfactory and chemosensory features
of the pups. If pups are scented with an artificial odorant during the exposure phase, on test 10 days
later dams respond more rapidly to pups labeled with the same scent than to those labeled with a
discrepant scent. That this effect depends on the association of the odors with the pups is shown by
the additional observation that preexposure to the odor on its own (in the absence of pups) does
not result in the same facilitation of responsiveness to similarly scented pups (Malenfant, Barry, and
Fleming, 1991). Moreover, how dams respond to pup-associated scents depends on the quality of
their interactions with pups during the initial pup-odor pairings. If mothers interact proximally with
pups during the pairing, and spend time sniffing and licking them, they develop a strong long-term
preference for that scent over a novel scent; however, if during the pairing mothers do not respond
maternally, and remain at a distance from pups, then the dams do not develop a preference for the
paired scent (Fleming, unpublished observation). Although pup-associated scents can be learned,
olfactory input is apparently not necessary for a maternal experience. Olfactory and accessory olfac-
tory denervations prior to an experience does not block the long-term influence of pup-associated
odors (Fleming et al., 1992; Mayer and Rosenblatt, 1977).
Long-Term Effects of Maternal Experience and Parity on Maternal Affect

Parity differences in anxiety. Postpartum and virgin animals differ in the number of respects, one dif-
ference is in the effects of a maternal experience on anxiety levels. Postpartum rats are often less
anxious than virgin females, as indicated by group differences in the elevated plus maze test (Lon-
stein, 2007a; Silva, Bernardi, Nasello, and Felicio, 1997a), the open field test, and light-dark box test
(Fleming et al., 1989a; Miller, Piasecki, and Lonstein, 2011). This reduced anxiety in dams persists
well into later adulthood (Bridges, 2016a; Byrnes and Bridges, 2006), as nonlactating primiparous rats
exhibit reduced anxiety levels on the elevated plus maze and open field test than their age-matched
nulliparous controls tested during the proestrus phase. Estrogens, possible estrogen receptor alpha
(ERα), may play a role in this parity-induced shift in anxiety (Byrnes, Casey, and Bridges, 2012).
Treatment of nonlactating, primiparous females with an Erα agonist increased time spent in the
open arms (Agrati and Lonstein, 2016b), and altered the “stress” peptide, corticotropin releasing hor-
mone (CRH) mRNA expression in both hypothalamus and amygdala in a parity-dependent fashion.
These findings indicate that reproductive experience can alter the Erα-mediated limbic systems that
mediate emotional responses.
Parity differences in pup reinforcement. Despite these parity differences in general olfactory learning
and emotion regulation, the primary reason the postpartum animal expresses such a robust maternal
experience is because rat pups are highly reinforcing for the maternal animal (Fleming, Korsmit,
and Deller, 1994a; Wansaw, Pereira, and Morrell, 2008). In a series of studies addressing this issue,
Fleming and colleagues (Fleming, Korsmit, et al., 1994; Lee, Clancy, and Fleming, 2000) compared
postpartum and virgin animals under a variety of different temporal, hormonal, experiential, depri-
vation, and stimulus conditions on a conditioned place preference (CPP) paradigm or in an operant
box using either rat pups or food as the reinforcing stimulus. CPP tests showed that dams spend
more time in the pup-associated side of the CPP apparatus than the food associated side, whereas
44
virgins spend more time in the food associated side than the pup-associated side. Hence, pups are
more reinforcing than food to postpartum animals, whereas food is more reinforcing than pups to
the virgins. Subsequent work by Morrell and her colleagues shows that dams in early (days 4–8)
postpartum period find pups more reinforcing than those in late (days 12–16) postpartum (Wansaw
et al., 2008), and young pups are even more reinforcing than cocaine for early postpartum females
(Mattson, Williams, Rosenblatt, and Morrell, 2001). When virgins are induced to become maternal
as a result of extensive experience with pups, pups take on heightened reinforcing properties in the
absence of the parturitional hormones. The fact that pups are more reinforcing to the maternal
postpartum animal than to the “maternal” virgin suggests, however, that hormones augment the sali-
ence of the pup stimulus ( Fleming, Korsmit, et al., 1994). Consistent with this interpretation, in the
virgin parturitional hormones enhance the reinforcing effects of pups, but only if the hormones also
activate the expression of maternal behavior in virgins. Hormones had no effect on the reinforcing
properties of food stimuli. In a second test of pup-reinforcement, Lee et al. (2000) found that during
pregnancy females will not bar-press for pups although they will bar-press for food, but that after the
birth of the litter, bar-pressing for pups increases tenfold. Again hormones augment this effect but are
not necessary for it, because animals commence bar-pressing outside the postpartum period, as soon
as they begin showing maternal responses in the home cage. That pups are the relevant reinforcing
stimulus, maintaining responses is indicated by the observations that, if pups are removed, the bar-
press response extinguishes almost immediately.
These studies indicate that for animals to respond maternally to pups during the initial encoun-
ters, pups do not have to be strongly reinforcing. Presumably the attraction to pups induced by
hormones is adequate to insure the mother will respond nurturantly. However, for maternal respon-
siveness to be sustained in the absence of hormones, females must gain experience interacting with
pups, which in general occurs in the presence of hormones; as a result of this experience pups acquire
strongly reinforcing properties.
Reinforcing characteristics of the mother-litter interaction. To determine precisely what aspects or features
of pups are reinforcing to the maternal animal, Magnusson and Fleming (1995) tested the relative
contributions of chemosensory and somatosensory stimulation during maternal interactions on the
development of a conditioned place preference. They found that pups do not acquire reinforcing prop-
erties as readily if dams are exposed to pups placed in a Plexiglas cube, permitting visual, auditory, and
olfactory stimulation, but preventing the proximal somatosensory and chemosensory input normally
associated with mouthing, licking, retrieving, and crouching. Thus, for pups to be reinforcing they must
provide proximal stimulation to the dam, and especially of her ventral and mouth regions. Finally, the
reinforcing properties of pups also depend on their ages and odor characteristics. Younger pups are
generally more reinforcing than older ones (Wansaw et al., 2008), possibly because they provide more
sensory stimulation to dams. Pups will not sustain a conditioned place preference if dams are rendered
anosmic. Thus, both tactile and olfactory pup input contribute to the reinforcing properties of pups.
Taken together, these studies show that maternal learning is a robust phenomenon that is based
on activation of both chemosensory and somatosensory systems during mother-litter interactions.
Although the new mother seems primed to respond to certain cues over others by the action of hor-
mones, the primary effect of the maternal hormones is to activate maternal behavior. Once maternal
behavior has been exhibited, general mechanisms of learning and memory are utilized to further
consolidate experiences acquired during mother-litter interactions. These experiences include the
activation in the mother of both chemosensory and somatosensory systems.
Neuroanatomy of Maternal Behavior

Although there is substantial evidence that some of the parturitional hormones exert their effects on
maternal responsiveness by acting on neural substrates in the brain, which hormones exert central
45
Aya Dudin et al.
effects, what brain systems are implicated, and by what specific behavioral mechanisms, are not totally
understood. Of interest also is whether the systems that mediate the onset of maternal behavior and
the maternal experience effect are the same, different, or overlapping.
As can be seen in Figure 2.1, the neural systems that are most important include the olfactory
systems, the limbic system, and the hypothalamus (Numan, 1994; Numan and Sheehan, 1997). The
olfactory system mediates the sense of smell and is composed of two parts: The main olfactory sys-
tem detects odor molecules in the air that activate the olfactory receptors in the nose when an animal
sniffs an object, and the accessory olfactory system detects molecules in a liquid medium that activate
receptors in the vomeronasal organ, also situated in the nose, when an animal touches an object with
its snout. The limbic system involves groups of neurons and their axons which have been implicated in
the regulation of emotional behavior, reward, and memory processes, as well as species-characteristic
behaviors. Brain regions included in this system are the hypothalamus, the amygdala, the nucleus
accumbens (NA), the hippocampus and medial prefrontal cortex. The hypothalamus sits at the base
of the brain above the pituitary gland, consists of different groups of cells involved in the control of
a variety of reproductive behaviors and the release of hormones such as oxytocin and vasopressin
from the pituitary gland, and is responsive through specialized receptors to hormones that activate
maternal behavior. Two important nuclear groups within this region are the medial preoptic area
(MPOA), which is situated somewhat anterior to the hypothalamus and the ventral part of the bed
nucleus of the stria terminalis (vBNST), which sits adjacent and dorsal to the MPOA. Also within
the hypothalamus are the ventromedial hypothalamic nucleus (VMH) and paraventricular nucleus
(PVN), which are positioned close to the midline, but posterior to the MPOA, closer to the pitui-
tary. All four hypothalamic structures respond to environmental stimuli and to circulating hormones.
These different brain areas are interconnected by a series of neural pathways. For instance, the two
olfactory systems (main and accessory) have direct connections with the limbic system (especially
the amygdala) and the hypothalamus by means of the lateral olfactory tract. The amygdala, in turn, is
interconnected with the nucleus accumbens, and both are connected with the hypothalamus.
Prior to Numan (1974), little was known about the neuroanatomy of maternal behavior. The early
work by Beach (1937) focused on neocortical structures and suggested no one area of the cortex is
crucial to the expression of maternal behavior. However, the greater the cortical mass removed, the
greater the deficits in behavior. Subsequent to these early studies on neocortex, other studies focused
on the midline cortex and associated limbic structures, the hippocampus, and septum (Fleischer
and Slotnick, 1978; Slotnick, 1967; Stamm, 1955; Terlecki and Sainsbury, 1978; Wilsoncroft, 1963).
Although small lesions of these regions disrupt maternal behavior, the deficits were primarily related
to motor sequencing and patterning, not to maternal motivation. Thus, animals with these forebrain
lesions continued to respond to pups but in a disorganized fashion (Slotnick, 1967; Stamm, 1955).
Maternal “Core”: Two Antagonistic Neural Systems

The neural bases involved in the regulation of maternal behavior expression at the time of partu-
rition are well understood. There are at least two basic antagonistic neural systems governing the
expression of maternal behavior—referred to as the maternal core. One is the excitatory neural
system that deals with the activation of maternal responses towards pups; the other is an inhibitory
neural system that may regulate avoidance and aversive responses to pups or pup-related stimuli. The
balance between these two systems determines whether maternal behavior will be expressed at the
time of parturition. The combination of sensory cues, parturitional hormones, and experiential fac-
tors exerts effects on the excitatory system to bring animals close to pups by increasing the attractive
quality of pups or pup-related stimuli, and initiate and maintain maternal care; these factors exert
effects on the inhibitory system to inhibit animals’ naturally fearful responses toward novel pups
(Rosenblatt, 1990; Schneirla, 1959).
46
The excitatory system. The excitatory system is controlled primarily by neurons in the medial
preoptic area (MPOA) and ventral part of the bed nucleus of the stria terminalis (vBNST) and
their efferent projections to the brainstem, such as the ventral tegmental area (VTA) (Numan, 1988,
1994; Numan and Sheehan, 1997). Lesions of MPOA/vBNST cell bodies, or knife cuts transect-
ing their lateral projections completely abolish maternal behavior (Numan, Corodimas, Numan,
Factor, and Piers, 1988; Numan, McSparren, and Numan, 1990; Numan and Numan, 1996) in the
new mother or maternal virgin, whereas “kindling-like” electrical stimulation of this site facilitates
maternal responding (Morgan, Watchus, Milgram, and Fleming, 1999a). Hormones that activate
maternal behavior act on receptors in the MPOA; implants into MPOA of either estradiol, prolactin,
or oxytocin (albeit under somewhat different conditions) facilitate maternal responding (Bridges,
Numan, Ronsheim, Mann, and Lupini, 1990; Insel, 1990; Numan, Rosenblatt, and Komisaruk, 1977),
whereas the infusion of oxytocin antagonists or morphine or β-endorphin in the MPOA impair
maternal behavior (Mann and Bridges, 1992; Pedersen, Caldwell, Walker, Ayers, and Mason, 1994;
Rubin and Bridges, 1984). Implants of antiestrogen in the MPOA also delay the onset of maternal
behavior (Ahdieh, Mayer, and Rosenblatt, 1987; Numan and Insel, 2003; Numan and Young, 2016).
Another line of evidence supporting the importance of MPOA/vBNST neurons in the control
of maternal expression comes from studies using c-fos immunohistochemistry. The proto-oncogene
c-fos is one of a class of genes (known as immediate early genes) which are expressed in response to
a variety of stimulus conditions (Sagar, Sharp, and Curran, 1988) by producing a protein called Fos
protein. Fos expression is often used as a marker for detection of neuronal activation. Several stud-
ies have demonstrated that a population of neurons in the MPOA/vBNST is involved in regulating
maternal responsiveness independent of sensory input. For example, Fleming, Suh, Korsmit, and
Rusak (1994) found that postpartum rats exposed to pups had higher numbers of cells showing Fos
within the MPOA nuclei than did those exposed to adult conspecifics or left alone. Numan and
Numan (1994) also found postpartum rats exposed to pups had more Fos-labeled neurons in the
MPOA and vBNST than did postpartum control females exposed to candy. These effects require
that the animal is actively maternal and crouches over pups; however, they do not depend on activa-
tion by many of the pup-associated sensory inputs. Heightened MPOA c-fos expression persists in
maternally active animals even after animals are rendered anosmic (unable to smell), anaptic (unable
to feel touch sensations around the muzzle) or after temporary anaesthetization of the ventral nipple
area (Numan and Numan, 1995; Walsh, Fleming, Lee, and Magnusson, 1996).
To understand the function of the MPOA/vBNST in maternal behavior, Numan and col-
leagues investigated the MPOA/vBNST projections implicated in maternal responding (see Numan
and Sheehan, 1997). By combining the c-fos immunohistochemistry technique with the neural
tract-tracing technique, Numan and Numan (1997) found that “maternal” neurons (visualized by
Fos-labeling) in the MPOA mainly project to the lateral septum (LS), ventromedial nucleus of
the hypothalamus (VMH), and the periaqueductal gray (PAG), whereas “maternal” neurons in the
vBNST project to the retrorubral field, PAG, and VTA. The activation of these projections in the
control of maternal behavior is consistent with the involvement of these regions in maternal behav-
ior. For instance, the LS has been implicated in the control of the sequential organization of the
maternal pattern (Fleischer and Slotnick, 1978); the VTA has been linked to the motivational aspect
of maternal behavior (Hansen, Harthon, Wallin, Löfberg, and Svensson, 1991b); the VMH is involved
in the control of aversive reactions toward pups (Bridges, Mann, and Coppeta, 1999; Sheehan and
Numan, 1997); and the PAG specifically for the regulation of the upright nursing posture and mater-
nal aggression (Lonstein and De Vries, 1999; Lonstein, Simmons, and Stern, 1998). These projections
may contribute to different aspects of maternal behavior control. As stated by Numan and Sheehan
(1997, p. 105),“the hormonally primed preoptic area projects to some regions to facilitate the appeti-
tive aspects of maternal behavior, projects to other regions to potentiate consummatory components,
and projects to still other neuronal groups to depress aversive reactions to pup stimuli”.
47
Aya Dudin et al.
The inhibitory systems. The MPOA/vBNST not only projects to the midbrain and motor system
involved in the expression of maternal behavior, these nuclear groups also receive input from other
parts of the brain, in particular the olfactory and limbic systems (see Figure 2.1), which exert inhibi-
tory influences on the functions of the MPOA/vBNST (Fleming, 1987; Numan, 1988). For instance,
a major input to the MPOA comes from the amygdala which in turn receives input from the main
and accessory olfactory bulbs (Fleming, 1987). Thus, removal of main or accessory olfactory input
facilitates the expression of the maternal complex in nonresponsive virgin animals, while at the same
time reducing certain olfactory-mediated components (like licking) in both virgins and postpartum
animals who are maximally responsive. These data are consistent with the observation that pup odors
within the context of other pup cues are aversive to virgin animals but attractive to the postpartum
dams (Fleming et al., 1989b; Fleming and Rosenblatt, 1974b, 1974c; Fleming et al., 1979). The find-
ings that infusions of oxytocin into the olfactory bulb facilitate the expression of maternal behavior,
whereas infusions of oxytocin antagonist markedly delay all components of maternal behavior sug-
gest that the olfactory bulb is one such site where parturitional hormones act to antagonize the
inhibitory control on maternal behavior (Yu et al., 1996). Behavioral inhibition is also exerted by sites
that receive chemosensory projections and that project to the MPOA, such as the medial and cortical
nuclei of the amygdala and the VMH, and the anterior hypothalamic nucleus (AHN). The amygdala
receives inputs from both olfactory systems and AHN/VMH and projects directly to the MPOA
and the AHN/VMH (Canteras, Simerly, and Swanson, 1995). Activation of the olfactory systems
increases medial amygdala neuronal activity, whereas electrical stimulation of the medial amygdala
predominantly inhibits MPOA neurons as well as the onset of maternal behavior (Gardner and Phil-
lips, 1977; Morgan et al., 1999a). Lesions of the medial amygdala, the stria terminalis (the major effer-
ent pathway from the medial amygdala), the BNST, or the AHN/VMH all result in the disinhibition
of maternal retrieving and crouching in virgin animals exposed to foster pups (Bridges et al., 1999;
Fleming, 1987; Fleming et al., 1992; Fleming, Vaccarino, and Luebke, 1980; Fleming et al., 1979;
Numan, Numan, and English, 1993). The facilitation of maternal behavior produced by amygdala
lesions is abolished by lesions of the MPOA, confirming that the input from amygdala acts through
MPOA to exert its inhibitory role on maternal behavior (Fleming, Miceli, and Moretto, 1983).
Komisaruk and colleagues (2000) provided support for the notion of an active inhibitory system
in the regulation of maternal behavior. They combined 14C-2-deoxyglucose (2-DG) autoradio-
graphic method with c-fos immunocytochemistry to visualize the neural activities in specific brain
regions under different maternal conditions. The 2-DG method provides information about the
metabolic activity of neuronal input. By contrast, c-fos immunocytochemistry detects post-synaptic
neuronal activity, therefore it indicates the metabolic activity level of neuronal output. Information
from the combined methods can reveal input-output relations in certain brain areas. They found that
in parturient and hormonally primed maternal animals, there were elevated 2-DG and c-fos activi-
ties in the MPOA and in sites that receive accessory olfactory bulb input (e.g., medial amygdala),
indicating an increase in the input and output activity of these areas. In contrast, maternal virgin
animals showed a decrease of 2-DG activity but an increase of c-fos activity in the medial amygdala
(ME), indicating a decreased input but increased output activity. These results suggest that for the
virgin animals to become maternal through pup-induction, the input activity in the vomeronasal
nuclei must be decreased, which in turn, disinhibits the output neurons to stimulate the neurons in
the MPOA and, in so doing, activates the whole excitatory system.
The amygdala, especially the medial part, has an inhibitory influence on the onset of maternal
behavior. The inhibitory circuits consist of olfactory systems-to-amygdala-to-MPOA/vBNST and
olfactory systems-to-amygdala-to-AHN/VMH-to-MPOA/vBNST (also the downstream periaq-
ueductal grey, PAG; Numan, 2015). Because it has been demonstrated that virgins sustaining amyg-
dala lesions differ from controls in not withdrawing from pups and in maintaining closer proximity
to them, and they are less fearful in a series of emotionality tests (Fleming et al., 1980), it is proposed
48
that these circuits “depress[es] maternal behavior by activating a central aversion system” (Numan
and Sheehan, 1997). These effects of amygdala lesions on the animal’s affect are consistent with an
extensive literature relating the amygdala to emotional behavior within other contexts (Davis, 1992;
Everitt and Robbins, 1992; LeDoux, 1992). One interesting feature of these olfactory limbic struc-
tures is that a number of these sites contain receptors for estradiol (Pfaff and Keiner, 1973), proges-
terone (Numan et al., 1999), and oxytocin (Brinton, Wamsley, Gee, Wan, and Yamamura, 1984), as
well as opiates (Bridges, 1990; Hammer, 1984), and therefore constitute likely sites for the hormonal
and neurochemical alteration of maternal affect. However, no studies to date have been published
showing that maternal hormones or neurotransmitters act on these limbic sites to influence maternal
affect in general or maternal behavior specifically.
Maternal behavior is under the joint control by two antagonistic neural systems: the excita-
tory system, which mainly consists of the efferents from MPOA/vBNST neurons to various brain
areas, and the inhibitory system, which mainly refers to the projections from medial amygdala to
the MPOA/vBNST and VMH. These systems may coordinately regulate neuroendocrine, sensory,
and autonomic components necessary for the elaboration of maternal behavior. This circuitry is the
“core” system mediating the expression of maternal behavior. They are likely modulated by other
higher cortical regions, such as the auditory cortex (Afonso et al., 2007; Febo et al., 2010; Marlin
et al., 2015; Pereira and Ferreira, 2016; Wu et al., 2016), possibly via its projections to the “core”
system of maternal behavior. This core system also does not incorporate other brain structures
known to be involved in maternal behavior, such as the lateral habenula (LH; Corodimas, Rosenblatt,
Canfield, and Morrell, 1993; Corodimas, Rosenblatt, and Morrell, 1992; Felton, Linton, Rosenblatt,
and Morrell, 1999; Matthews-Felton, Corodimas, Rosenblatt, and Morrell, 1995), and the nucleus
accumbens (Li and Fleming, 2003a, 2003b).
Neurochemistry of Maternal Behavior
The Mesolimbic and Mesocortical Dopamine Systems

and Maternal Motivation
Maternal behavior is a robust, goal-directed, motivated behavior, and pups are reinforcing stimuli
for the maternal animal (Fleming, Korsmit, et al., 1994; Fleming, Suh, et al., 1994; Lee et al., 2000;
Nissen, 1930; Stern and Mackinnon, 1976; Wilsoncroft, 1968). Most studies on maternal motivation
have focused on pup retrieval, because it is the most dramatic behavior and is very easy to quantify.
By looking at pup retrieval latency and the number of pups retrieved during a certain test period,
one can assess an animal’s interest in pups and the intensity of her “motivation” to be in proximity
to them. Besides the “core” MPOA/vBNST excitatory neural system in the regulation of specific
maternal motivation, the other important system in the regulation of maternal “motivation” is the
mesocorticolimbic dopamine system, which originates in the midbrain VTA and releases the neu-
rotransmitter, dopamine, in the nucleus accumbens and the medial prefrontal cortex (mPFC). The
current view is that the mesolimbic and mesocortical dopamine systems are part of a nonspecific or
general motivational system which serves to increase an organism’s responsiveness to a wide variety
of biologically significant stimuli, including pups (Numan, 2007; Numan and Young, 2016; Olazábal
et al., 2013a, 2013b).
In Vivo Dopamine and Enhanced Maternal Motivation

Hansen et al. (1993) were the first to demonstrate increased dopaminergic activity in the ventral
striatum (nucleus accumbens) of mother rats when they were reunited with their newborns after an
overnight separation. Building on the early work of Hansen et al. (1993), Fleming and her colleagues
49
Aya Dudin et al.
undertook a series of studies of the role of extracellular dopamine in the nucleus accumbens. In these
studies the authors provided evidence for the relation between DA and (1) hormonal profiles associ-
ated with late pregnancy, parturition, and parity and (2) salience of pup stimulation.
Dopamine and hormones. When provided with rat pups, postpartum females show large and sus-
tained elevations in DA in the NA. These elevations occur against a background of a hormonally
mediated suppression in basal tonic DA release (Afonso et al., 2009). In the absence of any prior pup-
experience, ovariectomized rats treated with progesterone and estrogen (which induces maternal
responsiveness to foster pups) display a similar reduction in basal DA responses as postpartum intact
mothers and a similar initial increase in response to pup presentation (Afonso et al., 2009). The more
rapidly a rat becomes maternal under the influence of hormones, the lower basal DA prior to pup
exposure. Thus, hormones that facilitate maternal responsiveness in the absence of previous maternal
experience have the same effects on NA DA functioning as in the intact postpartum experienced
female, and this hormone-induced basal suppression is related to heightened maternal responsiveness.
It can be argued that basal DA suppression aids the rapid expression of maternal behavior through
a reduction in the DA noise in the absence of other stimuli. In this way, it might serve as a mecha-
nism for sharpening the DA signal in response to pup stimuli and augmenting the pup saliency to
develops. Impairments to basal suppression (e.g., by manipulations of the early environment; Afonso
et al., 2011) would be expected to result in a consequent DA signal decrease and a reduction in pup
salience, culminating in impaired mother-pup interaction.
Dopamine and stimulus specificity. Does this signal-to-noise mechanism result in sharpened DA sig-
nals to all stimuli, or only to pup stimuli, in postpartum females? In subsequent studies Afonso et al.
(2009) compared the behavioral and DA profiles of female rats when exposed to pups and to palat-
able food (sweet cereal). Both nulliparous and parous females showed increased DA responses when
ingesting the sweet treat. However, only postpartum females showed pup-evoked DA responses
greater than the food-evoked DA responses; a finding not observed in cycling females even after
pup-experience. The enhanced DA responsiveness to pups in postpartum females is sustained even
when the dams are exposed to pups in a perforated box at a distance and cannot interact with them
(Afonso et al., 2013).
In summary, microdialysis studies suggest that, with maternal suppressed basal DA, pups take
on robust saliency to hormonally primed or postpartum females, as reflected in larger increased
pup-evoked DA release, even in the absence of actual interactions with pups. In the absence of the
hormones of parturition, DA release also occurs in response to pups in maternal animals but not in
nonmaternal animals; however, this elevation is not preceded by a reduction in the baseline char-
acteristic of the hormonally primed animal and is sustained for a shorter duration. There is, never-
theless an additive effect of maternal experience on DA release. DA therefore likely mediates pup
salience brought about through exposure to hormones or to sustained pup exposure and thereby
contributes to the expression of maternal behavior.
Pereira et al. (2011) also found a robust DA increase in NA core associated with pup-seeking
behavior (presentation of pups behind a screen). Such DA release was further augmented dur-
ing active (but not passive) maternal interaction with pups. Although the pattern of DA release in
relation to maternal behavior was similar in late postpartum females, the magnitude of release was
considerably attenuated in late, compared to early, postpartum females behaving maternally (Pereira
et al., 2011).
DA transients (spike-dependent fast increases in DA; Grace, 1995) during mother-offspring inter-
action have been investigated using in vivo voltammetry. Champagne et al. (2004) found increased
DA signal in the NA in mothers that showed heightened licking and grooming of their newborns;
however, the onset of the DA signal preceded the behavior. DA transients were also analyzed by
Robinson et al. (Robinson, Zitzman, and Williams, 2011) during pup investigation, and immedi-
ately before or during retrieval. The major transient DA signaling was found while the mother was
50
investigating the cage and newborns prior to retrieving them, and during the first retrieval; subse-
quent retrievals were not associated with DA transients. Thus, Robinson et al. (2011) proposed that
DA transients might play a role in facilitating initiation and maintenance of seeking and appetitive
aspects of maternal behavior. However, as the authors also suggest series of studies in a more natu-
ralistic environment could contribute to our understanding of the role of DA transients in the NA
prior to and during mother-offspring interaction, and in the transitions among the different behav-
ioral components (retrieving, licking, nest building, hovering over the pups, and nursing). It might
also be interesting to determine if those transients of DA change across lactation.
In addition to seeing changes in extracellular dopamine or dopamine transients associated with
measures of mothering, the injection of certain dopamine receptor agonists (e.g., apomorphine) or
antagonists (e.g., haloperidol or raclopride) disrupts various maternal responses, including pup lick-
ing, pup retrieval, and nest-building and/or nursing behavior (Giordano, Johnson, and Rosenblatt,
1990; Li, Davidson, Budin, Kapur, and Fleming, 2004; Stern and Protomastro, 2000). In all these stud-
ies, the drug-treated animals generally have longer latencies to retrieve pups and retrieve fewer pups
than vehicle-treated animals, although at lower drug doses they show only minimal motor deficits.
That these effects reflect motivational deficits is reflected in similar effects of dopamine antagonists
on many types of “approach” behaviors. For instance, if new mother rats are fitted with a muzzle so
they cannot retrieve pups, they will push pups with their snouts and paws. Dopamine antagonists
also block this response, at concentrations that are too low to affect actual pup retrieval (Stern and
Keer, 1999).
That dopamine function within the nucleus accumbens and mPFC is important for these effects
is suggested by studies that show that, if dopamine containing cells within the nucleus accumbens
or the midbrain (VTA) are destroyed with a neurotoxin (6-OHDA) or by lesions or if dopamine
antagonists or GABA agonists are infused into the nucleus accumbens or mPFC, new mother rats
retrieve pups more slowly (Febo et al., 2010; Hansen, Harthon, Wallin, Löfberg, and Svensson, 1991a;
Hansen et al., 1991b; Keer and Stern, 1999b). In contrast, pup nursing is elevated by dopamine
blockade (Keer and Stern, 1999a), indicating that it is the motivation rather than consummatory
behavior that is affected by the activity in the mesocorticolimbic dopamine pathways. This idea is
further supported by results showing that dopamine antagonists, administered to new mother rats,
also block the formation of a conditioned place preference for an environment that was previously
paired with rat pups (Fleming, Korsmit, et al., 1994). The fact that these maternal deficits only occur
if mothers have recently interacted with pups, but not if they have experienced a period of separation
from them, suggests that blocking brain dopamine primarily affects motivation to retrieve and not
the motor mechanisms of retrieval. What happens behaviorally when an animal experiences a deficit
in dopamine function is an open question. Many behavioral mechanisms can be invoked. Disruption
of DA function might (1) produce a general anhedonia, where pups are no longer experienced as
pleasurable stimuli (Wise, 1985; Wise, Spindler, and Gerberg, 1978); (2) dampen animals’ incentive
motivation to approach pups (Bindra, 1978; Bolles, 1972); or (3) inhibit flexible approach responses
towards pups by impairing the behavioral invigoration processes induced by pups (Ikemoto and
Panksepp, 1999).
The mesolimbic system controls maternal motivation, and the integrity of this system is required
for animals to exhibit normal motivated behavior. This conclusion is consistent with a large body
of evidence that has implicated the mesolimbic dopamine system in other motivated behaviors,
including feeding (Bassareo and Di Chiara, 1999; Wilson, Nomikos, Collu, and Fibiger, 1995), food-
foraging (Whishaw and Kornelsen, 1993), drinking (Miyazaki, Mogi, Araki, and Matsumoto, 1998),
drug-seeking (Di Chiara, 1998; Wise, 1998), and sex (Everitt, 1990; Mitchell and Gratton, 1994).
Dopamine and the maternal “core”. We hypothesize that the mesolimbic and mesocortical dopamine
systems exert their influence on maternal motivation by interacting with the MPOA and vBNST.
First, the mesolimbic and cortical systems have reciprocal neural connections with MPOA and
51
Aya Dudin et al.
vBNST at both the nucleus accumbens and VTA (Stolzenberg and Numan, 2011). Thus, in response
to pup cues, the MPOA may cause an increase in dopamine release into the ventral striatum via its
action on the VTA, possibly via the oxytocin-expressing projecton neurons (Hansen et al., 1993;
Hansen et al., 1991a; Shahrokh, Zhang, Diorio, Gratton, and Meaney, 2010; Stolzenberg and Numan,
2011). The increased dopamine in the nucleus accumbens acts to suppress the accumbal GABAergic
inhibitory input to the ventral pallidum (VP), allowing VP efferents to promote active maternal
responses. This idea is consistent with observations that bilateral inactivation of the VP disrupts rat
maternal behavior, as does the unilateral inactivation of VP paired with a contralateral depression of
MPOA activity (Numan et al., 2005). Second, the nucleus accumbens has long been regarded as the
limbic-motor interface and is responsible for converting information from the limbic systems (hip-
pocampus, amygdalal and prefrontal cortex, and so forth) into motor actions (Willner and Scheel-
Kruger, 1991). It is interesting to speculate that the function of the MPOA/vBNST in the control
of maternal motivation is to provide specific “maternal” information and pass it to the dopamine
systems, which in turn, activate the “motivation” and “motor” control systems (extrapyramidal motor
system) to execute the motor outputs. The mesolimbic and mesocortical systems may also feed back
to the MPOA/vBNST to regulate the appetitive component of maternal behavior (Numan and
Young, 2016).
The Serotonin Systems in Maternal Aggression

and Maternal Performance
Serotonin is a neurotransmitter implicated in many psychological processes relevant to maternal
behavior, such as anxiety, depression, affiliation, impulsivity and aggression (Cools, Roberts, and
Robbins, 2008; Graeff, Guimaraes, De Andrade, and Deakin, 1996). It also innervates several mater-
nal brain regions, including the MPOA, BNST, olfactory bulb, hippocampus, and amygdala (Angoa-
Perez et al., 2014; Steinbusch, 1981), and influences the release of parturitional hormones (Bagdy,
1996; Barofsky, Taylor, Tizabi, Kumar, and Jones-Quartey, 1983). There are surprisingly few studies
focusing on 5-HT in maternal behavior, possibly due to several earlier studies showing that disrup-
tion of 5-HT neurotransmission only causes a transient and nonspecific deficit (Barofsky, Taylor, and
Massari, 1983). However, genetic work on mutant mice mothers suggests that genetic mutants lack-
ing full synthesis of 5-HT display impaired pup retrieval, nursing, and nest building (Alenina et al.
2009). Other mouse mutants with impaired 5-HT metabolism also showed reduced reproductive
fitness and abnormal maternal behaviors (Girirajan and Elsea, 2009). Because these 5-HT related
genes are critical for natural brain maturation and homeostatic modulation of neural circuits, lack of
these genes throughout the lifetime may disrupt the development of the neural circuits governing
maternal behavior.
Work focusing on the roles of specific 5-HT receptor subtypes (>14 known 5-HT receptors in
the brain) demonstrates the distinct functions of serotonin receptors in the mediation of maternal
behavior. De Almeida and Lucion (1994) first reported that acute administration of a 5-HT1A recep-
tor agonist (8-OH-DPAT) into brain ventricles or into some brain regions (median raphe, periaque-
ductal gray, and corticomedial amygdala) reduced maternal aggression, whereas later studies showed
that, when the same receptor agonist was infused into other brain regions (medial septum and dorsal
raphe nuclei), it increased maternal aggression (da Veiga, Miczek, Lucion, and de Almeida, 2011; De
Almeida and Lucion, 1997). These data suggest that the 5-HT1A receptors located in different brain
regions play different roles in maternal aggression.
Similar work also shows that activation of 5-HT2A/2C receptors in the brain decrease maternal
aggression (De Almeida and Lucion, 1994) as well as affect other aspects of maternal care. For
instance, atypical antipsychotic drugs like clozapine (CLZ), all possessing a strong antagonistic action
52
against 5-HT2A/2C receptors, disrupt active components of maternal behavior (e.g., pup retrieval, lick-
ing, and nest building) in a dose-dependent fashion (Li, Budin, Fleming, and Kapur, 2005; Li et al.,
2004). The involvement of these 5-HT2A/2C receptors in clozapine-induced maternal deficits is sup-
ported by the finding that pretreatment with DOI (a selective 5-HT2A/2C agonist) dose-dependently
reverses the clozapine-induced disruption of pup retrieval (Zhao and Li, 2009). Furthermore, admin-
istration of DOI alone is able to disrupt maternal performance (Zhao and Li, 2010). Because both
5-HT2A/2C receptor antagonist CLZ and agonist DOI disrupt maternal behavior, this study suggests
that balanced 5-HT2 neurotransmission is critical for the normal expression of maternal behavior.
Too little or too much 5-HT signaling via 5-HT2A/2C receptors could lead to maternal disruptions.
Because both atypical antipsychotics and DOI are nonselective for 5-HT2A versus 5-HT2C recep-
tors, their relative contributions in the regulation of maternal behavior have been the focus of a series
studies conducted by Li and his colleagues. Using highly selective agonists and antagonists against
5-HT2A and 5-HT2C, Chen et al. (2014) and Wu et al. (2016) demonstrated that activation of either
5-HT2A or 5-HT2C receptor causes a disruption of maternal behavior, although blockade of them
has little impact.
Although activations of 5-HT2A and 5-HT2C receptor cause a similar disruption of maternal
behavior, they may do so through different behavioral mechanisms. Wu et al. (2016) provided evi-
dence showing that pharmacologic activation of the 5-HT2C receptor disrupts maternal behavior via
its suppression of mother rats’ motivation to take care of the young, whereas activation of 5-HT2A
receptors likely exerts its disrupting maternal behavior by disrupt the well-programmed sequential
order of behavioral acts, and not by affecting the incentive (“liking”) or instrumental (“wanting”)
value of pups (Chen et al., 2014; Wu et al., 2016).
To identify the brain regions where 5-HT2A and 5-HT2C receptors might be involved in maternal
behavior, Gao et al. (submitted) used c-Fos immunocytochemistry and found that acute pharmaco-
logic activation of 5-HT2A receptors increases neural activation (c-Fos expression) in the ventral bed
nucleus of stria terminalis (vBNST), central amygdala (CeA), and dorsal raphe (DR), whereas Wu
et al. (2016) reported that acute activation of 5-HT2C receptors decreases c-Fos expression in the
ventral part of lateral septal nucleus (LSv), MPOA, dentate gyrus (DG), and dorsal raphe (DR), and
increases it in the central amygdala (CeA). Furthermore, after microinjection of the 5-HT2c agonist
(TCB-2) into two brain regions important for the normal expression of maternal behavior: the
mPFC and mPOA. Only acute intra-mPFC infusion suppressed pup retrieval, whereas intra-mPOA
infusion had no effect. These findings suggest that the 5-HT2A receptor in the mPFC and possibly in
the vBNST, DR and CeA, and the 5-HT2C receptor in the LSv, MPOA, DG, and possibly CeA may
be required for the normal expression of maternal behavior.
Based on the available data, three neural systems where 5-HT2A and 5-HT2C receptors may achieve
their maternal effects are proposed. First, the 5-HT2A and 5-HT2C receptors in the VTA and NA
may modulate the neuronal activity of dopamine neurons and dopamine release (Alex and Pehek,
2007; Bailey et al., 2016) to affect incentive aspects of maternal care (Li and Fleming, 2003b; Numan,
2007). The observation that reunion with pups increases the concentration of both the metabolite
of 5-HT as well as the concentrations of dopamine (DA) and DA metabolites in the ventral striatum
is consistent with the DA × 5-HT interaction idea (Hansen et al., 1993). Second, the 5-HT2A and
5-HT2C receptors in the mPFC could modulate the glutamatergic and GABAergic neurotransmis-
sion and regulate impulsivity and motivation (Dalley, Mar, Economidou, and Robbins, 2008; Liu,
Bubar, Lanfranco, Hillman, and Cunningham, 2007; Pentkowski et al., 2010). Finally, the 5-HT2A and
5-HT2C receptors in the lateral septum (LSv), central amygdala (CeA), and median and dorsal raphe
may interact with other subcortical structures (e.g., medial preoptic area) to alter maternal behavior.
Apparently, more work is needed to pinpoint the exact neural circuitry through which these two
5-HT2 receptors influence maternal behavior.
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Aya Dudin et al.
Maternal Learning and Experience Effects

In this section, we discuss the neural and chemical controls of the maternal experience effect. Mater-
nal behavior beyond one week after parturition is maintained primarily by sensory influences and
processes of learning and reinforcement. Thus, the neuroanatomical and neurochemical substrates
of maternal experience effect may not be the same as those underlying the expression of mater-
nal behavior. Brain sites, which are involved in the control of sensory processing and learning and
memory processes, may be involved.
Neurochemical Bases of Maternal Experience

Because maternal experiences at parturition result in a long-term facilitation of behavior, similar to
other forms of learning, there must occur within the brain mediating structural (i.e., synaptic) or
functional (i.e., neurotransmitter, receptor) changes. One approach to understanding neuromolecular
changes associated with maternal experience was adopted by (Fleming, Cheung, and Barry, 1990)
who asked whether maternal memory requires the synthesis of proteins in the brain in the same way
as do other more traditional forms of memory (Davis and Squire, 1984). In their investigations of the
maternal experience effect (Fleming et al., 1990) and (Malenfant et al., 1991) found that, if a drug
that inhibits protein synthesis was injected into dams immediately after a 1-hour exposure, the long-
term retention of maternal behavior was blocked and at test animals behaved like inexperienced
virgin animals. Moreover, the consolidation of a specifically olfactory experience acquired during
the exposure phase was also blocked by drugs that inhibit protein synthesis. Later work showed that
the new protein synthesis in the NA shell, rather than in the NA core or MPOA, may be more
critical for the maternal experience effect (Li and Fleming, 2003a), as microinjection of a drug that
inhibits protein synthesis (cycloheximide at a high dose) into the NA shell immediately after 1 hour
of maternal experience disrupted maternal memory, whereas infusions in the MPOA had no effect.
It is unclear how these synthesis-blocking drugs interfere with the retention of the maternal
experience. One possibility is the noradrenergic system. Moffat, Suh, and Fleming (1993) injected
noradrenergic antagonists or agonists into dams immediately after a brief maternal experience and
tested them 10 days later for the retention of maternal behavior. Animals receiving the adrenergic
blocker exhibited reduced responsiveness, whereas those receiving the agonist exhibited elevated
responsiveness during subsequent induction tests. However, more than one neurochemical system is
probably involved in maternal memory acquisition and/or retention.
Possible other candidates include the cholinergic system (Ferreira, Gervais, Durkin, and Levy,
1999) and the neuromodulators oxytocin (Amico, Thomas, and Hollingshead, 1997; Broad et al.,
1999; Pedersen et al., 1995) and the endorphins and opiates (Bridges and Hammer, 1992; Byrnes
and Bridges, 2000; Mann and Bridges, 1992). These peptides are present at parturition, influence
maternal behavior, and have been shown in other contexts to influence learning and/or memory
formation (Martinez and Kesner, 1991). Hatton demonstrated that maternal rats, either real mothers
or virgins induced to be maternal by continuous exposure of pups, have profound morphological
and physiological changes in their oxytocinergic neurons in the supraoptic nucleus (SON), one of
the major sites responsible for oxytocin synthesis and release. These changes include increases in
multiple synapses and electrical synapses and decreases in astrocytic processes (Hatton and Tweedle,
1982; Hatton, Yang, and Cobbett, 1987; Perlmutter, Tweedle, and Hatton, 1984; Theodosis and
Poulain, 1984; Yang and Hatton, 1988).
Another neurochemical candidate that synthesis-blocking drugs may interfere with the retention
of the maternal experience is the dopaminergic system (Byrnes, Rigero, and Bridges, 2002). These
drugs may interfere with the dopamine D1 and the D2 receptor subtypes, especially those in the NA
to disrupt the consolidation of maternal memory (Parada, King, Li, and Fleming, 2008). Behaviorally,
54
dopamine in the NA shell may act on both receptor subtypes to alter the motivational salience of
pup stimulation (Afonso et al., 2008), as maternally experienced female rats show increased dopa-
mine levels in the nucleus accumbens when they are re-exposed to pups. More importantly, the more
experience a female has with pups, the greater is the dopamine response. Neurochemically, dopamine
may interact with oxytocin in the nucleus accumbens shell to facilitate the consolidation of maternal
memory (D’Cunha, King, Fleming, and Levy, 2011). Both dopamine and oxytocin are implicated
in the regulation of the salience of social cues, including pups (Shamay-Tsoory and Abu-Akel,
2016). Thus, both can facilitate maternal memory by enhancing the salience of maternal experience.
Indeed, oxytocin plays a role in the consolidation of maternal memory (D’Cunha et al., 2011).
Taken together, it is clear that changes in multiple systems are probably involved in maternal
memory acquisition and/or retention. The possible interactions of neural dopamine, oxytocin and
opioid systems in maternal memory are an important topic for future research.
Neuroanatomical Bases of Maternal Experience

Which specific brain sites are involved in the formation of a maternal memory has been the focus of
a number of immunocytochemical and lesion studies. A number of structures within the maternal
circuit are implicated in the formation of a long-term maternal experience. Besides the NA, the
MPOA and the amygdala might also be involved.
In a series of lesion studies we investigated experimentally which brain sites are implicated in the
formation of consolidation of the experience (Lee, Li, Watchus, and Fleming, 1999). We were inter-
ested to know whether lesions in neural sites that have been implicated either in the actual expres-
sion of maternal behavior or in the formation of memories within other behavioral contexts would
disrupt the long-term experience-based retention of maternal behavior. Of all the sites lesioned,
only lesions to the NA (especially the shell region) prevented the formation of a long-term mater-
nal experience effect. Animals receiving NA lesions either before or immediately after a maternal
experience did not show the faciliatory effects of the experience when tested 10 days later. Instead,
they responded to pups in much the same way as did the totally inexperienced animals, with long
maternal onset latencies. The involvement of the NA in maternal memory is consistent with its role
in the consolidation of other forms (e.g., spatial) of memory (Setlow and McGaugh, 1998; Setlow,
Roozendaal, and McGaugh, 2000; Winnicka, 1999). Because the NA has long been implicated in the
mediation of reward-related processes (Ikemoto and Panksepp, 1999) and is reciprocally connected
with both the MPOA and amygdala (Everitt et al., 1999; Holland and Soedjono, 1981), it is pos-
sible that the NA in combination with these structures mediates the formation of maternal memory
through its role in both reinforcement and associative processes.
Effects of Mothering and Its Absence on the Development

of Maternal Behavior
Many factors influence a new mother’s responses to her offspring. Some are situational; others are
physiological. These factors exert effects by acting on an organism that has a genotype and a devel-
opmental history. Adult characteristics and sensitivities do not emerge de novo, but come about
as a result of a host of earlier influences acting in relation to a genetic background, which include
influences exerted during the earliest stages of development, during the prenatal period; influences
exerted in the nest during the neonatal period; and influences during development through the juve-
nile and adolescent periods. These influences also take many forms; they include various forms of
physical stimulation (associated with variations in temperature, nutritional factors, endocrine factors,
and so forth) and include social influences experienced in the nest with mother and litter-mates, and
with litter-mates and conspecifics during later periods of development.
55
Aya Dudin et al.
There is now good evidence from a number of species that earlier experiences during preweaning
life can exert profound effects on the quality and intensity of mothering the new mother provides to
her offspring postpartum (Fairbanks, 1996; Fleming et al., 1999). Moreover, based on research with
rat and monkey, it seems that these earlier experiences exert their effects throughout development
via multiple routes to alter the neurobiology of the juvenile animal and of the adult animal (Fleming
et al., 1999; Francis, Caldji, Champagne, Plotsky, and Meaney, 1999; Francis and Meaney, 1999; Gon-
zalez, Lovic, Ward, Wainwright, and Fleming, 2001; Kraemer, 1992; Suomi and Ripp, 1983). Due to
space constraints we do not review the ever-growing literature on the effects of early experience on
the development of the neurobiology of mothering. For more on this topic, see Agrati and Lonstein
(2016a); Brett et al. (2015); Lomanowska et al. (2017); Lomanowska and Melo (2016); Pawluski et al.
(2017); and Ragan, Harding, and Lonstein (2016b).
Genetics of Maternal Behavior

The role of genetic factors in the expression of maternal behavior is poorly understood. There have
been attempts by geneticists to uncover quantitative trait loci corresponding to features of maternal
behavior (Sauce, de Brito, and Peripato, 2012). For example, a study using a cross-fostering approach
with genetically defined male and female founders identified a number of loci on offspring chro-
mosomes 5 and 7 in mice that modify maternal behavior (Ashbrook, Gini, and Hager, 2015). The
study went on to show that genetic variation in the mother can influence offspring development
independent of offspring genotype. That is, genetic variation associated with maternal provisioning
of care was found to be independent of genetic variation associated with the solicitation of maternal
care by offspring.
Many more studies using animal models have examined the interaction between specific genes
of interest and the expression of maternal behavior, using gene knockout approaches. Knockout
approaches involve specific genes of interest which are removed or mutated, leading to loss of func-
tion. For instance, deficits in maternal behavior have been reported for mice deficient in the Peg3
gene (Li et al., 1999; Li, Szeto, Cattanach, Ishino, and Surani, 2000), the Peg 1 (Mest) gene (Lefebvre
et al., 1998a), the dopamine beta hydroxylase (Dbh) gene (Thomas and Palmiter, 1997), the Fos B
gene (Brown et al., 1996), the 5-HT1B receptor gene (Brunner, Buhot, Hen, and Hofer, 1999), and
the prolactin (PRLR) receptor gene (Lucas, Ormandy, Binart, Bridges, and Kelly, 1998), to name a
few. In some cases the knockout involves a gene that regulates the synthesis of one of the “maternally
relevant” hormones, neurotransmitters, or their receptors. When this is the case, the expected deficits
in maternal behavior often occur, as with the PRLR receptor knockout (Lucas et al., 1998); however,
sometimes the expected deficits do not occur, as with the oxytocin knockouts which continue to
show robust maternal behavior (Young et al., 1997), despite the fact that oxytocin release occurs at
parturition and during lactation and has been implicated in the regulation of maternal behavior in
a wide range of mammalian species (Bell, Erickson, and Carter, 2014; Noonan et al., 1994; Pedersen
et al., 1994). In the case of oxytocin knockouts, only the synthesis of the hormone was eliminated,
not the oxytocin receptors which could theoretically be activated by other neurochemicals, such as
ligands (Gimpl and Fahrenholz, 2001). Other deficits in maternal behavior seen in the knockouts
may be due to the absence of genes that regulate activity and inhibition or ability to learn (Brun-
ner et al., 1999). These deficits that affect more than only maternal behavior itself illustrate a cen-
tral problem in interpreting the results of these classic gene manipulation studies. The deficit may
occur as a result of deficits in a number of core behavioral or physical features, and may therefore
not inform about the neurobiology of maternal care per se, but may relate to associated behaviors.
Where deficits do not occur, this may reflect either redundancy in neural systems mediating mater-
nal care or compensation as a direct result of the genetic manipulation. The interpretation of both
deficits or lack of deficits are particularly problematic with genetic approaches that affect germ cells,
56
as the manipulation could exert its impacts throughout all stages of life. Newer techniques involving
antibiotic- or chemical-sensitive transgenes used to control the temporal specificity of expression,
coupled with tissue-specific inducers of expression such as the CRE-LoxP recombination system, can
be used to overcome some of these limitations (Gierut, Jacks, and Haigis, 2014).
Other approaches have used optogenetic methods to identify highly specific neural circuitry
involved in the expression of maternal care. These techniques involve hijacking specific genetically
defined neural circuits to express light-sensitive ion channels and thus control neural activity using
laser stimulation. These genetic methods thus enable exquisite temporal and spatial precision of
neural activity in circuits hypothesized to underlie aspects of maternal behavior. The technique was
used to show that the stimulation of oxytocin neurons projecting from the periventricular nucleus of
the hypothalamus to the left auditory cortex can produce maternal retrieval in virgin mice (Marlin
et al., 2015). Importantly, these studies contrast with traditional pharmacological approaches in their
selective modulation of highly specified neural circuitry, as other maternal behaviors known to be
responsive to changes in oxytocin are unaffected.
New research has shown that a number of genes can also influence maternal care in a non-
Mendelian fashion. In these cases, the estimation of the influence of some genes on maternal behav-
iors depends on whether the genes originate from the maternal allele or the paternal allele. These
special genes, termed imprinted, rely on DNA methylation to silence one or the other allele, while
the other remains unmethylated and hence is expressed. DNA methylation is a form of epigenetic
modification, constituting a heritable mark on the DNA itself that does not change the underlying
DNA sequence. As a result, gene mutations in imprinted genes can have profound consequences,
because only one allele is expressed and, if mutated, cannot be compensated by the activity of the
other allele. Examples of imprinted genes that affect maternal behavior are the aforementioned Peg3
and Mest genes, where mutations of these paternal alleles lead to poor maternal care and impaired
milk letdown (Curley, Barton, Surani, and Keverne, 2004; Lefebvre et al., 1998b). The potential of
imprinted genes on maternal behavior has recently become more complex, as parental biases in the
expression of imprinted genes have been detected in dozens of genes, particularly those expressed
in the brain (Perez et al., 2015). These data indicate that the expression of imprinted genes may shift
between monoallelic expression to biallelic expression depending on tissue type and developmental
age. The role for parental bias in gene expression on maternal behaviors and ultimately offspring fit-
ness is currently unknown but an important question for future research in epigenetics.
While epigenetic modifications (of imprinted genes) can alter maternal behavior as a result of
genetic differences, environmental factors are also associated with modifications of epigenetic mech-
anisms relevant for maternal care. Work by Champagne and Meaney has revealed that relative differ-
ences in levels of care provided to pups is associated with modifications of the epigenome in female
offspring (Champagne and Curley, 2008). Specifically, they found that the promoter region of the
estrogen receptor alpha (ERα) gene in the medial preoptic area of the hypothalamus was hyper-
methylated and expression reduced in adult female offspring that had received low levels of care
compared to those receiving high levels of care relative to the mean of the cohort of dams examined
(Champagne et al., 2006). In females, ERα plays a major role in reproductive behavior and also in
maternal behavior, as evidenced by increased c-fos in ERα-positive cells during the postpartum
period (Cameron et al., 2008; Lonstein, Greco, De Vries, Stern, and Blaustein, 2000). These findings
are of potential importance for maternal care in light of the fact that levels of maternal care of female
offspring in adulthood are highly correlated with those of their mothers (Champagne, Francis, Mar,
and Meaney, 2003). Curiously, epigenetic modifications as a function of differences in maternal care
received during the first week of life emerge at weaning on postnatal day 21, indicating that other
mechanisms are likely involved in programing ERα activity in early life (Pena, Neugut, and Cham-
pagne, 2013). Nevertheless, these experiments showed by cross-fostering that these effects depend
on the care received by the foster mother, not the birth mother. These findings indicate that the
57
Aya Dudin et al.
mechanism of transmission of the epigenetic modifications was behavioral rather than by gametic
inheritance.
The above research underlines that complex interplay between genes and environment in influ-
encing maternal behavior. It is perhaps obvious, then, that a more complete understanding of the role
of genotype in maternal care will depend on understanding the biological mechanisms that mediate
gene and environment interactions to affect mothering.
Conclusions
Despite the extent of our knowledge of the control of maternal behavior in nonhuman rodents,
there is much still to learn. We know a great deal about the role of hormones in the onset of
maternal behavior, but very little about their role in its long-term maintenance. We know that the
parturitional hormones exert multiple effects on behavior, but we have very little understanding of
where they act in the brain to effect their behavioral changes; hormone-binding sites outside the
MPOA must surely be involved. We know that the MPOA/vBNST is a critical part of the neural
circuitry, but we do not really understand its precise function; it seems not to function as a sensory
integrator or a motivational “center”; the MPOA/vBNST may simply function as part of the effec-
tor mechanism. Similarly, we have learned a great deal about the multiple limbic and cortical systems
that interface with the core hypothalamic, midbrain systems, but precisely how they interact online
is not known. Despite our increasing knowledge about the role of individual neuropeptides and
neurotransmitters in the regulation of maternal behavior, we have limited information on how these
neurochemicals interact in the behaving animal. Our understanding of the sensory regulation of
maternal behavior is somewhat more complete, but even in this area there is some debate regarding
the extent to which single versus multiple modalities are involved in the onset and in the mainte-
nance of maternal behavior. We know that even very brief experiences acquired when the mother
interacts with the young can have robust long-term effects, profoundly altering the dam’s response
to subsequent external stimulation and to hormones. However, our understanding of where these
long-term effects are encoded in the brain and by what mechanism(s) is still not complete. Finally,
our knowledge of the genetics of maternal behavior and of the interactive effects of genes and the
developmental environment in which the young grow on adult maternal behavior is in its infancy.
The importance of epigenetic mechanisms in the regulation of maternal behavior is clear and yet
surprisingly few studies have demonstrated such effects. This area is wide-open for further study.
An additional set of issues, not touched on in this review, but which would be extremely fruitful to
pursue, is a consideration of the natural history and social factors in the regulation of both maternal
behavior and its physiological underpinnings. Many of the animals discussed, and rodents in particu-
lar, live in social groups in a natural environment characterized by seasonal variations in temperature,
food sources, and photoperiod. All these factors could substantially constrain the maternal system.
Although maternal behavior in the rat has a specific, quite stereotyped, species-characteristic
pattern, its expression depends on the activation of general processes that are stimulated in a variety
of diverse contexts. The onset of maternal behavior involves a hormonally mediated change in the
dam’s affective state and in the salience of pup-related olfactory and somatosensory cues. These
changes increase the likelihood that the dam will approach and maintain proximal contact with
pups, thereby creating the possibility for pup stimuli to elicit the specific maternal responses. Once
maternal behavior is expressed, other processes become activated to insure that the behavior will be
sustained beyond the period of hormonal priming. Once again, these other processes are not specific
to the maternal system, but occur within other functional contexts. For instance, when the mother
interacts with the young, pups acquire heightened reinforcing properties and through perceptual
learning and/or other associative processes, the mother sustains responsiveness to them until weaning
commences. All these processes occur in the adult animal that has had numerous experiences earlier
58
in its life—experiences being mothered. This expression in turns affects the subsequent development
of offspring and how they, in turn, will respond to their own offspring. Moreover, the general nature
of many behavioral processes activated in the maternal animal is accomplished through the media-
tion of physiological mechanisms that are recruited in a variety of functional and stimulus contexts
and that have had a particular developmental history. Thus, olfactory, limbic, and hypothalamic sys-
tems and their associated neurotransmitters, known to influence the expression and maintenance of
maternal behavior, are also activated during the acquisition and/or consolidation of learned behav-
iors within aversive, feeding, and sexual contexts.
Finally, it appears that many of these general processes also play a role in the regulation of maternal
behavior in other mammalian species (Pryce, 1992), including human beings (Barrett and Fleming,
2011; Fleming et al., 2016; Lonstein et al., 2015). As a result, our understanding of the control of dif-
ferent species-characteristic patterns may be enhanced by understanding some of these more general
processes, with the rat model providing a useful heuristic for their analysis.
Acknowledgments
Many thanks to all the lab technicians, animal care workers, undergraduate and graduate students
who made many of the studies reported herein possible. They know who they are. Thanks also to
our families who have patiently withstood the absences necessitated by work reflected in this chapter.
Short portions of the text contained herein are based on other review papers on which the authors
were co-authors, especially Lonstein et al., 2015; Olazábal et al., 2013a, 2013b. Work reported in this
chapter was supported by National Sciences Engineering Research Council (NSERC), Canadian
Institutes for Health Research (CIHR), and National Institute of Mental Health (R01MH097718).
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3
PARENTING IN
NONHUMAN PRIMATES
Kim A. Bard
Introduction
Conceptual and practical issues associated with parenting in nonhuman primates and with mater-
nal competence in chimpanzees, specifically, are addressed in this chapter. An important aspect of
parenting, emphasized here, is that caregiving differs across ontogeny in primates. There are both
short-term and long-term consequences of different parenting experiences, some of which differ
across primate species. Although we might like to know universally what makes a good primate par-
ent, the answer must be considered separately for different species. For some species, it seems evident
that there are specific behaviors that must be learned to be a competent parent (for example for
chimpanzees: Bard, 1994a, 1994b, 1995a, 2002), however, the need for learning parenting behaviors
may not be true for all nonhuman primate species. In addition, the influence of early experience
on maternal capability remains an unknown factor. These central questions of parenting in primates
have been asked for close to 50 years and still have not been satisfactorily answered (Rogers and
Davenport, 1970).
The foundation of basic knowledge, provided by field studies of the 1920s to 1960s (Altmann,
1967; Bingham, 1932; Carpenter, 1964; Dolhinow, 1972; Hall, 1962; Itani, 1959; Jay, 1962; Jolly,
1966; Kummer, 1967; Morris, 1967; Nishida, 1968; Nissen, 1931; Reynolds, 1967; Schaller, 1963; van
Lawick Goodall, 1968; Zuckerman, 1932), allowed specific issues to be addressed, such as the follow-
ing: (1) the genetic and evolutionary bases of parental care (Trivers, 1974), attachment (Chisholm,
2017), and male-infant relationships (Deag and Crook, 1971; Strum, 1984; Whitten, 1987); (2) the
functions served by nonparents providing infant care (Hrdy, 1976, 2009; Lancaster, 1971; McK-
enna, 1979; Quiatt, 1979; Rowell, Hinde, and Spencer-Booth, 1964); (3) explanations for infanticide
(Hausfater and Hrdy, 1984; Hrdy, 1976; Nicolson, 1987; Quiatt, 1979); (4) explanations for disrup-
tions/dysfunctions in parental care (Bard, 1994a; Maestripieri and Carroll, 1998, 2000; Nadler, 1980;
Reite and Caine, 1983); and (5) the manner in which social systems influence patterns of parental
care (Chisholm, 2017; Hawkes et al., 2017; Hinde and Spencer-Booth, 1967; Maestripieri, 1994).
Investigating these central issues has raised more specific questions. For instance, what are the
major factors that influence the evolution of cooperative care? It was thought that males would
provide care when paternity was certain, so monogamy was considered to be the best predictor of
cooperative care. When types of care were compared across monogamous primates, some did have
cooperative care but others did not (Wright, 1990). The ratio of infant(s) weight to mother’s weight
was also considered to be a predictor of cooperative care, but it was also found to be a poor candidate
78
Parenting in Nonhuman Primates
(Gursky, 2000). Cooperative care of infants is associated with increased vigilance and defense against
predators (Caine, 1993; Snowden, 1996). Comparing humans to other primates, the presence of
post-reproductive alloparents (i.e., grandmothers) may be an evolutionary driver for cooperative
care (Hawkes et al., 2017; Hrdy, 2009). Comparing primates to other mammals, however, shows that
most primate mothers receive some type of help in the care of their offspring (Isler and van Schaik,
2012). Additionally, modern humans, across many small-scale societies, as well as those living in urban
industrialized communities, have short interbirth intervals, with infants weaned while still in early
infancy (Dettwyler, 2004; Hawkes et al., 2017; Sellen, 2001). A perennial issue is how nonhuman
primate parenting relates to human parenting.
This chapter is concerned with describing parenting behavior in primates. There are many differ-
ent social and ecological environments and parenting behaviors experienced by primate subadults:
“behavioral plasticity is the very hallmark of primates” (Hawkes et al., 2017, p. 71). Chimpanzees are
used as the basis for comparison with other primate species. Chimpanzees are our closest evolution-
ary relatives, sharing over 90% similarity in genetic material (King and Wilson, 1975), and provide
important information relevant to human behavior. The majority of research on primate parenting
has been conducted in a relatively limited number of primate species, even though our knowledge
of different primate species has expanded across the decades. This chapter, as a product of our limited
knowledge, focuses on the well-studied species. Parenting behaviors in chimpanzees are described
in detail and explicit comparisons are made in parenting behavior between chimpanzees and other
primates.
This chapter contains two main parts, descriptions of species-typical parental behaviors and
longer-term consequences of infant experiences of parenting (or their lack). The section describ-
ing species-typical parental behavior is further divided by taxonomic divisions and by offspring age
beginning with parental behaviors directed toward newborn chimpanzees and ending with parental
behaviors directed toward adolescent prosimians. Social dynamics may also influence parenting styles
(Maestripieri, 1994), but are not discussed in detail here. This perspective provides insights on pri-
mate parenting because the focus is on different parenting behaviors that are required for offspring
of different ages for different primate species.
Parenting Behaviors in Nonhuman Primates

Different skills are required for parental care of infants of different ages (Tardif, Harrison, and Simek,
1993). The age of individuals within each period differs between the species due to different rates
of development (Table 3.1). Parenting behaviors are discussed separately for newborns, infants, juve-
niles, and adolescents. The newborn period is defined in humans as the initial 28 days after birth
during which the infant is most vulnerable and at risk. It is unclear whether there are objective mark-
ers of this period, in humans or other primates (Bard and Nagy, 2017). Infancy is the period when
the offspring is physically dependent on the mother’s milk. Weaning marks the end of the infancy
period. The juvenile period, between the end of weaning and the beginning of sexual maturity, is
distinguished by longer times spent further away from the parent(s), and sometimes accompanying
changes in coat color. The adolescent period begins at puberty and ends at the time when effective
reproduction occurs (Walters, 1987). The age estimates for weaning (the end of the infancy period),
puberty (the beginning of adolescence), and attaining adulthood (first birth for females, and full
adult size for males) are summarized in Table 3.1. The skills necessary for parenting offspring at each
developmental period may have different developmental histories. Therefore, both the skills and
their ontogeny are discussed within each age period separately. Parenting behaviors are additionally
presented within sections by order (see Figure 3.1).
There are major differences between species in the skills required for maternal competence. For
many species of monkey, infants are motorically capable soon after birth. Maternal competence in
79
Kim A. Bard
Table 3.1 Estimates of parenting life history variables from wild primate populations
Weaning Sexual maturity Full adult size
Females Males Females Males
Great and Small Apes

Chimpanzees ~5 yrs ~9 yrs ~10 yrs ~14.5 yrs ~18 yrs
Gorilla ~3 yrs ~6.5 yrs ~9 yrs ~10 yrs ~15 yrs
Orangutan ~5 yrs ~10 yrs ~14 yrs ~14.5 yrs ~17 yrs
Gibbon ~2 yrs ~6.5 yrs ~11 yrs ~13 yrs
Old World Monkeys
Baboon ~12 mos ~5 yrs ~5 yrs ~7 yrs ~7.5 yrs
Vervet ~12 mos ~4 yrs ~5 yrs ~5 yrs ~6 yrs
Blue monkey ~12 mos 6.7 yrs ~6.5–10 yrs ~8–10 yrs
Rhesus ~12 mos ~3 yrs ~3 yrs ~4 yrs ~8 yrs
New World Monkeys
Howler ~9 mos ~3 yrs ~4 yrs ~4.5 yrs ~7 yrs
Spider ~15 mos ~4.5 yrs ~7.5 yrs
Squirrel ~4–18 mos ~2.5 yrs ~3.5 yrs ~3 yrs ~8 yrs
Capuchin ~16 mos ~4 yrs ~4.5 yrs ~4 yrs ~4.5 yrs
Tamarin ~3 mos ~1.2 yrs ~3.5 yrs
Marmoset ~2 mos ~14 mos ~1.5 yrs
Prosimians
Ring-tailed lemur ~3 mos ~3 yrs ~3 yrs
Lesser galago ~1.5 mos ~9–10 mos ~1.1 yrs
Gray mouse lemur 4–6 wks ~3 mos ~3 mos
References: Chapman and Chapman, 1990; Cords and Chowdhury, 2010; Castanet et al., 2004; Cords (pers comm)
many monkeys, therefore, involves only acceptance of the infant (i.e., allowing the infant to cling).
For example, rhesus infants at birth are able to cling, climb on the mother’s body, and suckle. In
other words, rhesus infants can survive as a result of their own behavior, as long as the mother does
not actively reject them (i.e., pull them off her body and prevent them from clinging). In contrast,
for chimpanzees as in humans, maternal competence requires active cradling and nurturing. New-
born chimpanzees are as helpless to survive without maternal support as are human newborns. As
described in Bard (1995a, 2002), Winston, Barbara’s newborn baby, could not move into her arms;
Barbara needed to take the active role and to pick him up—but this was one of the behavioral skills
that she lacked. Maternal competence in chimpanzees therefore requires the mother to take positive
action, including picking up the helpless newborn. Competence in all species is defined broadly as
the ability to raise offspring to adulthood.
This section of the chapter concentrates on parenting during infancy for a number of reasons.
Primarily, parenting responsibilities are greatest during infancy when offspring are least capable of
coping on their own. The second reason is that there are already good reviews on juveniles (Pereira
and Fairbanks, 1993) and adolescents (Bernstein, Ruehlmann, Judge, Lindquist, and Weed, 1991;
Caine, 1986), although little is known about those parenting behaviors specifically directed to juve-
niles and adolescents. Adolescence is typically the time when emigration occurs, and offspring may
permanently leave the family group. In most primate species, it is the sons that leave, and the daugh-
ters that remain in the natal group with their mothers. Although parental status within the group
may be crucial to the long-term outcome for an adolescent, observable parent-offspring interactions
are minimal.
80
Marmosets
Tamarins
Capuchins
Squirrel monkeys
Night monkeys
Howler monkeys
New World monkeys
Wooly monkeys
and spider monkeys
Uakaris, sakis,
and titi monkeys
Macaques
SIMIANS
Mandrills
Old World monkeys Baboons

Vervets and guenons
Colobus monkeys
Langurs, proboscis,
CATARRHINES
and leaf monkeys
Gibbons
Orangutans
Chimpanzees
Great Apes
Bonobos
Humans
Gorillas
Tarsiers
Lemurs
Ruffed lemurs
PROSIMIANS
Aye-ayes
Indris and sifakas
Mouse lemurs and

dwarf lemurs
Galagos (Bushbabies)
Lorises
Figure 3.1 Classification of living primates frequently mentioned in this chapter.

Source: Reprinted with permission (Myowa and Butler, 2017, p. 54).
Kim A. Bard
Great Apes and Small Apes
Chimpanzee Demographics
Captive chimpanzee mothers are approximately 11 to 12 years old when they first give birth (Fra-
gaszy and Bard, 1997; Roof, Hopkins, Izard, Hook, and Schapiro, 2005). If they care for their infant,
the subsequent infant will be born 3 to 4 years later (interbirth interval), and they will have approxi-
mately 4 babies in their lifetime (Fragaszy and Bard, 1997; Roof et al., 2005). In the wild, chimpan-
zee mothers are approximately 11 to 17 years old when they have their first baby (Goodall, 1986;
Nishida et al., 2003; Sugiyama, 2004), although at some field sites, such as Mahale, many first babies
die or are victims of infanticide (Nishida, 2012). At most field sites, female chimpanzees transfer
out of their natal group before giving birth (Roof et al., 2005) and may be younger when first giv-
ing birth than those who stay in their natal group (Nishida et al., 2003). The interbirth interval in
wild chimpanzees whose infants survived was 5 to 7 years, but there is substantial variation by site
(e.g., 5.2 years at Budongo; 5.3 years at Bossou; 5.6 years at Gombe; 6 years at Mahale; 6.6 years at
Kibale; Emery Thompson et al., 2007), and by sex at some sites (e.g., 5.5 years following birth of
daughters versus 6 years following birth of sons at Mahale; Nishida et al., 2003). Wild chimpanzees
have approximately three babies in their lifetime (Emery Thompson et al., 2007; Potts, 2013). There
is some variation in these life history parameters across different field sites and between chimpanzees
and bonobos. Bonobo mothers in the wild first give birth when they are 14 years old, and their
interbirth interval is between 4.5 years at Wamba and 8 years at Lomako (Wich et al., 2004). The
oldest wild chimpanzee females to give birth to surviving infants varied from 40 years at Bossou to
55 years at Kibale (Emery Thompson et al., 2007).
Data on the birth of three chimpanzee infants found strong similarities in the delivery compared
with human newborns (Hirata, Fuwa, Sugama, Kusunoki, and Takeshita, 2011). Specifically, the
chimpanzee infants emerge from the birth canal facing away from the mother, as is found in humans.
There are considerable changes in orientation through the birth canal, as the head and body subse-
quently rotated, before dropping to the floor (two cases) or being gathered by the mother (one case).
This orientation at birth was previously thought to be unique to humans and was proposed as the
evolutionary basis for the use of midwives (Hirata et al., 2011). It appears that, among all the nonhu-
man primates, chimpanzees share the most similarities to humans in lifespan characteristics.
Parenting Newborn Chimpanzees and Apes

The newborn period is defined as the initial period after birth during which the infant is unable
to survive without parental support. For some primate species, there is really no clearly definable
neonatal period subsequent to the minutes after birth; for others the period lasts through the first
30 days as is true for human newborns (Bard, Brent, Lester, Worobey, and Suomi, 2011; Bard and
Nagy, 2017; Brazelton, 1984). This section discusses those special parental skills applied to newborns,
distinct from the parental behaviors to infants. The term infants refer to nonhuman primate infants:
Human infants are distinguished explicitly.
Field studies of free-living chimpanzees do not richly describe newborn chimpanzee behavior in
part because the very small neonate is difficult to detect on the body of the mother (weighing 1.5
kg, on average: Fragaszy and Bard, 1997) and in part because new mothers appear to be very cau-
tious or wary and are not often observed (Goodall, 1986; Plooij, 1984; van Lawick-Goodall, 1967).
Newborn chimpanzees do not have any distinctive coloration, although their skin is lighter in color
during infancy than the skin of adults (Nishida, 2012).
The chimpanzee newborn and mother are in constant ventral-ventral contact during the first 30
days of life (van Lawick-Goodall, 1968). Newborn chimpanzees are as helpless to survive without
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maternal support, as are human newborns. Although both newborn chimpanzees and newborn
humans have a strong grasping reflex (Bard, Hopkins, and Fort, 1990), this reflex is insufficient to
support the infant for more than a few seconds at a time. Unlike most primates, chimpanzees are
unable to support their own weight independently for at least the first two months of life (Bard,
Platzman, Lester, and Suomi, 1992; van Lawick-Goodall, 1968; Nishida, 2012; Plooij, 1984; Rijt-
Plooij and Plooij, 1987). Mothers provide the majority of physical support during this time although
they seldom help neonates to suckle. Feedings are short in duration and irregularly spaced (Brown
and Pieper, 1973; Dienske and Vreeswijk, 1987; Plooij, 1984).
Detailed observations of newborn chimpanzees with their mothers are possible in laboratory set-
tings, such as at the Yerkes Research Center of Emory University and the Primate Research Institute
of Kyoto University. In captive settings, nuzzling, rooting, and nursing account for approximately
20% of the chimpanzee infant’s waking time during the first month of life (Bard, Platzman, and
Coffman, 1989), but appears to account for only 2%–4% of infant’s time in field settings (Lonsdorf
et al., 2014). Infants nuzzle and root to find the nipple, typically without any assistance by the mother.
Suckling begins within the first day or two after birth, and averages 1 to 2 minutes every hour during
the day and night (Bard, 2002; Mizuno, Takeshita, and Matsuzawa, 2006; Nishida, 2012). Newborn
chimpanzees sleep during approximately 50% of observation time throughout the first 30 days of life.
During sleep, EEG reveals brain wave patterns (REM sleep, deep sleep, etc.) in chimpanzee newborns
that are similar to those of human newborns (Balsamo, Bradley, Bradley, Pegram, and Rhodes, 1972).
Behavioral observations reveal that newborns are in REM sleep ~35% of the nighttime and in deep
sleep ~35% of the nighttime (and awake at night, ~30%: Mizuno et al., 2006). The newborn is alert
and quiet for considerable periods, especially on the first day of life and increasingly through the first
month, an average of 25% of the time. Active alert states are apparent but account for less than 10%
of observation time during the first month of life. Newborn chimpanzees raised by their mothers
do cry and fuss, but these periods of moderate to mild distress are infrequent and brief (Bard, 2000).
The vast majority of maternal behavior with newborn chimpanzees is simply cradling, providing
the support they need to remain in physical contact (over 80% of the time; Bard, 1994a; Goodall,
1967; Nishida, 2012). Other maternal activities, in addition to cradling, occur for an average of 10
minutes per hour. These additional activities include grooming the infant (6%), playing with the
infant and eliciting some smiles (3%), examining the infant (2.5%), assessing the behavioral and
physical state of the newborn (2%), and encouraging the infant’s motor development with physical
exercises (1%) (Bard, 1994a; Bard et al., 2005).
Chimpanzees are responsive to their social environment, even as newborns. Bard et al. (2011)
conducted tests of the neurobehavioral integrity of newborn chimpanzees raised in one of three
different nursery environments and compared their performance with those raised with their moth-
ers in another captive setting (mother-raised data originally reported in Hallock, Worobey, and Self,
1989). These environments differed in the degree of exposure to humans and degree of cradling
contact. At 2 days of age, newborn chimpanzees significantly differed in the domains of Orienta-
tion (both to social and inanimate stimuli) and State Regulation (especially self-quieting skills) as a
function of rearing environment. By 30 days of age, almost all of the items and clusters (all 7 items
of the Orientation cluster, 4 of 5 items of the Motor cluster, 4 of 4 items of the Range cluster, and
all 3 items of the Regulation cluster) showed significant differences among the chimpanzee rear-
ing groups. In both the early part of the newborn period (2 days of age) and the latter part of the
newborn period (30 days of age), the largest differences were found when comparing the newborns
which were raised with their chimpanzee mothers to those raised in nurseries by human caregivers
(19 items and clusters from a possible 24 showed significant differences at 30 days of age). But there
were numerous significant differences among the different nursery chimpanzee groups as well (7 of
24 possible differences between the Southwest Foundation and the Yerkes Standard care nursery
groups, and 2 differences between the Responsive Care Intervention and Yerkes Standard care
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nursery groups). Surprisingly, the chimpanzees raised by their mother appeared to have lower levels
of neurobehavioral integrity in several domains (e.g., Orientation, Motor Performance, State Regu-
lation) compared to the human group and the Standard care nursery group at 30 days of age (Bard
et al., 2011). These lower scores are probably due to the fact that the mother-raised environment
is more protective and restrictive of the newborn infant than are nursery environments, with less
exposure to humans and human artifacts, and greater experience with constant cradling contact.
Gaze is an important aspect of primate behavior, and mutual gaze is an important aspect of
human mother-infant interaction (Trevarthen, 1979; Tronick, Als, and Adamson, 1979). On average,
chimpanzee mothers at Yerkes spend 12 minutes an hour looking at their newborn infants (Bard,
1994a). Half of that time is spent looking at the infants’ face, which occurs during maternal activities
of assessing, examining, playing, and grooming. Newborn infants also gaze at the face of their mother
(Goodall, 1986, p. 86). Numerous instances of mutual gaze occur between mother and newborn
infant, 10 times in an hour on the average (Bard, 1994a; Bard et al., 2005).
Newborn chimpanzees, like human newborns, can mimic facial movements of their caregivers
(Bard, 2007; Bard and Russell, 1999; Myowa-Yamakoshi, Tomonaga, Tanaka, and Matsuzawa, 2004).
Chimpanzee newborns appear to imitate mouth openings, tongue protrusions, and sequences of
mouth actions, including making sounds, such as a tongue click (Bard, 2007). Neonatal imitation,
therefore, is one mechanism by which parents can directly influence the behavior of their offspring,
during face-to-face interactions, especially when they are communicative (Bard, 2007). However,
there are additional influences on newborn behavior that may be more indirect or subtle, such as via
amount of cradling contact or degree of exposure to culture-specific practices, such as eye gaze pat-
terns or nurturing style of caregivers, for example (Bard, 2017; Bard et al., 2005, 2011). These early
interactions can have long-lasting consequences. For instance, the amount of nurturing experienced
by chimpanzees in the first month of life was found to be correlated with structural co-variation in the
gray matter of the basal forebrain (e.g., reward circuit) in adult chimpanzees (Bard and Hopkins, 2018).
Chimpanzees have a neonatal period that appears to be as distinct as is the human neonatal period
(Bard and Nagy, 2017). Newborn chimpanzees are very responsive to different parenting practices,
some of which have long-term consequences (see the following and Bard and Hopkins, 2018).
Parenting Newborn Gorillas, Orangutans, and Small Apes

The social structures of orangutans and gorillas differ from that of chimpanzees, and one might
expect differences in parenting, even parenting newborns. Orangutans are the most solitary of
the great apes (although orangutan mothers seem to spend as much time alone as do chimpanzee
mothers: Galdikas and Teleki, 1981). Wild orangutan mothers give birth when they are approxi-
mately 15 years old and have the longest interbirth interval among the great apes, at 9.3 years at
Ketambe (Sumatran subspecies), 7.7 years at Tanjung Puting, and 7.0 years at Gunung Palung and
at Tuanan (Bornean subspecies; van Noordwijk, Willems, Atmoko, Kuzawa, and van Schaik, 2013;
Wich et al., 2004).
Gorilla groups consist of a dominant male silverback and five to seven unrelated females (i.e.,
harem). Mountain gorilla females first give birth when they are 10 years old, and their interbirth
interval is 4 years (Wich et al., 2004).
Newborn orangutans and gorillas appear more capable motorically compared with chimpanzees
(Figure 4.2 in Bard, 2002). Orangutan mothers with newborns do not travel far or quickly, and they
rest frequently (Galdikas, 1982). Maternal support of the infant may be minimal even on the first
day of life, and the placenta may or may not be eaten (Fossey, 1979; Galdikas, 1982). Gorilla mothers
with newborns are given preferential proximity to the father, the silverback male. Newborn gorillas
can cling, unsupported by the mother, for up to three minutes (Fossey, 1979). In gorillas, the social
group is important for the maintenance of maternal competence, perhaps heightening protective
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responses (Nadler, 1983). In captive settings, new gorilla mothers isolated from the group typically
exhibit abusive behavior ( Joines, 1977; Nadler, 1983).
The small apes, gibbons and siamangs, have long been thought to be exclusively monogamous
and territorial, but several species regularly include more than two adults, and females engage in
copulations outside of their “pair-bond” (Hawkes et al., 2017; Sommer and Reichard, 2000). Inter-
birth interval is 2 to 3 years in siamangs (Lappan, 2008). As expected the males engage in paternal
care (mostly carrying) when offspring are older, but in the newborn and infancy periods, mothers
provide exclusive care in the small apes (Lappan, 2008). Gibbon mothers provide cradling support
to newborns and reposition the infants to a safer spot on the mother’s body, for example, prior to
leaping between trees (Carpenter, 1964).
The first weeks after birth has been identified as the newborn period for gorillas, orangutans,
bonobos, and the small apes (gibbons and siamangs). Although it is not clear whether this is a distinct
developmental period, it appears that caregivers do attend with extra care to the newborn.
Parenting Chimpanzee Infants

Infancy is the period when the offspring is physically dependent on the mother’s milk. The infancy
period is often differentiated into an early period, during which no independent locomotion occurs,
and a later period, during which there is some independent locomotion although the infant remains
close to the parent(s) during the day and night. Great apes (chimpanzees, orangutans, bonobos and
gorillas) remain in infancy for 4 to 6 years.
Goodall (1986) classified infancy in chimpanzees as the period from birth to the time of weaning
(and cessation of travel on the mother’s body), which occurs at approximately four to five years of
age in the wild (Clark, 1977; Nishida, 2012) but weaning occurs earlier, around 3.5 years, in cap-
tive settings (Fragaszy and Bard, 1997). The early infancy period is characterized by almost constant
physical contact. The first break in contact is typically initiated by the 3- or 4-month-old laboratory
infant (Miller and Nadler, 1981; van Lawick-Goodall, 1968). By 3 months the amount of maternal
restraint of infant movement has increased fivefold to about 15% of the maternal interactive time
(Bard, 1994a; Bard et al., 2005), indicating how active the infant has become and that the mother is
responsible for maintaining the infant’s proximity to her at this age.
In the first 3 months, chimpanzee mothers (with good maternal competence) engage their infants
in a variety of interactions (Bard, 1994a; Goodall, 1967; Plooij, 1984), although the majority of
maternal time is still spent simply cradling their young infants (71% at Yerkes, but less at PRI ~40%:
Bard et al., 2005). In captive settings, mothers spend almost 30% of their interactive time groom-
ing 3-month-old infants, and 14% of their interactive time playing with their 3-month-old infants
(Figure 3.2). Although there is variation across captive settings, when their infants are 4 months old,
caregivers encourage a variety of species-typical skills at a rate of 7 to 11 minutes per hour (Bard,
Bakeman, Boysen, and Leavens, 2014), and nurture social communicative skills, in particular, at a
rate of 2 to 3 minutes per hour (Bard, Dunbar et al., 2014). Infant motor development is stimulated
through maternal maneuvers, such as standing infants while holding their hands. Mothers repeat-
edly and alternatively stimulate their infants to hold their weight with legs and then with arms. At
3 months, there was significantly more exercising of infants at PRI (33% of noncradling time) com-
pared with at Yerkes (7%; Bard et al., 2005). Encouragement of early crawling is accomplished in a
similar way. Because mother-infant contact is rarely broken in these early months, these stimulating
exercises are typically performed on mother’s body. “Sooner or later every mother encourages and
variously aids her baby to learn to creep, stand erect, climb, and finally to walk and run” (Yerkes and
Tomilin, 1935, p. 333).
Early mother-infant communication in chimpanzees is often accomplished with touch (Plooij,
1979) and can be accompanied by vision and audition. Mothers monitor their infant’s behavioral
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Kim A. Bard
Figure 3.2 Mother and 3-month-old infant chimpanzees, Pan troglodytes, engage in mutual gaze 10 times per
hour, on an average, at the Yerkes National Primate Research Center Field Station, Atlanta, Georgia
(Bard et al., 2005).
Source: Photo credit: Joshua A. Schneider.
state by stretching and moving infants’ toes, fingers, arms, and legs and sometimes just by looking at
them. Chimpanzee infants smile with a playface, which consists of very little lip corner withdrawal,
but lots of jaw dropping with an open mouth (Figure 3.1 in Bard et al., 1992). During play, human
infant smiles are sometimes “marked” as critical features by the mother with an emphasized touch
(Adamson and Bakeman, 1984). When the chimpanzee infant smiles in response to a tickle in the
neck or groin, the chimpanzee mother may place her index finger on the infant’s lower gums and
exaggerate the smile by pushing gently on the lower gums, “marking” it in a similar way as do some
human mothers. In a responsive care nursery program for abandoned chimpanzee infants, human
researchers used this technique to highlight the importance of different skills as they emerged in
young chimpanzee infants (Bard, Dunbar et al., 2014).
Previous reports indicated very limited episodes of mutual gaze in mother-infant great apes
(Papousek, Papousek, Suomi, and Rahn, 1991; Plooij, 1979; Rijt-Plooij and Plooij, 1987). Chimpan-
zee mothers spend considerable amounts of their time gazing at their young infants’ body (~14% of
observation time) and gazing at the infant’s face (~11% of observation time). Chimpanzee infants
gaze at their mothers’ faces. Very young infant chimpanzees appear to have a greater visual acuity
at 30 cm than 15 cm but see quite comparably to human infants at 15 cm (Bard, Street, McCrary,
and Booth, 1995). Early social environments influence the expression of behavior in chimpanzees
and humans, and eye gaze patterns between mother and infant vary during the first 3 months of life
(Bard, 1994b; Bard 1992; Bard et al., 2005). At Yerkes, mothers appeared to be sensitive to infants’ eye
gaze and shifted their own gaze away whenever mutual eye gaze was attained (Bard, 1994a), in strik-
ing contrast to the extended mutual gaze encouraged by some human mothers (Trevarthen, 1979;
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Tronick et al., 1979), or when humans are interacting with infant chimpanzees (Bard, 1998b; Bard
et al., 1992). However, some captive and wild chimpanzee mothers actually encouraged extended
durations of mutual gaze by lifting the infants chin to establish and maintain mutual gaze (Bard et al.,
2005; van Lawick-Goodall, 1967). Chimpanzees at the Primate Research Institute, Kyoto University,
had an average of 27 bouts of mutual gaze per hour with their 3-month-old infants, a rate that is
very similar to that of Western human mothers with their 3-month-old infants (Bard et al., 2005).
For infant chimpanzees, as well as infant humans, these differences in eye gaze patterns tend to
accompany differences in proximal versus distal caregiving patterns (Bard et al., 2005; Keller, 2007).
When caregiving is more proximal, with extensive cradling contact (as in Yerkes chimpanzees and
many subsistence rural human communities), there is less mutual gaze, whereas when caregiving is
more distal (as at PRI laboratory and in middle-class Western human communities) then mutual
gaze is more extensive (Bard et al., 2005). Thus, it may be that one of the early behaviors that is
“culturally” regulated in chimpanzees (and humans) is eye gaze (Bard, 2017; Bard and Gardner, 1996:
Bard et al., 2005; Figure 3.2).
Independent quadrupedal steps and climbing appear as early as 4 months of age in field settings
(Rijt-Plooij and Plooij, 1987), and independent quadrupedal location is common by 5 months
in captive settings (Bard, 1996). In wild settings, in the first 6 months, infants spend about 12% of
observation time clinging to the mother’s belly or chest while she travels (i.e., riding ventrally; Lon-
sdorf et al., 2014). From 5 to 7 months, wild chimpanzee infants begin to ride on the mother’s back
(males ~4%; females ~1% of observation time; van Lawick-Goodall, 1968; Lonsdorf et al., 2014).
Male infants make the shift to more dorsal than ventral riding earlier (~1 year) than do female infants
(~1.5 years; Lonsdorf et al., 2014). Typically, it is not until 1.5 years of age that infant chimpanzees
reliably respond to the mother’s communicative signals to “climb aboard” (van Lawick-Goodall,
1968). One mother was explicitly observed to teach her young infant to climb on her back when
she displayed a hunched posture while looking over the shoulder, which constitutes a communicative
signal (Rijt-Plooij and Plooij, 1987). From 5 or 6 months, the mother places the infant on her back
or repositions the infant from ventral to dorsal position. Some argue that mothers act roughly to
attain dorsal riding and breaks in contact, but careful reading suggests that typical maternal behavior
at Gombe is determined, rather than aggressive (Rijt-Plooij and Plooij, 1987).
Observations of 19 chimpanzee infants from the Kanyawara community in Kibale National Park,
and the Tai South community in Tai National Park, provide details about the subtle and frequent
gestures employed by mothers and infants in coordinating joint travel (Frohlich, Wittig, and Pika,
2016a). Mothers used an average of 10 gestures and four actions to initiate carrying of 10- to
12-month-old infants, including multimodal signals, such as uttering a soft “hoo” vocalization while
presenting their back for the infant to climb on or while reaching their arm to the infant (Frohlich
et al., 2016a). Infants used an average of two or three gestures, and two actions, to initiate maternal
carrying. As infants develop, they change from using whimpering alone, for example, to using more
visual gestures, such as a reach, either alone or in multimodal combinations. Thus, chimpanzee moth-
ers provide both physical support and encouragement for their young infants’ motor developments
(Bard, 1994a; Frohlich et al., 2016a; Goodall, 1967; Yerkes and Tomilin, 1935). The responsive care
nursery program at Yerkes used these infant motor milestones and maternal encouragement behav-
iors to design a program to encourage 5-month-old chimpanzees, for example, to engage in dorsal
riding (Bard, 1996).
The subtle communication between mother and infant is documented in “meshing”. Rijt-Plooij
and Plooij (1987) discussed meshing only in the locomotor context and defined it as maternal
anticipation of and coordination with the infant’s contact behavior. Meshing occurs from 8 to 24
months but monthly levels rise and fall in correspondence with the infants’ responsibility for contact
maintenance. “It is the mother’s role to (force) teach the infant how to use newly emerged abilities it
might not, or not fully, have used otherwise” (Rijt-Plooij and Plooij, 1987, p. 72).
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From 8 months, infants and mothers are comfortable out of physical contact and within arm’s
reach, but infants whimper when mothers move too far away (Rijt-Plooij and Plooij, 1987). It is per-
haps no different from increased separation anxiety observed in human infants from 7 to 9 months,
when human infant’s developing cognitive processing allows them to distinguish novel from familiar,
and they show wariness of strangers (Ainsworth and Bell, 1970; Bard and Gardner, 1996; Bowlby,
1969, 1973; Fritz and Fritz, 1985; Plooij, 1984; Rijt-Plooij and Plooij, 1987; van IJzendoorn, Wil-
lems, Atmoko, Kuzawa, and van Schaik, 2009). A reasonable conclusion from Rijt-Plooij’s descriptive
data is that between 8 and 11 months the infant becomes more responsible for maintaining contact
and proximity with the mother, in contrast to the earlier ages when the mother is mostly insisting
on proximity with the infant.
From 12 months to 18 months infants return to being comfortable within their mothers’ arm
reach. By 18 months, dorsal riding becomes the predominant mode of joint travel (~8% of the time;
Lonsdorf et al., 2014). By 2 years of age, infants are spending time outside of arm’s reach (between 1
and 1.5 meters away), and as they get older they spend increasing amounts of time more distant from
the mother. By 3.5 years of age, sons move further away than daughters (3 meters versus 2 meters:
Lonsdorf et al., 2014). By 4.5 years, older infant chimpanzees spend less than 1% of their time rid-
ing dorsally on their mothers, spend between 5% (females) and 10% (males) of their day traveling
independently, and move up to 4.5 meters (males) and 2.5 meters (females) away from their mother
(Lonsdorf et al., 2014).
Social skills such as greeting social partners and using communicative signals to initiate play or
grooming, develop first in interaction with the mother and then are used in interaction with others,
first older siblings and peers, and then older unrelated individuals (Bard, Dunbar et al., 2014; Nishida,
2012). In the second month of life, infant chimpanzees respond to mothers’ tickles with smiles and
very quiet laughter (Bard, 1998b; Bard, Dunbar et al., 2014; Plooij, 1979). In the third month, infants
reach, with a smile, to initiate tickle games with the mother and sometimes with older siblings (Bard,
Dunbar et al., 2014; van Lawick Goodall, 1968; Plooij, 1979).
In the first 6 months, the primary social partner for infants is their mother (Goodall, 1967, 1986;
Nishida, 2012). Mothers, however, can be more, or less gregarious in exposing their infants to poten-
tial future social partners. Although there are individual differences, on average, mothers with sons
spend more time with both kin and nonkin, including more time with adult males, than do mothers
with daughters (Murray et al., 2014). Infant chimpanzees, in the wild, play on their own, up to 20%
of the time at 1 to 1.5 years of age, and engage in social play, with peak levels at ~15% for males when
they are 2 to 2.5 years, and at peak levels at 20% for females when they are 3 to 3.5 years old (Lons-
dorf et al., 2014). In a captive setting, 12- to 15-month-old infants played during 55% of observation
time and spent only 34% of that time in social play (Ross, Bard, and Matsuzawa, 2014). Tickle play
was not observed with peers, but only between infants and their mothers (and other older individu-
als). Mothers monitor their infants’ interactions with others and can rush to pick them up at the first
sign of the infant becoming distressed. Older play partners engage in “self-handicapping” behaviors,
such as laying supine, or lowering the intensity of play, when they engage with infants in play, thereby
increasing the chance that the infant will play with them and lowering the likelihood that the mother
will intervene (Frohlich, Wittig, and Pika, 2016b).
In the second half year of life, infants may initiate social interactions with others by approaching
them with vocal greetings. Nishida (2012) argued that infants first learn the appropriate greetings
by imitating their mothers when they encounter others. Imitation of greetings, in this case, is a type
of co-action, which is easy as the infant is riding on the mother’s belly which moves as the mother
utters her vocal greeting. Knowing whom to greet may be accomplished when the infant rides on
the mother’s back, as it is easy for the infant to see whom the mother greets.
Infant chimpanzees learn a great deal of social communicative signals in the first two years of life.
Communicative signals constitute all the ways that social partners negotiate social interactions. Some
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might label these communicative signals the natural “language” of chimpanzees. Chimpanzee infants
may learn communicative signals through co-constructed interactions (Bard, Dunbar et al., 2014),
in social negotiations (Frohlich et al., 2016a, 2016b), or mutual shaping (King and Shanker, 2003),
although there is continued discussion about developmental processes (Call and Tomasello, 2007)
and the role of genetics (Hobaiter and Byrne, 2011). Because young chimpanzee infants interact pri-
marily with their mother (in the wild and some captive settings) or with a caregiver (in some captive
settings), the caregiver plays an important role in communicative learning. Infants might learn greet-
ing vocalizations by uttering greetings at the same time, with the same intensity as the mother, when
she directs greetings to a particular individual (Nishida, 2012). Grooming gestures can be learned as
early as the first year of life in a captive setting (Bard, Dunbar et al., 2014), but typically are seen in
the 2- to 3-year-old wild chimpanzee. The grooming hand clasp, observed originally at Mahale, was
observed in older infants/young juveniles (at least 4 to 6 years of age), and sometimes appears to be
molded or scaffolded by the mother (Nakamura and Nishida, 2013; Nishida, 2012; Wood, Bruner,
and Ross, 1976). This social custom, more accurately called high-arm grooming, occurs in many, but
not all wild communities, and can take many forms, including palm-to-palm grasping, wrist-to-wrist,
and others (Wrangham et al., 2016). The mother has a major influence on the form learned by her
offspring. The form used by adult chimpanzees does not conform to that used by the social group,
but rather individuals retain the form they learned with their mothers (Wrangham et al., 2016).
The sharing of food is a negotiated event that involves communicative signals (Bard, Dunbar
et al., 2014) and occurs regularly between infants and caregivers in chimpanzees (Goodall, 1986) and
in orangutans (Bard, 1992: Jaeggi, van Noordwijk, and van Schaik, 2008). Mothers typically allow
young infants, around 4 or 5 months, to take some food from her mouth or hand, but by 9 to 12
months, infants typically use communicative gestures to request the sharing of food (Bard, Dunbar
et al., 2014; Plooij, 1984; van Lawick Goodall, 1968). It appears that chimpanzee mothers selectively
share the more difficult to process or difficult to obtain foods as the infant matures (Nishida, 2012;
Silk, 1978, 1979). Food sharing between mother and infant typically ends when the infant transi-
tions to juvenile status, around six or seven years of age. The same food begging gestures observed in
infants, however, are used by older individuals to request the sharing of meat from adult males after
a hunt (Boesch, 2012; Goodall, 1986; Nishida, 2012).
Data from a large sample of wild chimpanzees (Lonsdorf et al., 2014) show that infant chim-
panzees spend between 2% and 4% of their time nursing, approximately 1.5 to 2 minutes per hour
(Nishida, 2012). Although the level of nursing does not change substantially throughout the infancy
period, eating solid food increases from ~5% at 6 to 12 months, to 20% of the time from 1.5 to
2.5 years, ~33% from 2.5 to 4.5 years, and almost 50% at 4.5 to 5 years (Lonsdorf et al., 2014). In cap-
tive settings, however, suckling changes with age, with approximately 3%–17% of time spent nursing
in the first 4 months of life (Mizuno et al., 2006), and suckling tending to stop altogether within 2 to
4 years, much earlier than in the wild (Goodall, 1986; Nicolson, 1987; Nishida, 2012).
From 9 months, wild and captive infant chimpanzees request food from their mother, typically
with a cupped hand, held underneath the mother’s chin (Bard, Dunbar et al., 2014; Plooij, 1978,
1984; van Lawick-Goodall,1968). Ueno and Matsuzawa (2004) found, in an experimental setting
consisting of giving the chimpanzee mother food items, that these captive mothers sometimes spon-
taneously offered food to their young infants. However, these offers were often the unpalatable parts
of the food items. Nishida (2012) reported that mother chimpanzees at Mahale never spontaneously
offer food to infants, but that infants, through their own initiative, develop their mother’s food rep-
ertoire based on eating those food items being eaten by their mothers, through both “food retrieval”
and successful food sharing (Goodall, 1986).
From 2 to 5 years there is much to be learned about food, food processing, traveling, and hunting.
Chimpanzee mothers monitor what infants eat and prevent them from manipulating or eating unde-
sirable objects, in an example of what Nishida (2012) labels “education by discouragement”. Mothers
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serve as models for older infants to learn termite fishing (Goodall, 1986), ant fishing (Nishida, 2012),
tool manufacture (Wright, 1972), plant foraging (McGrew, 1974, 1977), food processing (Lefebvre,
1985), and locomotory behaviors (Bard, 1993, 1995b). Many food processing skills are thought to be
learned via “education by master-apprenticeship” as Matsuzawa et al. (2001) described for nut crack-
ing. Lonsdorf (2005, 2006) found that the amount of time mothers spend termite fishing alone with
her offspring predicts the number of critical elements acquired by the offspring, but only at 2.5 years.
Proficiency in termite fishing requires mastering five critical elements: identifying termite holes,
manipulating tools, modifying tool material, inserting tools into the hole, and the crucial element of
extracting termites. No maternal characteristics predicted the offsprings’ overall proficiency in ter-
mite fishing (Lonsdorf, 2006). Daughters were more proficient than sons, and daughters spent more
time watching others engage in termite fishing, and more closely matched maternal termite fishing
techniques (selecting similar lengths of tool; Lonsdorf, 2005, 2006). Lonsdorf (2005) suggested that
female offspring actually imitate some aspects of maternal termite fishing, but all offspring use goal
emulation (a social learning process involving copying the end result, but not imitating the means) to
develop understanding that using a tool achieves acquiring termites. The five critical elements were
mastered at different ages: identifying termite holes was mastered around 1.5 years of age, manipulat-
ing tools began around 2.5 years, modifying tool material began around 3.5 years, inserting tools into
the hole was mastered by all offspring by 4.5 years, and extracting termites was not mastered by all
offspring until 5.5 years of age. However, there is a great deal of individual trial-and-error learning
involved in mastering the final critical element of extracting termites (Lonsdorf, 2006).
There is an increasing amount of evidence that Great Ape mothers actively instruct their infants
under some, perhaps limited circumstances. Boesch (1991) argued that chimpanzee mothers “take
an active part in the apprenticeship of their female offspring” (Boesch and Boesch, 1981, p. 592) to
crack nuts with a hammer tool. Chimpanzee mothers facilitate arboreal locomotory behavior by
“bridging” gaps between trees allowing the young infant to cross the gap on her body and allowing
older infants to cross the gap on branches that she holds close together (Goodall, 1986). It is likely
that adult male chimpanzees play a teaching role in the apprenticeship of male offspring in coopera-
tive hunting (Boesch and Boesch, 1989, 2000). Much of the teaching found in small scale hunter-
gatherer communities with human infants involves demonstrations, giving infants tasks to perform,
scaffolding, redirecting, and providing positive or negative feedback (totaling over 10 times per hour),
rather than verbal instruction (less than once per hour) or ostensive cues that provide information to
infants (four times per hour; Hewlett and Roulette, 2016).
Chimpanzee males are remarkably tolerant when infants attempt to interfere with mating, and
males may reassure uneasy infants with a touch. The tolerance of infant behavior appears to continue
as long as the infant retains the “tail tuff ”, long white hairs at the base of the spine (Goodall, 1986).
Chimpanzee males in captivity engage in play with infants showing there is a capability (Bingham,
1927; Taub and Redican, 1984). The difference between gorilla and chimpanzee fathers may be that
paternity in chimpanzees is usually not known either by observers or apparently by the chimpanzees
(Gagneux, Woodruff, and Boesch, 1997; Goodall, 1986), but in gorilla harems paternity is certain.
Male orangutans rarely engage in interactions with infants.
Clark (1977, p. 235) described the 2-year gradual weaning process in 2- to 4-year-old chim-
panzees at Gombe as a period when infants may “display many elements of depression”. It begins
with mothers preventing access to the breast by holding the infant away, pushing the infant away, or
physically blocking access with an arm or knee pressed firmly against their own chest. Mothers often
distract the infant with play or grooming when they attempt to suckle, and mothers may move away
from the infant as the infant approaches to suckle. It was extremely rare for any mother to exhibit
aggressive behavior in relation to weaning her infant. Similar tactics were observed in chimpanzee
mothers in a captive setting (Horvat and Kraemer, 1982). In response to these tactics infants whim-
per and become physically more intrusive in their attempt to access the nipple. As the infant grows
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older and weaning is more strictly enforced, temper tantrums ensue at Gombe but not in the captive
setting. But as weaning progresses elements of depressive response are seen in the wild, including
decrease in play, loss of appetite, huddled posture, and resumption of infantile behavior with the
mother including ventral riding and increased contact (Yoshida, Norikoshi, and Kitahara, 1991).
In the wild, all 4- to 5-year-old infants exhibit distress when the mother’s milk is no longer
available and within months make no further attempts to suckle. Clark (1977) noted that all moth-
ers appear “remarkably tolerant and gentle with the infants during the weaning period” (p. 252)
and increased their attentiveness to the infant through grooming and waiting for them in traveling.
Infants, however, appear depressed through the period of the birth of younger siblings. Their depres-
sion is exhibited in lethargic movements, lack of positive emotions, and sometimes decreased appetite
and moderate weight loss. Although weaning occurs at an earlier age in captive chimpanzees (~2 to
4 years of age compared to 4 to 6 years in the wild), captive chimpanzees appear less affected, as there
is less distress related to the cessation of suckling and no signs of depression (Horvat and Kraemer,
1982), probably due to the abundant availability of food in captive settings.
Parenting Infant Orangutans

Orangutan infants, younger than 2 years, are less frequently out of contact with the mother com-
pared with chimpanzee infants (Figure 3.3; Bard, 1993, 1995b). Mothers may tolerate relatively close
proximity with other mothers in order to allow their infants to play, but fewer peers are available for
socialization in orangutans (compared with chimpanzees) during the entire infancy period (Bard,
personal observation). In the wild food context, mothers respond positively to their infants’ goal-
directed behaviors most often when directed toward herself (rather than directed to the food), and
Figure 3.3 Orangutan mother and 9-month-old infant, Pongo pygmaeus, at Camp Leakey, in the Tanjung Puting
National Park, Kalimantan Tengah, Indonesia.
Source: Photo credit: Kim A. Bard.
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Kim A. Bard
when infants request food with a gesture directed toward the mother, around 3.5 years of age (Bard,
1992; Jaeggi et al., 2008). In captive settings, gestures appear to emerge at earlier ages, and orangutans
(compared to chimpanzees, bonobos, and gorillas) gesture more often in the food context than other
contexts (Schneider, Call, and Liebal, 2012).
Mothers provide support for infants developing foraging skills. In the wild, it is likely that the
initial maternal influence for infants is akin to stimulus enhancement, supporting infant’s learning of
what to eat (co-feeding in the same patch occurred during 90% of bouts; Jaeggi et al., 2010). Infants
watched mothers while they extracted embedded foods, providing observational learning experi-
ences about how to process foods. Infants were more successful in food sharing if they timed their
solicitations for when the mother was finished processing the food and had already taken a bite (Bard,
1992). Additionally, mothers allowed infants to scrounge discarded and partially processed pieces
( Jaeggi et al., 2010; Russon, 2006). Analyzing the complexity of infant’s object manipulations, Bard
(1995b) found that more complex manipulations were used in locomotion compared with foraging.
More direct maternal assistance was found in locomotor, compared to the foraging, contexts in
wild orangutans. Basic assistance is provided by mother orangutans in arboreal traveling, by the infant
clinging to the mother while she travels 85% of time (in contrast to 50% for juveniles; Bard, 1995b).
For infant orangutans, a large proportion of the nonclinging traveling events consisted of maternal
assistance provided to infants in crossing a gap between trees. Infants climbed across on the mother’s
body while she held two trees to bridge a gap or rode in the tree that the mother swayed to cross a
gap (Bard, 1995b). Bard (1995b) suggested that these maternal behaviors could be considered a form
of tuition or scaffolding for young infants, as older infants and juveniles appeared to contribute their
weight to maternal swayings, prior to learning to sway trees independently. Orangutan infants are
weaned between 4 and 8 years of age (Bard, 1993; Galdikas, 1979), but can continue suckling until
the next offspring is born (van Noordwijk et al., 2013).
Parenting Infant Gorillas

The gorilla in early infancy is motorically more advanced than chimpanzees, chewing food items
in the first 2 months of life, and clinging to the mother’s hair without support by 2 months, and
reaching for objects earlier. Mothers spend time grooming the infant and begin to rebuff suckling
attempts before infants’ first birthdays. Gorilla infants play very little with their mothers or with sil-
verbacks, even in captivity, but choose to play alone primarily in the first year, and with same-aged
peers, preferring male partners through the next 2 years (Maestripieri and Ross, 2004). Mothers
do not seem to encourage any social activities of their infants, although they occasionally discour-
age them, especially when infants are young (Maestripieri and Ross, 2004). One study found that
communicative competence was scaffolded for infant gorillas by all older social partners (Luef and
Liebal, 2012). Juveniles, adolescents, and adult gorillas modified their gestural communication when
directed to infants, through using more repetition and higher rates of sequences with a tactile com-
ponent, in a type of communicative “motherese” (Luef and Liebal, 2012) or scaffolding (Wood et al.,
1976). Mothers share food when infants request it with communicative gestures (Maestripieri, Ross,
and Megna, 2002). In captive settings, lowland gorilla mothers encourage the development of infant
locomotor skills in a manner similar to that of chimpanzees, and infants show interest in maternal
activities, especially food processing, and appear to “create opportunities for their own social learn-
ing” (Maestripieri et al., 2002, p. 225; Whiten, 1999). By 2 years of age, mountain gorilla infants travel
primarily independently but they retain the white tail tuff that indicates their infant status through
part of the third year (Fossey, 1979).
Male gorillas undoubtedly play an important role as protector for infant gorillas, as paternity is
usually known (Stewart, 2001). One function of adult males may be as a “social magnet”, being the
focal point for the gathering of many infants and juveniles (Figure 4.5 in Bard, 2002; Stewart, 2001).
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In some wild settings, gorilla infants and their fathers play frequently, in contrast to chimpanzees
(Figure 3.4), but in other captive settings, infant play is almost exclusive with peers, although male
infants play more than female (Maestripieri and Ross, 2004). In addition, gorilla fathers carry some
young infants (Tilford and Nadler, 1978). Some gorilla infants spend more time near or interacting
with their father than with their mother (Fossey, 1979, 1983; Harcourt, 1979). “Gorilla males often
groom, cuddle, and nest with their 3- and 4-year old offspring” (Whitten, 1987, p. 346). The father
also monitors play between infants and stops it before it becomes too rough (Fossey, 1979). Gorilla
fathers, through these early interactions, appear to form a particularly close relationship with at least
one male infant. This preferred infant will grow up to remain in the father’s group (Harcourt and
Stewart, 1981; Tilford and Nadler, 1978).
Parenting Infant Gibbons and Siamangs: Small Ape Infants

Infancy in the small apes lasts 12 to 19 months (Chivers, 1976; Fox, 1977). When the gibbon mother
rests, 6-week-old infants begin to move a little distance from her. Infant gibbons in the first weeks
of life may eat some solid food and engage in locomotor play (Carpenter, 1964), but continue to
nurse for 2 years (Figure 3.5). One of the most striking behaviors exhibited by gibbon parents is
their vocal duet, songs are given morning and evening. “Infants often squeal during a mother’s great
call” (Leighton, 1987, p. 140). As older infants travel independently, they sometimes are unable to
Figure 3.4 Among the great apes, gorilla (Gorilla gorilla) fathers spend the most time interacting with infants.
Source: Photo retrieved from iStock.com/omersukrugoksu.
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Kim A. Bard
Figure 3.5 A white-handed gibbon mother, Hylobates lar, cradles her 8-month-old infant while she eats figs in
the Khao Yai National Park, Thailand. The group’s unrelated adolescent male sits nearby.
Source: Photo credit: Ulrich H. Reichard.
cross gaps between trees, and they “cry” until the mother retrieves them (Carpenter, 1964). Mother
siamangs do almost all of the carrying of infants younger than 8 months. In the second year of life,
fathers carry gibbon and siamang infants (up to 20% of the time) and spend some small amounts of
time grooming and playing with them (Lappan, 2008).
Paternal care in the small apes can be as high as 78% of the day (Whitten, 1987); the infant returns
to the mother to nurse and to sleep at night (Alberts, 1987). For the siamangs, living in the Way
Canguk Research Area in Sumatra, male care replaced female care, and the amount of care by males
differed as a function of whether the group was socially monogamous or not (Lappan, 2008). These
siamang infants began to travel independently around 1 year of age, were weaned between 10 and 15
months of age, and spent most of their time traveling independently by 21 months of age (suggesting
an early emergence of the juvenile period; Lappan, 2008). Older siamang infants still rely on adults
to facilitate their travel under difficult traveling conditions—“across gaps, at high speeds, or when
fatigued” (p. 1311).
Parenting Juvenile Chimpanzees and Apes

In chimpanzees, the end of infancy is indicated by cessation of nursing (weaning) and the disap-
pearance of the white tail tuft (Nishida, 2012; van Lawick Goodall, 1968). The juvenile period is
distinguished by longer times spent further away from the parent(s). Attention turns from mothers
to peers in the juvenile period (Horvat and Kraemer, 1981). Maternal responsibilities in terms of
providing milk and transportation diminish or cease, and responsibility for offspring to become fully
independent increases. It is when offspring reach this period of semi-independence that chimpanzee
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mothers facilitate learning of travel techniques (Goodall, 1986), of food processing (McGrew, 1977;
including tool use; Boesch, 1991), and of socialization. Puberty marks the end of the juvenile period.
After juvenile chimpanzees are weaned, they remain in close association with their mothers,
traveling with the mother but not being carried by her. Mothers are still the primary grooming
partners for juvenile chimpanzees, but daughters more often groom family members compared with
sons (Preuschoft, Chivers, Brockelman, and Creel, 1984). Juvenile sons and daughters play, groom,
and carry younger infants who may or may not be siblings (Nishida, 1983). All juveniles exhibit
submissive behaviors to adult males, for example presenting their hindquarters and pant grunting.
Occasionally juveniles display and attack adolescent females, but this level of aggression only occurs
when the mother joins to support her son or daughter (Pusey, 1990).
During the juvenile period, chimpanzees, gorillas, and orangutans eat the full range of food items
found in the diet of adults. The mother is the main source of information about what to eat, how to
eat, and when to eat different foods, and for chimpanzees this information includes learning about
tool use for termite fishing and nutcracking (Boesch, 2012; Lonsdorf, 2006). Many of these skills are
exhibited during the late infancy period, but adult-like success is not attained until the juvenile or
even adolescent periods in chimpanzees and orangutans (Russon, 2006).
Gorilla juveniles receive most grooming from their mothers (Watts and Pusey, 1993), but the rate
of grooming appears to be much less than found in chimpanzees. Similarly, there are few bouts of
grooming in mother-offspring orangutans (Galdikas, 1995).
Siamang juveniles spend most of their time traveling independently by 21 months of age (Lappan,
2008), but remain with their family group. Juveniles (2 to 4 years of age) begin to receive aggressive
behaviors from their parents: typically, mothers harass daughters and fathers harass sons (Preuschoft
et al., 1984). Fights ensue most often over access to food (Leighton, 1987). Juvenile gibbons and
siamangs may join in singing the duet with their parents. The song tends to be sex-appropriate but
imperfect (Leighton, 1987). During the juvenile period, mother gibbons initiate co-singing episodes
with their daughters, but later, adolescents begin co-singing episodes (Koda et al., 2013).
Parenting Adolescent Chimpanzees and Apes

The adolescent period begins at puberty and ends at the time when effective reproduction occurs
(Walters, 1987). The beginning of adolescence, signaled in female chimpanzees with small sexual
swellings, occurs around eight to nine years of age in wild chimpanzees (Nishida, 1988; Watts and
Pusey, 1993), and orangutans (Galdikas, 1979) and 5.5 years in the laboratory (Sarah Phythyon, per-
sonal communication, 1993). Menarche and full sexual swellings occur when a chimpanzee is 11 to
12 years of age in the wild and 8 to 10 years in the laboratory. The adolescent period includes the
time when offspring travel independently throughout days and nights, sometimes engaging in sexual
activity and reproductive behavior, but the individual is neither fully socially or physically adult.
Adolescence lasts from the age of 9 to 14 years in wild female chimpanzees (Nishida, 1988), until
15 years in male chimpanzees (Nishida, 1988), 16 years in gorillas (Watts and Pusey, 1993), and until
21 years in male orangutans (Galdikas, 1979). Mothers now have a more general influence on their
offspring. For example, maternal rank significantly influences the age of their daughter’s first full
sexual swelling. There is as much as a 4-year difference between the ages for daughters of high versus
low ranking mothers (Pusey, Williams, and Goodall, 1997). Some physical changes that character-
ize the adult status include coat color (e.g., silver-colored hair on the backs in the dominant male
gorilla), secondary sexual characteristics (e.g., cheek flanges in male orangutans), and attainment of
full growth (e.g., canines, testes, and general body size).
At adolescence, there are striking differences between chimpanzee daughters’ and sons’ behavior
in whom they groom, and with whom they spend their time. Sons are more often in the company
of adult males than daughters. Mothers provide support to their daughters in agonistic encounters,
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Kim A. Bard
whereas sons solicit and receive support from older brothers. The behavior of adolescent males is
molded by adult males who touch to quiet adolescent males during boundary patrols (Pusey, 1990)
and guide adolescent males in assuming complementary and cooperative roles while hunting colobus
monkeys (Boesch and Boesch, 1989, 2000). It is during mid-adolescence, when female chimpanzees
exhibit adult-sized, sexual swellings that daughters may leave their mothers by joining a new group,
and then are solicited and protected by adult males (Pusey, 1990).
Adolescent orangutans (7 to 8 years to 10 to 15 years: Wich et al., 2004) are relatively sociable,
traveling in mixed-gender groups (Galdikas, 1995). Females preferentially consort with adult males
but continue to spend some time with mothers as well, before settling into a home range than will
overlap, or be nearby the mother’s (Bard, 1995b; Galdikas, 1979; Watts and Pusey, 1993).
During adolescence in the small apes (6 to 8 years in gibbons: Koda et al., 2013), fighting occurs
over breeding access and adolescents leave (or may be evicted) from the family. Male adolescent gib-
bons appear to be inhibited from singing with their parents but females emit great calls simultane-
ously with their mother. The calls of the daughter are acoustically similar to those of the mother,
and it appears that the mother adjusts her calls when her daughter sings along (Koda et al., 2013).
Thus, mother gibbons have flexibility in their calls, and they play an important role in the “acoustic
learning” of their daughters (Koda et al., 2013). Call rates of daughters decrease during the adoles-
cent period with their increasing independence and become more adult-like in vocal structure. The
call rates of daughters predict when they leave the family group to begin their own family (Koda
et al., 2013). Fathers and adolescent sons may form a coalition in territorial defense against intruders.
Fathers may facilitate the process of their sons establishing their own territory, either by joining the
sons to usurp a neighbor’s territory or by expanding the home territory and then leaving the son in
the new area (Leighton, 1987). Young adult male gibbons sing solo apparently to attract unmated
females, but unmated females rarely sing alone (Leighton, 1987).
Summary
Apes spend 10 to 18 years in subadult states (Table 3.1). Apes both require longer periods of parental
care, and benefit from parenting for the longest periods of time. Among nonhuman primates, parent-
ing in the apes most closely approximates that seen in humans, as parents nurture the development
of many social, communicative, and motor skills in their offspring.
Old World Monkeys
Parenting Newborn Old World Monkeys

Macaques (e.g., rhesus, cynomologus, pigtails, and bonnets) have a strong crawling and grasping reflex
that may aid in the birth process (Rosenblum, 1971; Tinklepaugh and Hartman, 1932). Their “strong
righting reflexes and negative geotropism . . . function to produce the proper orientation” to emerge
from the birth canal (Rosenblum, 1971, p. 324). Mothers must provide a supportive base when expel-
ling the newborn, especially important when the mother is in an arboreal environment, or else the
infant may not initially get a grasp of the mother’s hair (Rosenblum, 1971; Timmermans, 1992).
Little attention is paid to the newborn in many species, but in some, particularly those with allo-
maternal care, newborns are of great interest to other adult females, for example in vervet monkeys
(Fairbanks, 1990; Lancaster, 1971) and to large juvenile females, for example in blue monkeys (Forster
and Cords, 2005). The placenta is eaten by some mothers during the initial birthing period. This
period is followed by a period of intense grooming of the infant. Mothers may gaze at their newborn
infants and infants may gaze at their mother’s face (Higley and Suomi, 1986). Newborn macaques,
through reflexive behaviors, suckle without maternal aid. Nipple contact is maintained over 80% of
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the time during the first month of life in captive rhesus macaques (Higley and Suomi, 1986), and
over 50% of the time in wild blue monkeys (Föster and Cords, 2002), both are in striking contrast to
the 20% nipple contact of chimpanzees.
New findings demonstrate that there is a short time window, or maybe a sensitive period (Born-
stein, 1989), during which rhesus monkey newborns imitate lip smacking and tongue protrusion
(Ferrari et al., 2006). Mother and newborn rhesus monkeys can show a great deal of reciprocal
affectionate and affiliative face-to-face behavior, including mutual gaze and lip smacking (Ferrari,
Paukner, Ionica, and Suomi, 2009).
Paternal behaviors toward newborns varies by species but generally ranges within the indifferent
category. Mothers with newborns may stay in close proximity to adult males in baboons (Papio anubis;
Hrdy, 1976). One-week-old Barbary macaques are carried by adult males as well as by juvenile and
subadult males (Deag and Crook, 1971), and there is a report of the dominant male holding an infant
four times on the day it was born (Burton, 1972, cited in Hrdy, 1976).
Parenting Infant Old World Monkeys

Most macaques remain in infancy for only 1 to 1.5 years. Baboon infants have black hair and pink
skin in contrast to the light hair and black skin of adults and have a new sibling when 1.5 to 2 years
of age, so are weaned at 1 to 1.5 years of age (Altmann, 1980; Strum, 1987). Macaque infants, and blue
monkey infants, mature quickly and begin to crawl and climb, respectively, within a few days after
birth (Förster and Cords, 2002). In the first few weeks, mother blue monkeys restrict their infants
from moving out of contact. When these infants are older, mothers move away more, although
they monitor the infant and remain ready to retrieve and protect the infant (Figure 3.6; Förster and
Cords, 2002). In contrast, mothers of some terrestrial species encourage infant locomotion (in pig-
tails; Bolwig, 1980; Hinde and Simpson, 1975; Ransom and Rowell, 1972; in macaques; Ehardt and
Blount,1984; Ferrari et al., 2009 [supplemental material]; Maestripieri, 1996). Macaque infants typi-
cally engage in independent excursions in the third and fourth weeks of life. The mother provides a
“secure base” from which the infant travels (Harlow and Harlow, 1965) and a “safe haven” of comfort
to return to when distressed (Maestripieri and Roney, 2006). “Mothers now become psychologically
more than physically important for their infants” (Higley and Suomi, 1986, p. 160). However, during
this time mothers are providing kinesthetic and vestibular stimulation through grooming and physi-
cal contact while traveling, and contingent responsive stimulation in their social interactions (Ferrari
et al., 2009). Mothers respond to infant cries and seem to respond selectively, or at least differentially,
in positive, negative, or neutral manners to all infant social communicative signals. It is this type of
selective and contingent social responsiveness that mothers, in particular, provide to infants that peers
do not. However, it is rare that macaque mothers engage in extended play with their infants (Suomi,
1979). Play in macaques is primarily a peer activity.
Paternal behavior in Old World monkeys varies considerably among species. Mothers stay close
to adult males in baboon species, and male langurs may respond to infants in distress with protection
and rescue (Hrdy, 1976). Young mangabeys spend most of their time with an adult male rather than
their mothers. Intensive caregiving by males is found in one species of Old World Monkeys, the
Barbary macaque. Adult males “groom, nuzzle, and mouth infants, lick and smell them, manipulate
their genitalia, and teeth chatter at them” (Whitten, 1987, p. 345), engaging in interactions analogous
to those of mothers (Taub, 1984). Males may temporarily foster an infant or permanently adopt an
orphaned infant (anubis baboon: DeVore, 1963; Japanese macaque: Itani, 1959; or hamadryas baboon:
Kummer, 1967). In fact, Kummer (1968) reported that motherless infant hamadryas are invariably
adopted by young adult males. But males in each of these species, and in langurs and vervets, also
use infants as an “agonistic buffer”, which puts the infant at risk for injury or death. Males carry an
infant to or near another male: the presence of the infant inhibits aggression and the males interact
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Kim A. Bard
Figure 3.6 Mother and 6-month-old infant blue monkey, Cercopithecus mitis stuhlmanni, in the Kakamega Forest,
western Kenya.
Source: Photo credit: Marina Cords.
in a less tense environment. Cases of infanticide in langurs and rhesus by adult males typically involve
nonfathers and appear to be cases whereby the adult male is maximizing his inclusive fitness (Trivers,
1974), whereas cases of infant care, play, and other affiliative interactions are by dominant males who
are likely fathers (Hrdy, 1976). Infants are used by adult males as agonistic buffers regardless of genetic
relatedness (Whitten, 1987).
Infants in some Old World monkey species experience considerable care from nonmothers (up
to 10% of the time). Nonmaternal handling of infants is common in blue monkeys, as well as vervets,
patas, and langurs. In blue monkeys, for example, during the first months of life, older juvenile
females attempt to kidnap infants, but by the time they are 3 to 4 months of age, allomothers act
much like biological mothers, providing a secure base and a safe haven, and protecting infants from
aggression and providing comfort when distressed (Förster and Cords, 2005). Infant blue monkeys
receive more grooming from nonmothers than from their mother.
Maternal “style” is the term used to differentiate both species differences and individual differ-
ences in maternal behaviors toward older infants that reflect the balance between permissive and
restrictive rearing (Hinde and Simpson, 1975). Maternal style in early infancy is reflected by how
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contact and proximity are regulated and how much contact the infant is allowed with others. In rhe-
sus and pigtail macaques, and baboon species, none may touch the newborn infant for many weeks.
In contrast, colobus and langur mothers may allow others to carry away their newborns (Bennett,
1988; Fimbel, 1992). Bonnet macaque mothers allow their infants to interact in a limited fashion
with others in the social group but not immediately after birth. It is common, however, to allow
older siblings limited access to infants. Mothers exhibit consistent rejection rates with each of their
offspring and across the development of each offspring. Moreover, there is consistency in maternal
style across generations. So, it appears conclusive that maternal style is a characteristic of the mother,
and not the mother-infant bond, in rhesus (Berman, 1990) and vervet monkeys (Fairbanks, 1989).
Maternal “style” is also apparent at weaning. Frequency of rejection and punishment have been
documented to be stable maternal characteristics in rhesus monkeys (Berman, 1990). Rejection and
punishment as a weaning style clearly differentiate species; rhesus have high rejection rates compared
with bonnets for instance (Rosenblum, 1971). By the end of the first year of life, weaning is complete
in Old World monkeys. Baboon mothers hit, push, grab, and bite at infants. Individual differences
between mothers occur, but by the time that infants are 5 months of age they have experienced
maternal aggression at least once (Altmann, 1980). Weaning occurs between 4 and 6 months. Wean-
ing begins somewhat later in Hanuman langurs, and different responses are noted from male and
female offspring, but all mothers are harsh, punitive, or indifferent (Rajpurohit and Mahnot, 1991).
Baboon mothers encourage independent locomotor behavior when infants are 7 months of age.
The mother may descend from sleeping trees without carrying the infant. Infants “protest” with
whimpers. Sometimes mothers return halfway up the tree still requiring that the infant descend part
of the way independently. Mothers always monitor the tree, even if they do not facilitate any of the
travel, until the infant travels down. Typically, the infant immediately runs to the mother and nurses.
By the end of 1 month of these “lessons”, the 8- or 9-month-old infants descend from the sleeping
tree independently.
Competing theories have suggested either that punishment/rejection facilitates, if not causes,
independence (Hansen, 1966; Hinde and Spencer-Booth, 1967), or that high levels of punishment/
rejection cause increased dependence and delays in the attainment of independence (Kaufman and
Rosenblum, 1969; Rosenblum and Harlow, 1963). In experiments on cross-fostered rhesus mon-
keys, Suomi (1987) evaluated the independent contributions of inherited reactivity, foster caregiver
reactivity, and foster caregiver style on infant reactivity. Foster caregiving “style” is a better index of
infants’ behavioral reactivity than inherited temperament, or caregiver temperament, under stable
environmental conditions. However, when presented with environmental challenges, such as a brief
separation from the caregiver, infants’ reactivity was best predicted by inherited reactivity. When ini-
tially returned to the foster mother, then the caregivers’ reactivity best predicted the infants’ behavior
(Suomi, 1987).
Parenting Juvenile Old World Monkeys

In rhesus macaques, baboons, and Japanese macaques, the age of menarche is 5.5 years on the average
but can be influenced by hierarchical dominance status (Pereira and Fairbanks, 1993). Male testes
descend in baboons around 5.5 years of age (Altmann, Altmann, Hausfater, and McCuskey, 1977).
Infants and juvenile baboons are given the preferential center location during group travel.
Mother macaques allow juvenile daughters to handle infant siblings and juvenile sons to play with
infant siblings. Mothers also allow unrelated juvenile females access to infants. Sometimes mothers
are intimidated by juveniles of high-ranking matrilines and their infants are kidnapped (Hrdy, 1976;
Maestripieri, 1994). In agonistic encounters the entire matriline will support their kin. Juveniles
begin to acquire ranks similar to their mothers and exercise dominance toward all females that are
subordinate to their mother.
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In Hamadryas baboons, adult males form “special relationships” with juvenile females. These
begin as friendships but grow to be consortships. Juvenile males may have strong affiliative bonds
with an adult male or with male peers, which develop into coalitions later in life (Kummer, 1967,
1968; Walters, 1987).
Parenting Adolescent Old World Monkeys

There is a great deal of variance in the age of adolescence in Old World Monkeys. In baboons, full
adult size is reached in females around 7 years of age, and full secondary sexual characteristics are
developed by 10 years in males (Altmann et al., 1977). In vervets, breeding is seen at 3 years, but
females are not adult in size until 4 years and males not before 5 years (Fairbanks, 1990). Rhesus
monkeys are in early adolescence between 2.5 and 4.5 years (Bernstein, Judge, and Ruehlmann,
1993) but may remain in adolescence for 4 years or more (Bernstein et al., 1991).
In rhesus, the males leave the birth group during adolescence. Few specifics on parenting are
known to account for the large differences between male and female behavior. Female rhesus spend
significantly more time grooming, and males spend more time with male peers. The result of these
differences in behavior is a loosening of the sons’ bonds with mother and sisters, and a strengthening
of the daughters’ bonds with mother and sisters. The role played by the mother during the process
is unspecified. “Male macaques and vervets frequently emigrate in the company of brothers or natal
group peers” (Walters, 1987, p. 365). In hamadryas baboons, the alpha male acts aggressively toward
subadult males, eventually evicting them from the natal group (Caine, 1986). It is the adolescent
females in red colobus groups, however, that migrate as adolescents (Caine, 1986).
Summary
Old World monkeys spend 5 to 8 years as subadults. Parenting in Old World monkeys is less about
nurturing skills (a characteristic of parenting in apes) and more about exerting an influence on their
path of development. This influence takes many forms that differ across species, from adherence
with strict dominance hierarchies to experiencing extensive alloparental interactions. There is good
evidence in many Old World monkey species, that parents continue to have an influence on their
offspring, even after they become adult.
New World Monkeys
Parenting Newborn New World Monkeys

New World monkeys are arboreal and forest dwelling. The newborn period has not been defined for
New World monkeys but is likely to consist of only the first day or so after birth. Care of newborns
is different for the species that are pair-bonded (e.g., marmosets, tamarins, titi monkeys) and for the
species that live in large social groups (e.g., squirrel monkeys, howler monkeys, and capuchins).
Newborn capuchin monkeys can cling to the mother unaided, but the mother supports the
infant while moving from ventrum to her back (Fragaszy, 1990). Squirrel monkey newborns can
move independently from the ventral position used in nursing, to the dorsal position used in travel.
Early field studies of newborn howlers documented that mothers regularly restrained and pulled the
infants as they continually climbed up the mother’s ventrum (Carpenter, 1964). Although newborn
New World Monkeys have the physical strength to support their weight with their tails, it appears
that newborns are uncoordinated with their tails (Carpenter, 1964: Fragaszy and Bard, 1997).
From the day of birth, squirrel monkey infants respond visually and vocally to the communica-
tion of others. Adult females and juveniles are allowed to touch newborn squirrel monkeys but many
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mothers avoid adult males or prevent them from touching the newborns (Hopf, 1967). Adults vocal-
ize “caregiver calls” to newborns, and newborns vocalize responsively (Biben, 1994). These vocal
exchanges occur when infants and adults are engaged in mutual gaze. It is noteworthy that these
communicative exchanges are not between infant and mother but rather between infants and other
adult members of the group.
Marmosets, tamarins, and titi monkeys are monogamous species, that typically give birth to twins
(80% for marmosets and tamarins); triplets are as common as single births. In marmosets and cebus,
mothers are primarily responsible for carrying, and totally responsible for feeding newborns (Box,
1977; Rothe, Darms, Koenig, Radespiel, and Juenemann, 1993; Wamboldt, Gelhard, and Insel, 1988).
Specialized behavior has developed to cope with twin births (fraternal twinning is typical: Tardif,
Carson, and Gangaware, 1992), specifically “helpers” to carry the infants which includes fathers.
Experienced fathers are highly motivated to pick up infants, even if the infant is not their own
and are hormonally responsive to the scent of their infants (Ziegler, Prudom, and Zahed, 2009).
The amount of maternal care relative to paternal care varies among the species as does the amount
of care by nonparents, however, in all these species there are substantial amounts of infant care by
individuals other than the mother (Goldizen, 1987). Helpers become an important factor after the
newborn period. Systematic research indicates that helpers of specific age, gender, and experience
levels participate in care of infants of different ages (Price, 1992). In marmosets, fathers are allowed
to carry newborns (Ziegler et al., 2009). In titi monkeys, the mother and father work as a team from
birth. From the first week of life, the father carries the infant more than 70% of the time (Mason
and Mendoza, 1998).
Parenting Infant New World Monkeys

The infancy period in many species of New World Monkeys is relatively short (approximately five
months). Infant capuchins and howlers first break contact with their mother in the second month of
life (Figure 3.7) and spend 50% of their time off the mother when they are 5 months of age (Fragaszy,
1990; Pavé, Kowalewski, Zunino, and Leigh, 2016). In wild settings, howler monkey infants begin
eating solid food around five weeks of age and are weaned around nine months of age (Pavé et al.,
2016). Infant capuchin monkeys explore their environment and spend much of the time off of their
mother, directing visual attention to the activities of the mother (Fragaszy et al., 1991; O’Malley and
Fedigan, 2005). Squirrel monkeys are weaned within the first 3 months of life (Biben,1994) as are
capuchins (Wamboldt et al., 1988). Marmoset and tamarins, however, appear to be weaned around six
months of age. Both adult and adolescent squirrel monkey females carry and play with infants, and
mothers groom their infants regularly (Robinson and Janson, 1987). Cebus infants, 2 months old or
older, are occasionally left with the dominant male who either huddles with them or is tolerant of
their play (Robinson and Janson, 1987). Many primatologists emphasize the commonalties in infant
care patterns within cooperatively breeding species of New World Monkeys, but “variation and
flexibility emerged as major themes” when considering parenting in cooperatively breeding species,
such as tamarins, marmosets, and tit monkeys (Snowden, 1996, p. 682). In some monogamous species,
mothers have primary responsibility for infant care at birth. For these species, however, maternal cra-
dling of newborns does not occur. In fact, New World monkey infants, even at birth, have sufficient
strength and motor maturity to support their own weight and to maneuver to the nipple. However,
the survival rate of infants born to first-time tamarin mothers is quite low (i.e., 10%, Snowden, 1996).
This high mortality rate appears to be due to the first-time tamarin mother doing most of the car-
rying of her twins: experienced mothers typically carry their infants only 50% of the time during
their first week of life.
The infant is able to crawl ventrally to nurse and otherwise rides on the mother’s back. In some
monogamous species, such as marmosets, fathers have primary responsibility for the infant from
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Figure 3.7 Mother and nursing month-old infant capuchin, Cebus capucinus, in the Refugio de Vida Silvestre
Curú, Costa Rica.
Source: Photo credit: Mary Baker.
shortly after birth (Figure 3.8). In titi and night monkeys, fathers carry the infant up to 90% of
the time and the infant forms a strong attachment to the father (Spence-Aizenberg, DiFiore, and
Fernandez-Duque, 2016; Wright, 1990). In these species, mothers only hold infants for nursing.
Fathers, not mothers, groom, play, and share food with their infants. These observations of parental
care in the wild mirror those reported from captive settings (Spence-Aizenberg et al., 2016). Infants
transfer to other carriers for the majority of their time. Initially, the infant’s transfer is facilitated by
the helper by adopting a different posture, “including extension of the arms toward the infant and
direction of the infant’s crawling motions” (Tardif et al., 1992, p. 156).
Infants can be actively rejected by mothers who have insufficient prior experience ( Johnson,
Petto, and Sehgal, 1991; Tardif, Richter, and Carson, 1984), and these mothers show some fear and
avoidance of the newborn presumably due to the lack of prior experience (Pryce, Abbott, Hodges,
and Martin, 1988). Although deaths of newborns occur due to lack of sustenance, most early deaths
appear to be due to falls (i.e., skull fractures found in autopsies of newborn squirrel monkeys; Hopf,
1981). Tamarin, marmoset, squirrel monkey, and capuchin mothers actively reject an infant by rub-
bing the infant off their body. It is for this reason that the presence of helpers is crucial, as they can
collect and carry newborns, or even take the infant before the mother tries to remove it.
Both male and female nonparents act as helpers with equal frequency in cotton-top tamarins, but
which gender helps may be constrained by age. Female subadults carry more than male subadults,
but as adults, males carry more than females. Very young infants are carried more often by adults
compared with subadults, but subadults carry more than juveniles (Yamomato, Box, Albuquerque,
and Arruda, 1996). In some species (red-bellied tamarins) younger, less experienced helpers are not
allowed access to very young infants (Pryce et al., 1988). Weaning is accomplished by the eighth
Figure 3.8 Marmoset fathers (Callithrix jacchus) carry young offspring on their back. The mother sits nearby.
Source: Photo credit: Judith Peterson, with permission by Toni Ziegler, and reprinted with permission from Myowa and
Butler (2017).
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month of life, when mothers have been observed to act aggressively toward infants who attempted
to nurse (Snowden, 1996).
The squirrel monkey pattern of mutual gaze and responsive vocalizing, which occurs between
newborns and reproductively active females, becomes a characteristic pattern between the squirrel
monkey infant and mother when the infant begins to leave mother in independent forays in the
third or fourth week of life (Biben,1994). Infant position, riding on the mothers’ back, influences
the infant’s ability to engage in eye contact with the mother. Constant physical contact in the early
days of life also makes it unlikely that contact-resuming vocalizations will be directed at the mother.
Perhaps these vocal exchanges or dialogues serve socialization purposes, acquainting infants with the
sounds of group members (Biben, 1994).
In captive capuchins, Weaver and de Waal (2002) were able to classify the quality of attach-
ment between mother and infant capuchins. There were significant consequences in the behavior
of infants and juveniles as observed through longitudinal study (Weaver and de Waal, 2003). Infants
and weanlings initiated more reconciliations than they received from adults. Infants with an inse-
cure relationship with their mothers (i.e., above-average amounts of agonistic behavior relative to
affiliative behavior) were highly aroused by conflict with adults, exhibiting distress vocalizations, and
found reconciliation to be comforting. Infants with a secure relationship with their mother (i.e.,
above-average ratios of affiliative relative to mildly agonistic behavior) exhibit a more appeasing style
of reconciliation, imbued with positive friendly overtures to adults that previously aggressed against
them. Thus, infants developed characteristic styles of reconciliation, labeled homeostatic regulation
styles since they involved ways in which infants resolved the distress emerging from social conflict.
Weaver and de Waal (2003) speculated that the attachment relationship facilitated the development
of infant’s homeostatic regulation style, supporting interpretations of a strong biological and physi-
ological influence on infant behaviors (Maestripieri et al., 2009).
A noticeable aspect of paternal care in tamarins, Cebus apella, and Ateles is food sharing. In some
species of tamarins and marmosets, food is shared with infants through both active offering and
tolerated taking. The possessor vocalizes and extends their hand holding the food while maintaining
eye contact with a particular infant, or infants are allowed to take food from the hands or mouth
of the possessor (Feistner and McGrew, 1989). Some males, in some species, are reported to initiate
food sharing with specific vocalizations (Brown and Mack, 1978), but primarily it is the infant who
begs. Older siblings also participate in sharing food with infants. Food sharing is primarily directed
at weanlings. In golden lion tamarins, for instance, up to 90% of the food eaten by infants is obtained
from social partners (Brown and Mack, 1978). But, as in wild chimpanzees, meat is shared equally
across all social partners in wild capuchins (Perry and Rose, 1994).
Fathers will defend infants against intruders and rescue them, at some risk to their own welfare.
Typically, parental caregiving of this sort is given only to very young infants—until infants can loco-
mote independently (Whitten, 1987). Titi and night monkey fathers, however, are reported to guard
their infants for their first full year of life (Wright, 1990). Adult male howler monkeys may carry and
play with infants for short periods of time (Vogt, 1984).
Parenting Juvenile New World Monkeys

In tamarins, marmosets, and squirrel monkeys, the juvenile period begins around five months and
ends at puberty, around 14 months of age (Tardif et al., 1992). In some of these callitrichid species, the
presence of the mother inhibits their daughter’s sexual development (Walters, 1987). In cotton-top
tamarins, juveniles, 6.5 to 15 months of age, react with stress to the birth of new infants. Parents and
juveniles were seen to be more often in conflict. Initially, it was thought that this conflict was due to
competition for parental resources (that is, conflict between the juveniles and the infants). But, when
the parents were not carrying the new infants there was no conflict, so it was clear that the conflict
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was due to competition for the infants. When juveniles were allowed to carry the infants, about four
to six weeks after they were born, all conflicts ceased.
Juveniles take more responsibility for learning and for maintaining the relationship with their
parents. In wild titi monkeys, juveniles, not the parents, are responsible for maintaining proximity
with their parents (Spence-Aizenberg et al., 2016). Juvenile capuchins, more than subadults or adults,
spend the most time watching others process food. Although there is no evidence that they learn
food processing techniques, they may benefit from learning what can and what cannot be eaten.
Experimental studies find that capuchins can learn how different parts of a “plastic food” can be
manipulated or the way in which different parts move (Custance, Whiten, and Fredman, 1999). In
cases where prey was eaten, observation by juveniles preceded begging or scrounging (O’Malley and
Fedigan, 2005).
Parenting Adolescent New World Monkeys

Adolescents play less than juveniles do, and when the newest set of twins is born then adolescent
behaviors become more adult-like (Caine, 1986). Adolescent sons and daughters help with infant
care by grooming and caring for younger siblings. For marmosets and tamarins, adolescence appears
to end at approximately three years (Santos, French, and Otta, 1997). Prior to achieving adult stature,
however, both males and females leave the family. In squirrel monkeys for example, primiparous
females give birth around three years of age (Hopf, 1981), but males are not fully mature before
5 years (Biben, 1992). Adolescent squirrel monkeys continue to play, but play takes on sexual ele-
ments (Caine, 1986). It appears that adult females actively reject adolescent males who are forced to
the periphery of the group (Caine, 1986).
Summary
New World monkeys spend 3 to 5 years as subadults. In some species, there is extensive paternal
involvement and cooperation among family members in carrying offspring and sharing food with them.
Prosimians
Parenting Newborn Prosimians

Some prosimian newborns cling and are carried ventrally (without support) by mothers. In some
species transport of infants is by mouth, and infants are left unattended in nests or left “parked”,
meaning grasping on tree branches (Higley and Suomi, 1986; Klopfer and Boskoff, 1979). In some
prosimian species, mothers may leave their infants unattended for up to 12 hours. Mothers may scent
mark the infants for identification, an indicator that mothers do not visually identify their infants as
individuals (Niemitz, 1979). Prosimians are the most quickly maturing of primate species and reach
sexual maturity by 1 year of age.
Parenting Infant Prosimians

Infant prosimians may ride dorsally on the mother. This is accomplished by 2 weeks of age in ring-
tailed lemurs (Lemur catta) and by 4 weeks in the brown lemur (Lemur fulvus). In the ruffed lemur
(Varecia variegate), however, infants engage in independent excursions away from the mother by 3
weeks of age (Klopfer and Boskoff, 1979). Mothers groom their infants frequently. In contrast to
monkeys, but like chimpanzees, prosimian mothers may play for extended periods of time with their
infants (Charles-Dominique, 1977; Niemitz, 1979). Prosimians, which are the most evolutionarily
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distant from humans, are weaned by 2, 3, or 4 months depending on the species ( Jolly, 1985; Klopfer
and Boskoff, 1979).
The most distant ancestral primate is considered to be the gray mouse lemur, which lives in
Madagascar in mixed sex social groups and is nocturnal. Mothers give birth to one or more infants
(Figure 3.9), and may form sleeping groups with one or two closely related females (Eberle and
Kappeler, 2006). Lactating females nurse cooperatively within these sleeping groups. During the
night, mothers carry their own infants and park them nearby while they forage, mostly solitarily. In
the first 4 to 6 weeks of life, mother carry their infants in their mouth (Eberle and Kappeler, 2006).
Infants emit at least three different vocalizations that appear to be specific to different contexts.
Infant gray mouse lemurs purr when being groomed, emit whistle vocalizations when separated
from their mother, and tend to emit tsak and whistle vocations when they are threatened while
alone (Scheumann, Zimmermann, and Deichsel, 2007). Scheumann et al (2007) suggest that the
vocalizations of gray mouse lemur infants convey both infant emotion and a communicative mes-
sage to the mother: “the mother should perceive purrs as cohesion calls, whistles as attraction calls,
and tsaks as aversive calls” (p. 716). Additionally, infants and mothers engage in greeting calls, for
example, when mothers return to the sleeping box after a period of time away (Scheumann, Linn,
and Zimmermann, 2017).
Prosimian fathers tend not to participate at all in infant care (Vogt, 1984). The involvement of
male ring-tailed lemurs in parenting tends to be limited to occasional grooming or sniffing of the
infant, but if the mother is removed from the group the amount of time that the infant spends in
contact with the male increases (Vogt, 1984). If the gray mouse lemur mother dies, then a close
maternal relative from the sleeping groups may adopt the infant (Eberle and Kappeler, 2006).
Figure 3.9 Littermate gray mouse lemurs, Microcebus murinus, at 5 days of age, have not fully opened their eyes.
Source: Photo credit: Marina Scheumann, Institute of Zoology, University of Veterinary Medicine, Hannover.
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Parenting Juvenile and Adolescent Prosimians

Very little is known about parental influence on older prosimian offspring. Dispersal by sons appears
to be voluntary. Immature male bushbabies may leave by following an adult male when he travels
through the mother’s home range. Daughters remain close to their mothers; even as adults, daughters
sleep close to mothers at night (Charles-Dominique, 1977).
Summary
Prosimians mature most quickly among primates, and many reach full maturity in less than one year
(Table 3.1). Offspring are weaned in 3 to 12 weeks and can have relatively few interactions with
their mother during waking hours as they are often parked in nests while mothers forage. Some spe-
cies have extensive reciprocal vocal interactions between mothers and offspring. As adults, they may
adopt a home range that overlaps with that of their mothers.
Long-Term Consequences of Early Experiences
Effects of Responsive Care Intervention for Nursery-Raised Chimpanzees

Two studies have documented long-term consequences of differential early rearing in chimpanzees.
Whether nursery-raised chimpanzees had developed an attachment relationship with a human car-
egiver was assessed at 1 year of age (van IJzendoorn et al., 2009). Years later, when these chimpanzees
were adults, their health, behavior, and well-being were assessed (Clay et al., 2015). Chimpanzee
infants that did not possess an organized attachment system were found to have more respiratory
ill-health events into adulthood, more stereotyped rocking behavior, and an average level of well-
being, compared to those with an organized attachment system that had fewer adverse health events,
significantly less rocking, and above-average levels of well-being as adults.
The second study investigated differences in the structural connectivity of the gray matter in adult
chimpanzees’ brains based on their early experiences (Bard and Hopkins, 2018), and followed the
advice of Krasnegor and Bridges (1990) to look more closely at how the brain changes in response
to parenting behavior. At birth, chimpanzees were raised by the biological mothers, or, if their moth-
ers had insufficient maternal behavior (Bard, 1994a, 2002), they were placed in a nursery. Most were
given standard nursery care that provided all essential care to meet the infants’ physical needs and
provided same-aged peers to provide for the infants’ social needs. A responsive care intervention was
given to some nursery-raised individuals, however, for 4 hours per weekday, that included focused
efforts on nurturing chimpanzee species-typical motor, social, emotional, and communicative skills
from birth through the first year of life in a sort of intuitive parenting paradigm (Bard, 1994a, 1996;
Bard, Dunbar, et al., 2014). Chimpanzee mothers act responsively, contingently, and nurture develop-
ment in their infants. Of special significance are the intuitive parenting behaviors utilized to support
development in motor skills and in communicative behaviors (Bard, 1994a, 1996; Bard, Dunbar,
et al., 2014; Rijt-Plooij and Plooij, 1987). Intuitive parenting consists of a psychobiological preadapt-
edness to stimulate infants’ integrative development (Papousek and Papousek, 1987). The behaviors
are neither reflexive nor based on rational thought, but they appear to require prior experience with
infants. Intuitive parenting of locomotor skills in monkeys, for example, may have evolutionary ben-
efits, including early cessation of infant carrying, early weaning, and thus, increased likelihood of a
shorter interval to the next birth (Maestripieri, 1995).
Providing positive emotional responsiveness to infant chimpanzees, through intuitive parenting
processes, for as little as four hours per day, had dramatic effects on joint attention, cooperation,
organized attachment (van IJzendoorn et al., 2009), communication (Bard, Dunbar, et al., 2014),
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Kim A. Bard
and emotional expressiveness (increasing positive affect: Bard, Bakeman, et al., 2014). In contrast,
early stress had equally dramatic effects but on different systems—early stress caused deficits in
attention span, goal-directed efforts, emotional expressiveness (increasing fearful responses: Bard
and Gardner, 1996), and impacted the development of lateralized behavior (Bard, 1998a; Bard,
Hopkins, and Fort, 1990; Hopkins and Bard, 1993, 2000). There has been a great deal of research
on the effects of stress and fear, but scientists have not spent equivalent energy in investigating the
effects of positive emotions and attachment relationships (Bennett et al., 2017; Panksepp, 1986;
Sheridan and Bard, 2017).
Standard institutional nursery care for chimpanzees appeared to cause connectivity deficits in
the gray matter of the basal forebrain, dopamine-rich brain regions that in humans involves reward
circuits, and these deficits were prevented with a responsive care intervention (Bard and Hopkins,
2018). The lack of nurturing care of the standard nursery resulted in gray matter volume in the basal
forebrain that was greater than that of the responsive care intervention or the mother-raised group.
A possible cause for larger gray matter volumes would be a concentration of cell bodies related to
relatively fewer dendritic connections.
A special focus of the Bard and Hopkins (2018) study is the impact of the responsive care inter-
vention on brain structures. There were no brain components in which the Responsive Care Inter-
vention differed from the mother-raised chimpanzees. The Responsive Care Intervention, given to
chimpanzee newborns and infants through the first year of life at the Yerkes Primate Center (Bard,
1996), appeared to provide the experiences necessary to prevent the deleterious effects of atypical
rearing on structural connectivity evident in the brains of the Standard Care group.
The nurturing experiences of chimpanzee infants were measured for the responsive care inter-
vention group, the standard care nursery group, and for those raised by their chimpanzee mother, and
were found to differ (Bard and Hopkins, 2018). Nurturing motor development has been observed
in behaviors called “exercising” (Yerkes and Tomilin, 1935), which occur for up to 20% of the time
while mothers are engaged with their very young infants (Bard, 1994a). In addition to nurturing
motor skills, chimpanzee mothers also nurture the development of social and communicative skills
(Bard et al., 2014; Bard, 2017; van Lawick Goodall, 1968). It was surprising to find that the rate of
nurturing experienced by 1-month-old chimpanzees predicted the structural connectivity in the
basal forebrain of their adult brain (Bard and Hopkins, 2018). Thus, the Responsive Care Interven-
tion was an effective intervention that prevents the deleterious effects of atypical rearing on structural
connectivity in the chimpanzee brain.
Long-Term Consequences of Early Separations of Offspring

From Conspecific Caregiving
Rhesus monkeys have most often been the subjects for longitudinal studies, in which early rear-
ing experiences are linked with outcomes in adulthood. Many of these studies can be conceived as
investigating the long-term consequences of stress or adversity early in life (Parker and Maestrip-
ieri, 2011). These primate models have often used the lack of parenting or temporary separations
from caregivers as the stressor. Not surprising, these manipulations result in long-term behavioral,
physiological, immunological, and neurological consequences (reviewed in Parker and Maestripieri,
2011; Suomi, 2004). When infant rhesus monkeys are permanently separated from their mother
under experimental conditions, they can become hyperaggressive adults, and females do not provide
adequate care for their infants (Harlow, 1958). In rhesus, early separations in infancy can compromise
adult immunological responses (e.g., to simian immunodeficiency virus [related to HIV] challenges,
Capitanio and Lerche, 1991). In pigtailed macaques, separations lasting only 10 days in infancy can
result in juveniles and adolescents being deficient in the ability to develop close friendships and social
networks (Capitanio and Reite, 1984).
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Whereas isolation-rearing for the first months of life produces severe deficits in behavior, com-
promised physiology, and permanent changes in neuroanatomy, peer-rearing can have persistent but
less severe effects in rhesus monkeys, but can include problems in maternal behavior in adulthood
(Champoux, Byrne, DeLizio, and Suomi, 1992). It is common, however, for peer-rearing paradigms
to consist of an initial period of isolation (e.g., for the first 30 days of life, perhaps with a surrogate
and blanket provided at day 15) and peer interactions provided after day 30. This type of peer-rearing,
for instance, appears to cause a lowering of the physiological stress set-point (that can be upregulated
by greater peer exposure; Capitanio, Mendoza, Mason, and Maninger, 2005), decrements in the
development of the serotonin system (found as early as 14 days of age and lasting at least through
150 days of age; Shannon et al., 2005), and structural changes in the brain that last, at least, into the
late juvenile period (Spinelli et al., 2009). It is worth noting, however, that these long-term conse-
quences may not be found in other primate species or may not be found if social interaction expe-
riences are given in the first 30 days. Pigtailed macaques, for instance, given nursery rearing with
extensive infant handling by humans, showed no long-term behavioral or social deficits as juveniles
or adolescents (Sackett, Ruppenthal, and Davis, 2002).
Even when separations from caregivers during infancy are more limited in scope, there can still
be long-term consequences. Early separation experiences cause changes in immunology in rhesus
monkeys. Separation experiences in marmosets, involving daily separations from both parents in the
first month of life, result in changes in physiology, impairments in cognitive functioning, and behav-
ioral inhibition (Pryce et al., 2005), and impairments in the function of genes in the hippocampus
related to synaptic plasticity (Parker and Maestripieri, 2011). It is noteworthy that separations from
the mother later in life, especially around the ages when offspring are normally becoming independ-
ent, can serve to enhance resilience, at least in squirrel monkeys (Parker et al., 2004).
Maternal abuse has clear immediate and short-term consequences for the primate infant, ranging
from distress to injury and death. Abused infants cry more as a direct consequence of abuse, and they
cry more than nonabused infants, even when they are not being abused (Parker and Maestripieri,
2011). Abuse increases the infant’s tendency to cling to the mother, which may paradoxically cause
more abuse. Abused infants are developmentally delayed in initiating play with peers, and they play
less (Maestripieri and Carroll, 2000; Reite, 1987).
Stress experienced early in life may induces changes in emotional reactions to stress, and these
changed responses may last a lifetime (Suomi and Levine, 1998). Emotional behavior, mediated by
the limbic system, results in emotional reactions that are easily transferred to other individuals and
to the next generation (e.g., phobic reactions; Mineka et al., 1984). The magnitude of long-term
disturbance in adults could be related to the magnitude of the stress response in infancy. Many separa-
tion paradigms involve both the loss of caregiving behaviors and distress caused by that loss; thus, the
independent contribution of each variable is difficult to determine. One advantage of investigations
into the consequences of normal variation in maternal behaviors is that it might disentangle these
variables.
Long-term consequences for infants and juveniles of being raised with different “maternal styles”
have been studied. In general, protectiveness is thought to prevent potential dangers, including har-
assment from others (Maestripieri, 1994). Infant and juvenile vervet monkeys with more protective
mothers are more cautious in response to novelty (Fairbanks and McGuire, 1993). Although these
infants do have less risk from predation, they appear to be less able to cope with stressors, such as the
loss of the mother (Hinde, 1987). Maternal style can include differences in abuse or rough handling,
typically reflected in how often offspring experience “rejection”. Rejecting vervet mothers have
infants that show more enterprise (if they survive): The infants develop independence at an earlier
age and are more resourceful (Fairbanks, 1996). Rhesus macaques that experienced more maternal
rejection early in life were more socially avoidant as adolescents (Maestripieri et al., 2006). Adoles-
cent male vervets with more rejecting mothers were bolder with adult male strangers (Fairbanks,
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Kim A. Bard
1996). For daughters, it appears that the mothers’ maternal style is the best predictor of their own
maternal style with their offspring (Berman, 1990: Fairbanks, 1996). Variability in maternal style has
long-lasting consequences in the social behavior and physiology of offspring (for more details see
review by Parker and Maestripieri, 2011).
Infant temperament and the maternal style experienced early in life interact in the attainment of
dominance status, which is a measure of social success. High-reactive rhesus monkey infants raised
with nurturant mothers attain high status in an adolescent peer group, whereas high-reactive rhesus
monkey infants raised with punitive mothers are lowest in dominance status (Scanlan, 1986). Sch-
neider (1984) found that high-reactive nursery-reared rhesus infants and low-reactive mother-reared
infants score highest on cognitive assessments when tested as juveniles even though there were no
overall group differences based on rearing or on reactivity. In chimpanzees of the Tai Forest of Côte
d’Ivoire, West Africa, there appears to be differential maternal investment in sons and daughters,
based in part on maternal dominance rank (Boesch, 2012). Mothers of high rank have a longer
period of time between the birth of a son and the next offspring—which decreases the mortality of
sons of high-ranking mothers, whereas mother chimpanzees of low rank tend to invest more in their
daughters (Boesch and Boesch-Achermann, 2000). Moreover, through their continuing support,
mother chimpanzees of high status aid their sons in achieving high dominance status as well (Boesch
and Boesch-Achermann, 2000).
Conclusions
There is no single “primate pattern” of parenting. Diversity, variability, and flexibility are among the
most important characteristics of primate parenting (Hawkes et al., 2017). There is a strong genetic
basis for maternal behavior, but an equally strong influence of experience. It would be a mistake to
expect any single variable to have an exclusive determination for parenting: “Maternal behavior is
obviously so important to the survival of a species that it has been ‘overdetermined’—that is, driven
by multiple behavioral and physiological systems” (Coe, 1990, p. 178). There is a danger in drawing
explicit comparisons between a particular primate species and a particular human society (Myowa and
Butler, 2017). Perhaps every facet of human parenting can be found to occur in some other primate
species, yet clearly no one other primate species exhibits a complete repertoire of human parenting.
Maternal competence in chimpanzees, expressed in interaction even with very young infants,
reflects sensitive responsivity during which the mother engages in contingent behavior and encour-
ages development of infant capacities (Bard, 1994a). These behaviors parallel those observed in intui-
tive parenting in humans (Papousek and Papousek, 1987). For chimpanzees, being raised with their
biological mother provides a myriad of benefits, some of which can be provided by humans with a
focus on nurturing the development of species-typical skills (Bard, 1996; Bard and Hopkins, 2018;
Clay et al., 2015; van IJzendoorn et al., 2009).
There is not a single theoretical account, barring evolutionary theory, that can explain the diver-
sity of parenting patterns in primates (Hawkes et al., 2017). Moreover, there is not a single theoretical
account that explains or predicts the diversity of patterns of infant interactions with individuals other
than the mother (Caine, 1993; Chism, 2000; Maestripieri, 1994; Manson, 1999; Paul, 1999; Ross and
MacLarnon, 2000; Silk, 1999; Snowden, 1996; Wright, 1990). It is likely that there will not be a single
theoretical account that explains the ontogeny of maternal competence in primates, because there is
a similar pattern of diversity in maternal behavior across species. For some species, such as tamarins,
mothers must learn to let others help in infant care. Rhesus mothers must learn to allow infants to
cling to their hair. Chimpanzee mothers must learn to provide support for the infant, and nurture
their physical, emotional, and communicative development.
Parenting in primates is diverse. Some Prosimian mothers park their infants in nests, carry them
in their mouth, and let them nurse and otherwise do not engage with their infants. In contrast,
110
chimpanzee mothers provide cradling support and spend up to 15 minutes each waking hour inter-
acting socially, communicatively, or didactically with their infants. Some New World monkey fathers
play an important role in parenting infants, as do gorilla fathers. Consideration of the influence of
monogamy or living in a harem illuminates the manner in which social organization contributes
to behavioral expressions of parenting. Cognitive influences on parenting, especially evident in the
Great Apes, typically provide for flexibility and richness in parenting behaviors, making great ape
parenting more similar to human parenting than that of other primates. The evolutionary risks,
however, include greater dysfunction in parental behavior when learning environments are altered.
The continuing influence of parents on older offspring, especially adolescents, requires more focused
study. The manner in which independence is achieved is also an important consideration. Evidence
of teaching complex locomotor behaviors, tool use, hunting, and subtle food searching patterns
suggests that primate parents, at least of some species, continue to influence offspring throughout
development (Boesch, 2012; Goodall, 1986; Lonsdorf, 2006). As we develop new primate models of
parenting, it will be important to consider emotional components of parenting with an emphasis on
the positive aspects of bonding and attachment (Keller and Bard, 2017), and on investigations of the
mechanisms underlying the reinforcing qualities of parenting (Panksepp, 1998). There is much that
remains to be learned.
Acknowledgments
Funding was provided by NIH Grant RR-00165 to the Yerkes Regional Primate Research Center
of Emory University, NIH Grant RR-03591 to R. B. Swenson of the Yerkes Center, NICHD Intra-
mural Research Program funds through the Laboratory of Comparative Ethology and S. J. Suomi, a
Max-Planck Society stipend in cooperation with the Developmental Psychobiology project directed
by H. Papousek at the Max-Planck Institute for Psychiatry, NICHD-NRSA Research Fellowship
HD-07105 to K. A. Bard, and NIH Grant RR-06158 to K. A. Bard. The Yerkes Center is fully
accredited by the American Association for Accreditation of Laboratory Animal Care. I thank my
research assistants, especially Kelly McDonald, Yvette Veira, Kathy Gardner, and Josh Schneider,
for all their work with the chimpanzees, students at the University of Portsmouth, including Chris
Brindle, Nadja Dreschler, Kriztina Ivan, and Judit Barta, and I thank Ms. N. Johns, my friend and
Librarian of the Yerkes Research Center. Special appreciation is extended to Marc H. Bornstein,
Steve Suomi, David Leavens, Mechthild Papousek, and to the memory of Hanus Papousek for their
intellectual contributions to this chapter.
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4
GENETICS AND PARENTING
Introduction
Parenting is a dynamic, multiply determined process that is influenced by a range of extra-familial
(e.g., cultural context) and intra-familial factors (e.g., interparental relationships) (Bornstein, 2016).
Generally, researchers have been interested in parenting as the means by which the environment
shapes the developing child. A number of theoretical models of how parents influence their chil-
dren (e.g., socialization, modeling) have been generated, with these models typically assuming that
parenting operates via social transmission of behaviors. Yet, the ability of children to shape their
own environments has been understood for nearly five decades (Bell, 1968), and within biologically
related families, parents have not only an environmental influence, but a genetic one as well. Thus,
unless explicitly modeled, associations between parenting and child outcomes may be due, in part,
to confounds from unmeasured heritable influences. It may be, for instance, that children’s heritable
characteristics evoke certain types of parenting, such as when parents respond negatively to adoles-
cents’ aggression (Narusyte, Andershed, Neiderhiser, and Lichtenstein, 2007). Alternatively, relations
between parenting and child outcomes may be due to shared genetics between parents and children.
For example, parents who are high in irritability may be more likely to use harsh parenting strategies
and to have children who are also irritable (Prinzie et al., 2004).
The aim of this chapter is to discuss the “genetics” of parenting by outlining how genetically
informed designs can provide important information for parenting research. Genetically informed
designs refer to a class of designs that measure genetic influences either by assessing family members
of varying relatedness (e.g., twins, full siblings, adoptive parents and adopted children) and explicitly
modeling relatedness or by directly collecting DNA. These designs (e.g., twin studies, adoption
studies) allow us to test an assumption underlying many theoretical models of parenting—that asso-
ciations between parent and child characteristics are due to environmental influences as opposed to
heritable influences, facilitating the further refinement of existing theories and ultimately informing
intervention efforts (Harold, Leve, and Sellers, 2017; Leve et al., 2017). Moreover, some genetically
informed approaches are beginning to identify how children evoke parenting at the level of the gene
(Elam et al., 2017).
We begin with a review of the significant changes that have occurred over the past century to
make behavioral genetics the field that it is today—one that includes examination of parenting. Next,
we describe various genetically informed research designs and discuss the utility of each. Then, we
summarize extant literature of the genetics of parenting during infancy and early childhood (ages
123
birth–6 years), middle childhood (ages 6–10 years), and adolescence (ages 10–18 years). We end with
a discussion of the ways in which the work of developmental scientists can inform that of behavioral
geneticists and vice versa.
Historical Considerations in Genetics and Parenting

The field of behavioral genetics has seen rapid growth since the middle of the 20th century, espe-
cially in regard to the types of constructs examined (Knopik, Neiderhiser, DeFries, and Plomin,
2017). Broadly, behavioral genetics research can be categorized into family-based approaches, which
capitalize on varying degrees of genetic relatedness within families to quantify relative contributions
of heritable and environmental influences on traits, and molecular genetic approaches, which aim to
identify the specific genetic underpinning of behavioral traits. Family-based genetic studies include
twin, extended family, and adoption designs and allow for heritable and environmental influences
to be disentangled by estimating heritability as well as shared environmental (i.e., nongenetic influ-
ences that account for similarity among family members) and nonshared environmental influ-
ences (i.e., nongenetic influences that account for differences among family members). All of these
designs, as well as combinations of the designs, can also help to clarify gene-environment inter-
play such as gene-environment correlations (rGE) and gene by environment interactions (GxE).
Although we will discuss each of these aspects of gene-environment interplay in detail in the sec-
tions that follow, it is important to note that genes and environments can systematically co-occur
(as in rGE) and can moderate one another (as in GxE) because it has implications for understanding
the processes involved in development. Molecular genetics studies use approaches such as candi-
date gene, genome-wide association studies (GWAS), and polygenic risk scores (PRS) to pinpoint
a gene or set of genes that explain variance within a phenotype. These studies can also examine
gene-environment interplay, although GxE is more commonly examined within molecular genetic
frameworks than is rGE.
Although humans have been thinking about genetics for centuries, in 1960, Fuller and Thompson
published Behavior Genetics, which was effectively the first comprehensive handbook of the study of
the genetic basis of behavioral traits. At that point, genetics research had largely focused on Mende-
lian inheritance. During the 19th century, Gregor Mendel studied pea plants and showed that both
members of the parent generation of pea plants provide a component (later called a gene) to the
offspring (Knopik et al., 2017). He measured observable traits (i.e., phenotypes) of the plants, such
as whether the seeds were wrinkled or smooth, and noticed that variation in offspring phenotypes
could be reliably predicted based on the parent generation. In other words, a single gene determined
these characteristics, and this work provided the basis of heredity, which refers to the degree to which
1969
1977
2002
1859
Bandura’s
1960 Social Plomin, Publicaon of
Darwin’s On the Learning Purcell’s GxE
Origin of Theory DeFries, &
Loehlin’s Models
Species Publicaon of 1991
Published 1940s Behavioral rGE paper
Genecs 1970s Early 2000s
1865 1983 Plomin &
Early Bergemen’s GE
Adopon interplay paper Early candidate
Baumrind’s
Mendel Studies gene work
Parenng Scarr &
presents his Styles McCartney’s
Laws of
rGE paper
2000
1860
1960
1970
1980
Inheritance
1990
Figure 4.1 Historical timeline of notable behavioral genetics and developmental events.
124
Genetics and Parenting
differences within a population on a phenotype can be attributed to differences on a genotype. Some

human behaviors or diseases, like Huntington’s disease or phenylketonuria, are caused by single
genes, but complex human behavioral traits (e.g., personality, psychopathology) are not attributable
to any single gene (Knopik et al., 2017). Complex traits can be better understood by studying the
degree to which they are explained by genetics.
Even in 1960, when contemporary scientists rejected the notion that biology played any role
in human behavior, behavioral geneticists acknowledged that behavior is not influenced by purely
genetic or environmental factors (i.e., “the Nature-Nurture Problem”), but instead, results from
complex processes that involve both genetic and environmental factors (Fuller and Thompson, 1960).
In a seminal adoption study, Heston and Denney (1968) compared adopted offspring whose birth
mothers had been diagnosed with schizophrenia with adopted offspring who had no maternal his-
tory of schizophrenia. Schizophrenia was only found in the offspring of biological mothers with
schizophrenia, lending evidence to genetic causes of schizophrenia. Previously, psychologists believed
that poor parenting caused schizophrenia (Fromm-Reichmann, 1948). Subsequent adoption studies,
such as the Texas Adoption Project (TAP; Horn, 1983; Horn, Loehlin, and Willerman, 1979) and
the Colorado Adoption Project (CAP; DeFries, Plomin, Vandenberg, and Kuse, 1981; Plomin and
DeFries, 1983) expanded both the parent and offspring characteristics studied. For instance, TAP
examined children ages 3 through 16 years and found that children’s intelligence was more similar
to their birth mothers than to their adoptive parents (suggesting heritable influences; Horn, 1983),
and CAP extended those findings downward to infancy, identifying genetic contributions to infant
intelligence and communicative competence, and provided preliminary evidence that the association
between maternal responsivity and infant intelligence is due to both heritable and environmental
influences (Plomin and DeFries, 1983).
The decades following the publication of Behavioral Genetics saw growth in both theory and
methodology. Researchers who were interested in human behavioral genetics at the time Behav-
ioral Genetics was published were limited by the methodological and statistical techniques available.
Geneticists who studied nonhuman populations had the benefit of experimental designs to test the
effects of manipulating genetic or environmental parameters, whereas the field of human behavioral
genetics relied heavily on correlational research designs that consisted of family members of varying
degrees of genetic relatedness. Advanced statistical tools, such as structural equation modeling, were
developed and subsequently applied to genetically informed designs, enabling scientists to estimate
the relative heritable and environmental variance underlying the covariation of multiple phenotypes
(Eaves and Gale, 1974; Martin and Eaves, 1977).
During the late 1970s and early 1980s, researchers began thinking about gene-environment inter-
play. Two early twin studies found heritable effects on adolescent twins’ reports of accepting and
rejecting parenting—a construct that had been previously assumed to be a measure of the environ-
ment (Rowe, 1981, 1983). Rather than considering heredity and environment as wholly independ-
ent forces, the study of gene-environment (GE) interplay is based on the assumption that, under
certain circumstances, heritable and environmental factors are systematically related due to rGE and
GxE (Plomin, DeFries, and Loehlin, 1977). In other words, heritable factors can control, at least in
part, which environments are experienced and how responsive an individual is to an environment
(Scarr, 1992; Scarr and McCartney, 1983). Passive rGE is present when children are exposed to
certain environments because of their parents’ inherited characteristics, such as a child of anxious
parents being exposed to more overprotective parenting. Evocative (also called reactive) rGE occurs
when children’s heritable characteristics evoke reactions from their environment, such as an anxious
child eliciting more over-protective parenting in stressful situations. Active rGE refers to children
choosing their environments based on inherited characteristics, such as an inhibited child partaking
in more solitary activities. Conceptually, evocative and active rGE are distinct, but in extant research
designs, they are difficult to differentiate. Additionally, rGE does not account for the entirety of the
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variance in a phenotype, but rather is one possible explanation for links between parent and child
characteristics.
In contrast to rGE, GxE refers to processes in which the effects of the environment vary as a
function of genetics or instances in which the environment modulates the expression of an indi-
vidual’s genotype. An example of GxE is a child with a predisposition for surgency who thrives in
the context of appropriately structured parenting but struggles in the context of overly permissive
parenting. GxE can be thought of in different ways based on whether the environmental construct
under consideration is conceptualized as negative/risky or positive/protective (Burt, 2011). When
the environment is risky, an inherited vulnerability may be amplified or activated (Burt, 2011). For
instance, at increasing levels of paternal punitive discipline, heritable influences on adolescent exter-
nalizing behaviors increase (Button, Lau, Maughan, and Eley, 2008). Also when the environment is
risky, an inherited vulnerability may be diminished, which is known as the bioecological interaction
model (Burt, 2011; Pennington et al., 2009). A finding consistent with this model is that heritable
influences on adolescent aggression are lower, and shared environmental influences are higher, in dis-
advantaged neighborhoods compared to advantaged ones (Tuvblad, Grann, and Lichtenstein, 2006).
Finally, when the environment is protective, the effects of an inherited vulnerability may be dimin-
ished, as in the compensation by social context model (Burt, 2011). For instance, at increasing levels
of parental warmth, heritable influences on adolescent antisocial behaviors are diminished (Feinberg,
Button, Neiderhiser, Reiss, and Hetherington, 2007). Notably, GxE can occur in the context of rGE
although consideration of these two types of GE interplay simultaneously is rare ( Jaffee, 2016; Jaffee
and Price, 2007; Price and Jaffee, 2008).
In light of evidence that parenting constructs are themselves subject to heritable influences
(Rowe, 1981, 1983), Plomin and Bergeman (1991) argued that genetic influences on measures
that had historically been considered “environmental”—such as parenting or family climate—do
not necessarily mean that environments themselves are heritable. Instead, they suggested that such
measures capture something about the individual’s relations with the environment that are due to
unmeasured, heritable characteristics. Parenting, then, could be considered a phenotype that is influ-
enced by both parents’ and children’s genetically influenced characteristics. More than a decade after
Plomin and Bergeman (1991) put forth this idea, Rutter (2005) pushed the field to engage in more
rigorous testing of purported environmental mechanisms of risk. In the intervening years, a number
of “environmental” risk factors (e.g., harsh parenting) had been explored using genetically informed
designs, but Rutter highlighted a number of steps required to strengthen the case that these are causal
mechanisms. In particular, he stressed the importance of longitudinal designs that include precise
measures of purported risk factors (and outcomes), that allow for testing of competing hypotheses,
and that explicitly test the theoretical limitations of the specific design (e.g., equal environments
assumption for twin studies, restricted range of environments for adoption designs). In many ways,
he argued for the field of behavioral genetics to adopt practices that were already in place in typical
developmental research, while encouraging developmental researchers to incorporate methodology
practiced by behavioral geneticists.
This theoretical shift that began in the early 1980s necessitated changes in research strategies
and methodological advances to allow the investigation of specific developmental processes of GE
interplay. For one, more fine-grained assessments of family processes were included in family-based
genetic designs. Parent-offspring adoption studies, by design, are well-suited to test GxE, but the
available data from some early studies were generally too vague to specify the inherited “risk” that
was being captured, due in part to the fact that many early studies examined adult adoptees. Meth-
odologically, GxE in parent-offspring adoption designs can be tested by statistically modeling the
interaction between constructs of birth parents (genetic influences) and constructs of adoptive par-
ents (environmental influences) to predict the offspring outcome of interest. One example of this
strategy was implemented by Cadoret, Yates, Troughton, Woodworth, and Stewart (1995) who used
126
hospital and prison records to diagnose the biological parents of adopted adults and also assessed the
adoptive family for marital, psychiatric, and legal problems. Adoptive family problems were related
to increased adoptee aggression, but only if the biological parent exhibited antisocial personality dis-
order. This study and others helped to establish evidence of GxE (Cadoret, Troughton, O’Gorman,
and Heywood, 1986; Wahlberg et al., 1997), but the mechanistic processes remained unexplored.
Registry data continue to be used and are critically important because studies using registry data are
able to examine the large sample sizes needed to detect small effect sizes; to understand the mecha-
nisms of effects, in-depth assessment of family process is also necessary. We describe findings from
parent-offspring adoption studies that have collected more in-depth assessments of family processes
thus helping to clarify processes by which parents and children influence one another.
A second important change involved methodological advances that provided straightforward
strategies for examining GxE within the context of both twin designs and molecular genetic designs
(Purcell, 2002; Purcell and Sham, 2002). Prior to the early 2000s, two approaches were used to test
GxE within a twin study. One required samples of twins in which co-twins were discordant on
exposure to discrete environmental constructs (e.g., adult twins discordant on marital status). For
instance, twins discordant on marital status were compared, and evidence suggested that being mar-
ried mitigated genetic liability for depression (Heath, Eaves, and Martin, 1998). The other approach
required instances in which co-twins (or whole samples) were exposed to vastly different levels of
a continuous environmental construct (e.g., twins who experienced high versus low levels of life
stress). Using this approach, Heath and colleagues (1985) compared twins who were educated prior
to the institution of liberal educational policies in Norway with those who were educated following
policy changes. The first cohort showed little evidence of heritability for educational attainment,
whereas following these policy changes, heritable factors played a substantial role in educational
attainment.
Thus, until the early 2000s, researchers using twin designs to investigate GxE were relegated to
examining characteristics or experiences that were discrete or allowed for the creation of distinct
groups. This design is reasonable for discrete factors pertinent to the study of parenting (e.g., paren-
tal divorce, daycare attendance), but many other relevant factors are continuous (e.g., socioeco-
nomic status, parental hostility). Furthermore, many statistical models of GxE assume that the
latent genetic factors are independent of measured environmental constructs—an assumption that
is violated in the context of rGE ( Jaffee and Price, 2007). To address this limitation, Purcell (2002)
developed a method to test GxE in twin samples if the moderator is continuous, and subsequently
revised the method to test GxE in the presence of rGE (Purcell and Sham, 2002), sparking a
number of analyses that would not have been possible previously. Now, continuous moderators,
such as those indexing parenting and broader family characteristics, could be tested in a twin GxE
framework, providing more information about developmental processes. One subsequent report
found that, for example, as family dysfunction increases, genetic contributions to conduct disorder
decrease, while shared and nonshared environmental influences increase (Button, Scourfield, Mar-
tin, Purcell, and McGuffin, 2005).
It is important to note that the model developed by Purcell (2002) and used in hundreds of stud-
ies since is not purely a strategy for testing GxE, but is rather more general. Specifically, this model
tests for moderation of heritable, shared environmental, and nonshared environmental influences by
a continuous variable, most often an “environmental” variable like parenting or socioeconomic status.
We would argue, however, that moderation of the shared and nonshared environmental estimates is
equally informative as moderation of heritability, especially when the focus is on understanding the
mechanisms of gene-environment interplay. For example, using an twin/sibling design, adolescent
temperament moderated heritable and shared environmental influences on parental negativity (Gan-
iban, Ulbricht, Saudino, Reiss, and Neiderhiser, 2011). The heritable influences on parental negativ-
ity were greatest at higher levels of adolescent negative emotionality, and both shared and nonshared
127
environmental influences were smallest. In other words, as adolescents exhibit more temperamental
negative emotionality, the child-based effects on parental negativity increase resulting in less-similar
parental negativity in sibling pairs.
Concomitant to advances in family-based approaches, the field of molecular genetics experienced
rapid growth with major initiatives such as the Human Genome Project (Collins and Mansoura,
2001), the International HapMap Project (Gibbs et al., 2003), and the 1000 Genomes Project (Siva,
2008) that have begun to characterize the human genome. The proliferation of research in psychi-
atric molecular genetics is reflected in the rapid increase in human molecular genetic publications,
from about 2,000 published between 2000 and 2004 to 9,400 published between 2010 and 2014
(Ayorech et al., 2016). Arguably, a catalyst of this growth was the publication of two early studies
demonstrating candidate gene-by-environment interaction (cGxE). The first found that adults who
were maltreated as children were likely to develop antisocial behavior as adults but only if they were
carriers of the neurotransmitter-metabolizing enzyme monoamine oxidase A (MAOA) gene coding
for low expression (Caspi et al., 2002). The second concluded that stressful life events were more
strongly linked to depression for carriers of two short alleles of serotonin-transporter-linked poly-
morphic region (5-HTTLPR) (Caspi et al., 2003).
This methodological approach made genetic work more accessible to nongeneticists and allowed
for theories like diathesis-stress and differential susceptibility to be tested in a molecular genetic frame-
work. However, subsequent attempts at replication of those particular findings have been inconsistent
(Culverhouse et al., 2017), and general concern has been raised about the validity of published cGxE
studies (Dick et al., 2015). Multiple meta-analyses (Culverhouse et al., 2017; Karg, Burmeister, Shed-
den, and Sen, 2011; Munafò, Durrant, Lewis, and Flint, 2009; Risch et al., 2009; Sharpley, Palanisamy,
Glyde, Dillingham, and Agnew, 2014) testing the initial cGxE interaction between 5-HTTLPR and
life stress on depression have themselves produced conflicting results, with some finding no evidence
of an interaction and concluding that if one exists, it is likely a small effect that is not generalizable
across groups (Culverhouse et al., 2017). The 5-HTTLPR and life stress interaction is an illustrative
example of the challenges posed by widespread interest in cGxE work, but it is by no means unique.
To address these problems, the field needs to adopt standards and practices that increase the meth-
odological rigor of cGxE work (Dick et al., 2015).
As we have reviewed, the field of behavioral genetics has seen major growth in theory and
method since the publication of the field’s namesake text in 1960. We next describe contemporary
approaches for studying parenting from a behavioral genetics perspective.
Research Designs and Methodology in Genetics and Parenting

We first describe the logic of family-based genetic designs (e.g., twin, adoption studies) before
discussing molecular genetic methodology. The foundation of family-based genetic approaches is
heritability, which refers to the extent that individual differences on a phenotype are attributable
to inherited differences. Estimates of heritability can be obtained by comparing the phenotypes
(i.e., measured behaviors) of individuals who vary in degree of genetic relatedness but provides no
information about phenotypes at the individual level or about the specific inherited factors that
influence a phenotype. For instance, heritability estimates of parental warmth range from 34% to
37%, indicating that 34%–37% of the population variance in parental warmth is due to heritable
influences (Kendler and Baker, 2007). By combining evidence from multiple research designs, we
can better understand GE interplay and the mechanisms of effects on child development. Below, we
further describe the most commonly used family-based behavioral genetic designs (twin, adoption,
extended family, and combination), outline the basic assumptions and limitations of these designs
and clarify how each can inform our understanding of GE interplay. We discuss family-based genetic
approaches before turning to a discussion of molecular genetic approaches.
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Twin Designs
Twin studies, as shown in Figure 4.2, compare monozygotic twins (MZ) with dizygotic twins (DZ)
and are particularly well-suited for detecting heritable effects. MZ twins share 100% of their segre-
gating genes, while DZ twins share, on average, 50% (like any full sibling pair). Twins are also the
same age and often the same sex (DZ twins can also be different sex, although many twin studies use
only same-sex twin pairs). Thus, if MZ twins are more similar to each other on a particular construct
than DZ twins, the best explanation for this similarity is that the construct is heritable. To illustrate,
within a sample of 8-year-old twin children, the intraclass twin correlations for child-rated anxiety
symptoms are .72 for MZ twins and .20 for DZ twins. Because the DZ twin correlations are less
than half that of the MZ twin correlations, heritable influences on child anxiety are indicated (Eley,
Napolitano, Lau, and Gregory, 2010).
Beyond identifying the heritability of a trait, genetically informed designs allow variance of a phe-
notype to be partitioned into additive genetic (A), shared environmental (C), and nonshared environ-
mental (E) influences. Shared environmental influences refer to nongenetic influences that account
for similarity among family members, such as residing in the same home. Nonshared environmental
influences refer to nongenetic influences that account for differences among family members, such
as having separate peer groups or differential parenting, and also include measurement error. This
approach does not allow for identification of the specific environmental influences acting on parent-
ing but highlights the importance of both heritable and environmental factors. Multivariate analyses
go a step further by incorporating multiple phenotypes.
Particularly relevant to twin studies is the assumption of equal environments, which states that the
effects of the environments on MZ twins and DZ twins are equivalent with respect to the construct
under study. It could be violated, perhaps, if parents of MZ twins systematically treat their children
more similarly than do parents of DZ twins and this more similar treatment results in greater MZ twin
similarity for the construct under study. This assumption has been empirically tested and supported
(Conley, Rauscher, Dawes, Magnusson, and Siegal, 2013; Eaves, Foley, and Silberg, 2003); however, if it
Child-Twin Based Design Parent-Twin Based Design

Parents
Twin Parent 1 Twin Parent 2
Child Twin 1 Child Twin 2

Child 1 Child 2
A = Influence of child’s genecally influenced A = Influence of parents’ genecally influenced
characteriscs on parents’ parenng. characteriscs on parenng. Interpreted as
Interpreted as evocave rGE passive rGE
C = Influence of children’s shared C = Influence of parents’ shared environmental
environmental characteriscs on parenng. characteriscs on parenng.
Interpreted as passive rGE E = Influence of parents’ unique environmental
E = Influence of child’s unique environmental characteriscs on parenng.
characteriscs on parenng.
Figure 4.2 Child and parent-based twin designs and gene-environment correlation.
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were to be violated, genetic estimates would be inflated. An early study used a small sample of “misclas-
sified” twins (i.e., MZ twins whose parents believed them to be DZ twins and vice versa) to investigate
this assumption (Scarr, 1968). If parental beliefs about zygosity impact similarity of treatment, misclas-
sified MZ twin correlations would resemble those of correctly classified DZ twins and misclassified
DZ twin correlations would resemble those of correctly classified MZ twins. By contrast, if genetic
relatedness impacts similarity of treatment, misclassified MZ twin correlations would resemble those
of correctly classified MZ twins, and misclassified DZ twin correlations would resemble correctly clas-
sified DZ twins. The latter was found for all examined measures, including maternal report of twins’
current behaviors, temperament and personality characteristics, and developmental milestones. In other
words, parents’ similar treatment of their children is attributable to actual genetic similarity rather than
parents’ perception of twin similarity. This finding strengthens the conclusion that, when twin studies
find parenting of MZ twins to be systematically more similar than DZ twins, it is not a violation of
the equal environments assumption but a reflection of heritable influences on parenting. Finally, it is
important to keep in mind, that the equal environments assumption is relevant only in relation to how
MZ twins’ more similar experiences influence the construct being examined.
Twin designs can be used to investigate GE interplay and particularly, by using combinations of
twins-as-parents and twins-as-children designs, it is possible to disentangle passive and evocative/
active rGE. The distinction between these two types of twin designs in regard to parenting is impor-
tant because they lead to different interpretations of heritable influences on parenting. Parent-based
designs (in which the twins are parents) answer the question how do parents’ genotypes influence their
parenting? Studies using these designs that find heritable influences on parenting constructs can be
inferred as passive rGE, even when child constructs are not explicitly modeled because passive rGE
occurs when parents’ inherited characteristics influence their parenting. Within child-based designs
(in which the twins are children/adolescents), shared environmental influences refer to nongenetic
factors that account for siblings’ similarity, including parents’ heritable influences on their parenting.
Taken together, when there is evidence of heritability for parenting within a parent-based design and
shared environmental influences within a child-based design, the case for passive rGE is strengthened.
Child-based designs, by contrast, are focused on how children’s genotypes influence their parents’ behavior.
When twins are children, heritability on measures of parenting suggest that parents are responding to
their children’s inherited characteristics (evocative rGE). Different research designs, then, can provide
complementary information about GE processes operating within families.
Adoption Designs
Adoption studies are particularly useful for testing parenting as an environmental mechanism because
similarities between adopted children and nonbiological adoptive parents cannot be due to shared
genes (Haugaard and Hazan, 2003) especially when the child is placed in the adoptive home at or
near birth. The basic adoption design is displayed in Figure 4.3 and includes data collected from birth
parents, adoptive parents, and adopted children and compares correlations between biological parents
and adopted children with correlations between adoptive parents and adopted children. If the adopted
child and biological parent(s) correlate for a given phenotype, heritable effects are indicated, but if the
adopted child correlates with adoptive parent(s) or with the rearing environment, there is evidence of
environmental influences (Haugaard and Hazan, 2003). As Figure 4.3 shows, birth mothers provide
children with both their prenatal environment and half of their genes, so by including birth fathers,
prenatal influences can be disentangled from genetic influences (Loehlin, 2016). Beyond comparing
correlations, the adoption design has been used with more complex modeling techniques (e.g., struc-
tural equation modeling) to study evocative rGE. When birth parent characteristics are related to
adoptive parent parenting, there is initial evidence for evocative rGE because the child is presumably
the factor linking the two (Ge et al., 1996; Leve et al., 2007). By subsequently testing child factors as
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Genec Environmental
Influences Influences
Birth Birth Adoptive Adoptive

Father Mother Father Mother
Prenatal
Influences
Adopted
Child
Figure 4.3 Traditional adoption design.
potential mediators between birth parent and adoptive parent characteristics (Harold, Leve, Barrett,
et al., 2013), the mechanisms by which evocative rGE is operating can be more clearly specified.
One criticism of adoption studies is that adoptive families are not generally representative of all
families and therefore may represent a selected range of environments (Stoolmiller, 1999). Because
adoptive families are screened prior to adoption, adoptive parents typically are older and better edu-
cated than nonadoptive parents and consequently may have higher incomes and more education. In
other words, there are concerns that adoptive families are systematically different from nonadoptive
families and that these differences reduce the generalizability of findings from these studies. If this
were the case, the restricted range could increase the likelihood that shared environmental effects will
be underestimated. However, comparisons of adoptive and nonadoptive families have not supported
this criticism and have shown that factors like parental anxiety and depression, family functioning,
and peer group exposure are roughly equivalent for adoptive and nonadoptive families (Leve et al.,
2007; McGue et al., 2007). In addition, for the constructs that do differ for adoptive and nonadop-
tive families (e.g., socioeconomic status [SES], parental antisocial personality disorder, and substance
dependence), those differences are not related to child behavioral outcomes (McGue et al., 2007;
Stoolmiller, 1999). A second possible limitation of adoption studies is that, if children are placed
with adoptive parents based on certain characteristics (e.g., parental education), correlations between
adoptive parents and adopted children could be inflated and consequently misinterpreted as envi-
ronmentally mediated. This phenomenon is referred to as “selective placement” and can be empiri-
cally tested by correlating birth parent characteristics with adoptive parent characteristics; if they are
uncorrelated, there is no evidence of selective placement. Selective placement has been assessed in
adoption studies on the basis of characteristics such as personality, temperament, cognitive ability,
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and financial needs, and no evidence of systemic selective placement has been found (DeFries et al.,
1981; Leve et al., 2013).
Combination Designs
Although twin and parent-offspring adoption designs are the backbone of family-based genetic
research, variations on these strategies help to refine the questions that can be addressed or increase
the generalizability and/or power of the approaches. Sibling designs can be used in much the same
way as twin designs (D’Onofrio, Lahey, Turkheimer, and Lichtenstein, 2013). Specifically, full siblings
(i.e., children who have the same biological mother and father) are as genetically similar as DZ twins,
sharing 50% of their segregating genes on average, although they are different ages and did not share
their intrauterine environment. Half-siblings, who share 25% of their segregating genes, and steps-
iblings, who share none of their segregating genes, are often present in families, especially because
of the incidence of divorce and remarriage. In the same way that twin studies compare correlations
between MZ and DZ siblings, sibling designs compare correlations between sibling pairs (e.g., full,
half, and biologically unrelated siblings). Similarly, sibling adoption designs can be used in two differ-
ent ways. More commonly, genetically unrelated siblings living in the same homes can be examined.
These genetically unrelated siblings may be the result of two unrelated children being adopted into
the home or of one child being adopted and another biological to the parents. A second type of
sibling adoption design comprises genetically related (full or half ) siblings living in separate homes.
In other words, one sibling is adopted, and the other child is parented by the biological parent(s).
If correlations for biologically unrelated siblings reared in the same home are significant, research-
ers can conclude that shared environmental influences are operating (Deater-Deckard, Petrill, and
Wilkerson, 2003), whereas correlations for biologically related siblings reared apart would bolster
conclusions that genetic influences are at play (D’Onofrio et al., 2013).
Children-of-Twins (COT) studies provide strong tests of causality between parent and offspring
phenotypes by including data from twin parents and the offspring of both twins (D’Onofrio et al.,
2003; McAdams et al., 2014). Children of MZ twin pairs share exactly 50% of their genes with their
parent (Twin 1) and with their aunt or uncle (Twin 2), while children of DZ twins share just 25%
of their genes with their aunt or uncle, like any other avuncular relationship. Yet the rearing envi-
ronment of children of MZ twins is not shared as it is provided by different individuals. Similarly,
the children of MZ twins share 25% of their genes with their cousin (like half-siblings), whereas
children of DZ twins share 12.5% of their genes with their cousin, like any other cousin pair. This
extended family design provides many combinations of family member correlations that can be
compared to test specific hypotheses about genetic and environmental influences. Pertinent to the
study of parenting, however, is that COT designs control for genetic and environmental factors that
make biologically related family members living in the same household similar. Thus, by comparing
parent-offspring correlations with avuncular correlations (i.e., the correlation between aunts/uncles
and nieces/nephews) for MZ and DZ twin families, COT designs can test the extent to which
parent-offspring associations are attributable to shared genetic influences (passive rGE) or within-
family environmental influences. In other words, if there is greater similarity for children of MZ
twins than children of DZ twins and the parent-offspring correlations are similar to the avuncular
correlations for MZ twin families, then passive rGE would be indicated. If parent-offspring correla-
tions are greater than avuncular correlations, associations between parent and child phenotypes are
attributable to rearing environmental factors.
An Extended Children-of-Twins (ECOT) design can more clearly distinguish among the effects
of passive rGE, evocative rGE, and direct environmental influences because it can test bidirectional
effects between parents and children (Narusyte et al., 2008). As shown in Figure 4.4, ECOT models
add a child-as-twin design to the parent-as-twin design of a COT, essentially combining two separate
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Child-Twin Based Component Parent-Twin Based Component

1.0 1.0/0.5
1.0 1.0
A1 C1 E1 E1 C1 A1 A1 C1 E1 E1 C1 A1 A = genetic influences
C = shared environmental
influences
E = nonshared environmental
influences
Parenting Parenting Parenting Parenting m=direct effect of parenting

on Twin 1 on Twin 2 by Twin 1 by Twin 2
s=passive rGE
0.5 0.5
0.5 m m 0.5 m m n=evocative rGE
n n n n
Child of Child of
Twin 1 Twin 2
Twin 1 Twin 2
Phenotype Phenotype
Phenotype Phenotype
s s s s
A1 A1 A1 A1
’ ’ ’ ’
A2 C2 E2 E2 C2 A2 A2 C2 E2 E2 C2 A2
1.0
1.0/0.5 0.25/0.125
Figure 4.4 Extended Children-of-Twins design.
but compatible samples (comparable on key demographic characteristics) to increase the power to
detect child-specific genetic effects (McAdams et al., 2014). The COT portion of the design meas-
ures the parenting of twin parents and therefore can model the effects from parents to offspring (i.e.,
passive rGE or direct environmental), and the child-twin portion of the design measures comparable
constructs from parents of twin children and can model the effects from offspring to parent (i.e.,
evocative rGE; Narusyte et al., 2008). To date, this powerful design is the only one in which passive
rGE, evocative rGE, and direct environmental influences between parenting and offspring pheno-
types can all be modeled simultaneously.
Summary
Together, family-based genetic approaches rely on the varying degrees of genetic relatedness and
shared rearing environment (or lack of a shared rearing environment) between individuals that natu-
rally occur within families. Beyond identifying whether traits are heritable, these designs can identify
and/or control for confounding genetic influences that account for similarity among biologically
related family members to provide strong tests of environmental causation. Child- and parent-based
designs (including twin and extended family designs) can provide evidence for passive and evocative
rGE, adoption designs are well-suited for detecting evocative rGE and environmental influences,
and more complex models parse passive rGE from environmental influences (COT) and passive and
evocative rGE from direct environmental influences (ECOT). Therefore, the use of multiple geneti-
cally informed designs allows for stronger conclusions to be reached about links between parenting
and child characteristics both in regard to how genes and environments may work together and for
the role of environmental effects “free” from the effects of heredity.
Molecular Genetic Approaches

Although molecular geneticists use a number of approaches (e.g., linkage association, candidate gene,
and genome-wide association studies), we limit our discussion to approaches that have been used
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to study parenting, namely, candidate gene and polygenic risk score approaches. These approaches
incorporate genotypic data known interchangeably as genetic variants or genetic markers, referred
to here as variants. Genetic variants are selected because they index known or purported functional
segments of the gene that may be causally related to the phenotype of interest. When these seg-
ments vary across individuals, they are referred to as polymorphisms. Genetic variants include single
nucleotide polymorphisms (SNPs; variation in a single nucleotide base pair), variable number tan-
dem repeats (VNTRs; variation in the number of repeated base pairs) in addition to other, less com-
mon genetic variants that will not be discussed here (Griffin, Schlomer, Cleveland, and Vandenbergh,
in press). To illustrate, a study that helped launch widespread interest in candidate genes examined
5-HTTLPR, known colloquially as the serotonin transporter polymorphism (Caspi et al., 2003). The
5-HTTLPR polymorphism is a well-studied VNTR in the promoter region of the SLC6A4 gene.
This variant exists as two alleles, long and short (16 and 14 repeats, respectively; Lesch et al., 1996),
and the short allele causes low expression of the serotonin transporter (Bradley, Dodelzon, Sandhu,
and Philibert, 2005). Individuals with at least one copy of the 5-HTTLPR short allele (s/s and s/l,
known as “short carriers”) are compared to homozygous long individuals (l/l).
The basic approach of candidate gene-by-environment interactions (cGxE) involves genotyping
individuals and statistically testing the interaction between selected genetic markers and measured
environmental characteristics on a phenotype of interest (Dick et al., 2015). This method has been
employed to test child DNA as a marker of vulnerability or susceptibility to particular parenting
behaviors. However, cGxE studies face a number of criticisms and have been difficult to replicate
(Duncan and Keller, 2011). cGxE work requires large (i.e., greater than 1000) sample sizes to be suf-
ficiently powered and studies have to failed to replicate previous research that used small to moder-
ate sample sizes (Duncan and Keller, 2011; Karg et al., 2011; Risch et al., 2009). Furthermore, many
published cGxE studies omit rGE entirely. Much like family-based genetic approaches, molecular
genetics designs can be used to assess gene-environment interplay including rGE and GxE ( Jaffee
and Price, 2007). When parents’ genetic markers are related to their parenting behaviors, it suggests
passive rGE, whereas when child genetic markers are systematically related to parenting behaviors via
child behaviors, researchers can infer evocative rGE. Certain genotypes may be linked with increased
exposure to certain environments and significant cGxE findings may, in reality, be capturing rGE
processes. The presence of rGE can be assessed by testing the direct association between genotype
and environment; if they are correlated, methods must be employed to control for rGE before testing
for GxE (Dick et al., 2015).
One criticism of candidate gene work is that, ultimately, any single genetic variant accounts for
only a tiny fraction of the variance in a complex phenotype. To address this criticism studies have
used polygenic risk scores, which are sums of multiple genetic markers used to index genetic liability.
To identify genetic variants of interest, both biologically based and data-driven approaches are used
(Dudbridge, 2013; Griffin et al., in press; Nikolova, Ferrell, Manuck, and Hariri, 2011). Biologically
based scores rely on extant literature to inform the selection of genes relevant to neurotransmit-
ters (e.g., dopamine and serotonin) and biochemical pathways. Data-driven approaches like GWAS
employ large sample sizes to identify genetic markers associated with the phenotype of interest.
Polygenic risk scores have been used to test for evocative rGE at the molecular genetic level while
controlling for passive rGE and as we review below, and suggest that, for the same PRS, there is
evidence of both passive and evocative rGE depending on the parenting constructs examined (Elam
et al., 2016, 2017).
Molecular genetic approaches have been used to study correlates of parenting at the level of
the gene. Candidate gene studies of parenting aim to identify genetic variants associated with par-
enting behaviors or, when genes are measured from the child, to identify individual differences
in susceptibility to environmental factors, such as parenting. Early candidate gene work sparked
widespread attention from genetic and nongenetic researchers, but concerns about the validity of
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these studies have followed difficulty with replicability, culminating in a number of journals, such as
Behavior Genetics, Drug and Alcohol Dependence, and the Journal of Abnormal Child Psychology, establish-
ing stringent criteria for the publication of such studies (Hewitt, 2012; Johnston, Lahey, and Matthys,
2013; Munafò and Gage, 2013). Another response to this issue has been the development of novel
approaches, such as polygenic risk scores, that include multiple genetic variants used to disentangle
passive and evocative rGE in a way that is complementary to statistical genetic approaches used in
family-based genetic research.
In the sections that follow, we review literature that uses the genetically informed designs outlined
above to better understand the etiology of parenting. We segment this discussion by developmental
period (i.e., infancy and early childhood, middle childhood, and adolescence). Within each section,
we first discuss work on the heritability of parenting, then review findings about the genetic influ-
ences underlying links between parenting and child behaviors (“genetics on associations between
parenting and child outcomes”), and finally examine evidence of GxE. While much of the extant lit-
erature relies on family-based genetic approaches, relevant findings from molecular genetic research
are interspersed throughout the discussion.
Parenting During Infancy and Early Childhood

Developmental studies of infancy and early childhood (birth–6 years) have typically highlighted
characteristics such as parental sensitivity, hostility, and secure parent-child attachment as crucial for
shaping children’s development. To date, very few twin studies have examined parenting in infancy.
The Leiden and London Twin Studies consisted of infant twins and their families recruited from
a twin registry in the Netherlands (Leiden) and from hospitals in the United Kingdom (London).
Both studies measured parenting when infants were approximately nine months old and found
that maternal sensitivity was explained by shared environment, with little evidence of heritability
(Fearon et al., 2006). The same general pattern was found in the Quebec Newborn Twin Study
(QNTS), which recruited a representative sample of families living in the greater Montreal area and
assessed twins beginning at 5 months of age (Boivin et al., 2005). When the infants were 5 months
old, mothers’ self-efficacy, perceived parental impact, and overreaction were all attributable to large
shared environmental influences. Only mothers’ hostile cognitions towards their infants were herit-
able when assessed when the infants were 5 months old and again when the twins were 30 months
old (Forget-Dubois et al., 2007). Together, these studies suggest that, for parents of infants, positive
parenting characteristics may be less due to child-driven heritable effects. There is, however, a need
for more research on parenting of infants using genetically informed designs, especially that includ-
ing negative parenting of infants.
During early childhood, twin and family studies have found evidence for heritability in some
aspects of parenting with positive parenting constructs tending to be less heritable than negative
parenting constructs. The Twin’s Early Development Study (TEDS) is a representative sample of
twins in the United Kingdom who were assessed repeatedly beginning at age 2 (Oliver and Plomin,
2007). Findings from TEDS have shown that parental positivity is modestly heritable when twins are
3 and 4 years old with the bulk of variance explained by shared environmental influences. Parental
negativity was explained primarily by heritable influences, with the rest of the variance explained
by both shared and nonshared environmental influences (Alemany, Rijsdijk, Haworth, Fañanás, and
Plomin, 2013; Knafo and Plomin, 2006; Larsson, Viding, Rijsdijk, and Plomin, 2008). In contrast, a
study using data from 4-year-old children and their next oldest sibling (including full, half, and steps-
iblings) participating in the Avon Longitudinal Study of Parents and Children (ALSPAC) found no
heritability for maternal negativity with variance due only to equal amounts of shared and nonshared
environmental influences, whereas mother report of their partners’ negativity toward the children
was mainly heritable with some significant shared environmental influences (Deater-Deckard, Dunn,
135
O’Connor, Davies, and Golding, 2003). The differences in findings for these two studies could be
due to differences in measurement or to differences in the samples. For example, twin studies like
TEDS are especially powerful for detecting heritable effects, while studies that include step or adop-
tive siblings like ALSPAC provide additional power for detecting shared environmental effects but
are limited in their power to detect heritable effects due to the absence of MZ twins.
Studies that measure parenting of young children observationally as well as via parent reports have
shown that estimates of heritable and environmental influences also vary based on the method of
measurement—generally, observational measures are less heritable than parent-reports, and estimates
of shared environmental influences tend to be higher when parenting is measured observationally
than when assessed via parent-report. For example, the TRACKS twin study, a study of 3-year-old
twins and their primary caregivers who were recruited from central and southern England, explored
the heritability of observed maternal behaviors and mother-reported parenting (Deater-Deckard,
2000). Overall, variance in observed parenting behaviors (i.e., positive and negative affect, harsh
discipline, positive and negative control, and responsiveness) was primarily attributable to shared
environmental influences with heritability estimates essentially zero. In comparison, maternal report
of parenting (i.e., positive and negative affect) was largely heritable with some variance due to
shared environmental influences. The Early Childhood Longitudinal Study–Birth cohort (ECLS-B),
a nationally representative sample of same-sex twin pairs from the United States who were recruited
at 9 months of age (Bethel, Green, Kalton, and Nord, 2005) has shown that variance in observed
parenting supportiveness when twins were 24 months old was largely explained by shared and non-
shared environmental influences (Roisman and Fraley, 2008). These findings may reflect that various
measurement approaches capture different aspects of parenting. For example, in child-based designs,
shared environmental influences on observed parenting may be due to shared rearing environment
accounting for twin/sibling similarity, but it may also suggest passive rGE (i.e., parent’s genes influ-
encing their parenting behavior).
A handful of child-based twin studies has generally found that variation in mother-infant attach-
ment is largely explained by shared environmental influences, suggesting passive rGE. This was the
case for twins in the Leiden and London Twin Studies assessed in infancy with the Strange Situ-
ation, as well as twins from the ECLS-B assessed at 24 months of age with an attachment Q-Sort
(Bakermans-Kranenburg, van IJzendoorn, Bokhorst, and Schuengel, 2004; Fearon et al., 2006; Rois-
man and Fraley, 2008). However, data from the Louisville Twin Study, a longitudinal study of infant
twins who were assessed beginning in the 1970s, did not find shared environmental influences to
be important for their measure of attachment. In this case, archival videos recorded when the twins
were 24 months old were coded using a measure of security that is similar to the Strange Situation.
The heritability estimate for attachment security was 25%, and the rest of the variance was attribut-
able to nonshared environmental influences (Finkel and Matheny, 2000). This difference may be due
to measurement differences, in that the Louisville Twin Study used a procedure in which co-twins
were in the room for much of the procedure and were occupied with an activity during the separa-
tions. Moreover, Fearon and colleagues (2006) observed mothers from the Leiden and London Twin
Studies interacting with their infants and found that the covariation between maternal sensitivity
and infant attachment security was attributable to shared environmental influences. The same pattern
was found for families in the ECLS-B; the covariance between attachment and parenting quality was
explained by shared environmental influences (Roisman and Fraley, 2008).
Using a candidate gene approach, some studies have investigated whether maternal geno-
type is associated with mothers’ sensitive behaviors towards their young children. Together, two
dopamine-related genes, catechol-O-methyltransferase (COMT), dopamine D4 receptor (DRD4),
and daily hassles each independently explained variance in maternal sensitivity (van IJzendoorn, Bak-
ermans-Kranenburg, and Mesman, 2008). In the same sample, maternal oxytocin receptor (OXTR)
and 5-HTTLPR were also associated with maternal sensitivity (Bakermans-Kranenburg and van
136
IJzendoorn, 2008). However, a separate study found no association between maternal dopamine
receptor D2 (DRD2) and maternal sensitivity (Mills-Koonce et al., 2007). These studies are subject
to some of the criticisms of candidate gene work that we highlighted above, including a reliance on
small sample size.
Summary of Genetic and Environmental Influences on Parenting

During Infancy and Early Childhood
The paucity of twin studies examining parenting during this developmental period which is char-
acterized by rapid developmental changes, means that there is no clear consensus about the rela-
tive importance of heritable, shared, and nonshared environmental influences in infancy and early
childhood. However, some conclusions can be drawn. First, negative aspects of parenting tend to
be more heritable than positive aspects. Second, parenting assessed via observational measures tends
to show evidence of shared environmental influences, whereas self-report parenting measures are
typically heritable. Third, studies of parent-infant attachment generally detect shared environmental
influences, which could be interpreted as passive rGE. Finally, candidate gene research has begun to
explore whether parents’ genetic variants are associated with maternal sensitivity during this period.
Associations Between Parenting and Child Characteristics

A surprising number of genetically informed designs have been used to explore links between par-
enting and infant and early childhood characteristics. As such, a breadth and depth of evidence have
been amassed, with evidence for passive and evocative rGE as well as support that parenting is associ-
ated with child outcomes above and beyond shared heritability.
Family-based designs have demonstrated that positive parenting qualities (e.g., positivity, warmth,
reinforcement) are related to child behaviors via both heritable and nonheritable pathways (Boeldt
et al., 2012). Within the TEDS sample, for instance, longitudinal associations across early childhood
between parental positivity and child prosocial behavior were attributable to shared environmental
influences (Knafo and Plomin, 2006). Data from the TRACKS study has shown that preschoolers’
cognitive ability and task orientation were related via shared environmental influences, which were
mediated via SES and maternal warmth. Specifically, children from affluent families experienced
increased maternal warmth, which in turn was linked to better cognitive ability and task orientation
(Petrill and Deater-Deckard, 2004). Another form of positive parenting behavior, facilitative reading,
was examined in the Northeast-Northwest Collaborative Adoption Project (N2CAP). N2CAP is
a national survey of adoptive families and an in-depth assessment of a subsample of families, with
a focus on socioemotional and cognitive development (Deater-Deckard et al., 2003). Within this
study, adoptive parents’ reading behaviors with their children (e.g., time spent on books) were related
to child literacy skills, suggesting that parents’ reading-related behaviors (i.e., shared environment)
facilitates literacy independent of heritable influences on literacy which were not estimated in this
study (Petrill, Deater-Deckard, Schatschneider, and Davis, 2005).
Much of the information about parenting and its correlates in infancy and early childhood has
emerged from a parent-offspring adoption study, the Early Growth and Development Study (EGDS;
Leve et al., 2007, 2013). EGDS is a longitudinal, prospective parent-offspring adoption design that
includes adoptive and birth families who were first assessed when adopted children were infants.
EGDS is well-suited to test evocative rGE and GxE. For example, positive parenting behaviors have
been linked to child outcomes across early childhood in a way consistent with a social transmis-
sion model. In one example, adoptive parents were observed interacting with their toddlers using a
clean-up task at child age 18 and 27 months (Natsuaki et al., 2013). Bidirectional longitudinal effects
between parenting and child social wariness indicated more directive behavioral commands (i.e.,
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structured parenting) from adoptive fathers at age 18 months were associated with a decline in child
social wariness from 18 to 27 months. In different reports using the same sample, adoptive parents’
use of structured parenting and maternal reports of higher levels of warmth were associated with
fewer behavior problems in children (Leve et al., 2009; Reuben et al., 2016). Finally, adoptive moth-
ers who were observed using positive reinforcement during a clean-up task had toddlers who later
exhibited lower levels of callous-unemotional behaviors (Hyde et al., 2016). Taken together, findings
from EGDS support rearing environment having a positive impact on child social and emotional
development.
Across infancy and early childhood, negative parenting characteristics have generally been studied
more than positive parenting in behavioral genetic research. Rather than investigating GE interplay
as discussed above, some studies have used an MZ twin differences approach to control for genetic
confounds within families, enabling them to test whether parenting and early childhood character-
istics are associated within families for nongenetic reasons. In an MZ twin differences approach the
focus is on differences between the twins within the twin pair. This allows for direct estimation of
nonshared environmental influences for MZ twins who were reared together. Therefore, if within
MZ twin pairs, the twin who receives higher levels of negative parenting also exhibits higher levels
of the child construct of interest (e.g., externalizing behavior) than the co-twin and if this occurs
systematically across families, one can conclude that nonshared environmental factors influence the
association between parenting and child characteristics. Using this method with a subsample of
twins from the kindergarten cohort of the ECLS study (ECLS-K), parental withdrawal was related
to lower child self-control (Wright and Beaver, 2005). This approach has also been employed with
the Environmental Risk Longitudinal Twin Study (E-RISK), a child-based study of twins recruited
from England and Wales when the twins were 5 years old that is notably one of the few twin studies
oversampled to include high-risk families (based on maternal age at twins’ birth). Within families,
differences in maternal negativity directed towards each twin were linked to differences in behavior
problems (Caspi et al., 2004). In other words, differences within twin pairs’ environmental experi-
ences explained the association between maternal negativity and child behavioral problems. The
twin-difference method was also used with Japanese preschool-aged twins recruited from the Tokyo
Twin Cohort Project (Ando et al., 2013). Twins were assessed longitudinally at 42 and 48 months
and the best-fitting model indicated that differences in authoritative parenting were associated with
differences in child peer problems at each time of assessment, but not longitudinally (Yamagata et al.,
2013). Thus, within families, children who received more authoritative parenting had fewer peer
problems at each time and this was not attributable to heritable characteristics.
Family-based studies that explicitly model heritable and environmental influences have shown
that negative parenting and child characteristics are linked via passive and evocative rGE. Within the
QNTS, the covariation between maternal hostile cognitions and infant difficultness was explained
by genetic influences with no evidence of shared or nonshared environmental contributions to this
covariation (Boivin et al., 2005). Similarly, the covariation between parenting negativity and child
behavioral and emotional problems was explained by common heritable influences for ALSPAC
families (Deater-Deckard, et al., 2003) and for TEDS twins (Alemany et al., 2013) with no evidence
for shared or nonshared environmental contributions to the covariation. Because these are herit-
able effects within a child-based design, these findings can be interpreted as evocative rGE. The
TRACKS twin study of preschoolers found that parental negative affect and control were linked to
child externalizing behavior problems (Deater-Deckard, 2000) with different patterns of heritable
and environmental contributions to the covariance as a function of reporter. For observed measures,
parenting and child behaviors were associated via shared environmental influences, whereas heritable
influences explained these associations for parent-report (Deater-Deckard, 2000).
Data from N2CAP and the TRACKS twin study, Deater-Deckard, Ivy, and Petrill (2006) dem-
onstrated that, for children ages 3–8 years, associations between maternal harsh discipline and child
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externalizing behaviors was moderated by maternal warmth. The inclusion of both study samples
allowed the researchers to test for passive rGE by comparing results in biologically related and adop-
tive families. Passive rGE would be indicated if harsh discipline and child externalizing behaviors
were more strongly linked for biologically related mother-child pairs. Because this was not the case,
the authors concluded that passive rGE was not at play, but this design does not allow for evocative
rGE and direct environmental causes to be disentangled. A separate study found the covariation
between parents’ use of corporal punishment and oppositional behavior in children was primarily
due to heritable influences (suggesting evocative rGE) in the E-RISK sample ( Jaffee et al., 2004).
However, the same was not true for the association between oppositional behavior and maltreatment;
this link was primarily explained by shared and nonshared environmental influences. Thus, chil-
dren’s heritable characteristics appear to evoke negative punishment, but child maltreatment does not
appear to be elicited or evoked by the child’s heritable characteristics. A separate report of E-RISK
families found that when fathers were engaged in caregiving during early childhood and exhibited
high levels of antisocial behavior themselves, their children had the highest levels of behavioral
problems; this was independent of the heritable risk of antisocial behavior ( Jaffee, Moffitt, Caspi, and
Taylor, 2003). In other words, antisocial fathers’ presence in their children’s lives is an environmental
influence that contributes to child behavior problems above inherited risk.
A series of papers has examined parenting and child behavior longitudinally in TEDS. Knafo and
Plomin (2006), for instance, found that heritable influences explained the longitudinal association
between parental negativity and child prosocial behavior from preschool into middle childhood.
Follow-up cross-lagged approaches have shown that, after controlling for stability over time, there is
evidence for bidirectional effects between parenting and child behaviors that are attributable to both
heritable and environmental effects (Lysenko, Barker, and Jaffee, 2013). For instance, “parent-driven”
effects from parental negativity towards preschoolers to middle childhood antisocial behavior were
primarily environmentally mediated, but the opposite “child-driven” cross-lagged effects between
preschoolers’ antisocial behavior and parental negativity measured in middle childhood were medi-
ated through heritable influences (Larsson et al., 2008). Similarly, parental negativity in the preschool
years was linked to adolescent behavior problems, which was due to both heritable and shared
environmental factors (Alemany et al., 2013). There were also effects of preschool child behavior
problems on increased parental negativity when twins were adolescents, and this was attributable to
heritable factors.
Longitudinal work in EGDS has shown that negative parenting behaviors are associated with
children’s heritable characteristics, suggesting evocative rGE. For example, infants whose birth par-
ents had elevated psychiatric symptoms or higher levels of reward dependence evoked more negative
parenting from both adoptive mothers and fathers (Fearon et al., 2015; Hajal et al., 2015). Elam and
colleagues (2014) found that birth mother low social motivation was related to child low social moti-
vation, and, in turn, those children experienced more hostility from their adoptive parents, suggest-
ing evocative rGE. Similarly, toddlers whose birth mothers had more antisocial behavior symptoms
had adoptive fathers whose negative parenting increased from 18 to 27 months (Klahr et al., 2017).
Finally, birth mother and adopted child attention deficit hyperactivity disorder (ADHD) symptoms
were indirectly associated via hostile parenting (Harold, Leve, Barrett, et al., 2013). Specifically, chil-
dren with an inherited “risk” for ADHD symptoms were more impulsive which evoked adoptive
mothers’ hostile parenting. In turn, hostile parenting was associated with later child ADHD symp-
toms. Because adoptive mothers and children are not biologically related, these paths are interpreted
as evidence of evocative rGE.
Other reports from the EGDS have demonstrated that parenting behaviors are related to changes
in child characteristics, for what appear to be rearing environmental reasons. Lipscomb and col-
leagues (2011) found a main effect of adoptive parents’ overreactive parenting on child negative
emotionality at 9 months of age and on increases in negative emotionality between 9 and 27 months.
139
Similarly, there were direct links between adoptive parents’ harsh parenting at 9 months and toddler
anger and frustration at 18 months, and adoptive parents’ hostile parenting was related to changes in
toddler’s aggressive behavior from 18 to 27 months and from 27 months to 4.5 years (Rhoades et al.,
2011; Stover et al., 2012, 2016). During the infant and toddler period, parents have direct environ-
mental influences on child emotional development.
Some candidate gene research has examined preschoolers’ characteristics as mediators between
their genetic variants and the parenting they receive. In an innovative approach of candidate gene
work within a child-twin design, Pener-Tessler and colleagues (2013) observed mothers interacting
with their preschoolers, collected genetic data from both mothers and children, and asked mothers
to rate their children’s self-control. The authors examined the serotonin transporter gene polymor-
phism and the inclusion of the twin design allowed them to test within- and between-family effects.
There were only significant results for families with male twins. Within families, 5-HTTLPR geno-
type was related to both self-control and positive parenting. Short allele carriers exhibited more
self-control and experienced more positive parenting, which is somewhat inconsistent with prior
work that showed that the short allele is related to lower self-control under certain circumstances
(Beaver, Ratchford, and Ferguson, 2009). Between families, the maternal 5-HTTLPR genotype was
only related to positive parenting (short allele carriers exhibited lower levels of positive parenting)
and when maternal genotype was added to the full model, child genotype remained a significant
predictor of self-control. In other words, because child genotype was linked to self-control even
when maternal genotype was accounted for, it strengthened the authors’ conclusion that evocative,
rather than passive, rGE was operating. Competing models were tested to evaluate whether child
self-control statistically mediated the relation between genotype and positive parenting or whether
parenting mediated the relation between genotype and child self-control. The first model (child
self-control mediating the link of child 5-HTTLPR to positive parenting), but not the second
(positive parenting mediating link of child 5-HTTLPR to child self-control), was supported. Thus,
for preschool boys, self-control appears to be a heritable behavior that evokes differential positive
parenting within families (Pener-Tessler et al., 2013). Other work in singleton preschoolers suggests
that child negativity mediates the relation between child OXTR and parent confidence (Kryski,
Smith, Sheikh, Singh, and Hayden, 2014) and between child DRD2 and parental support (Hayden
et al., 2010).
Summary of Associations Between Parenting and Infancy

and Early Childhood Characteristics
Child-based designs have shown that parenting during this developmental period is linked to child
characteristics via a number of pathways. Positive parenting and young children’s behavior has
received less attention than negative parenting but appears to be linked via both passive rGE processes
and direct environmental paths. Links between negative parenting and child characteristics have been
attributed to both passive and evocative rGE, as well as to nonheritable sources. Notably, multiple
methodological approaches have been used to examine this developmental period and the findings
bolster the assertion that parenting and child characteristics are associated for nonheritable reasons,
suggesting causality. Within a longitudinal adoption design, for instance, positive parenting behaviors
have been systematically linked to decreases in maladaptive child behaviors, whereas negative parent-
ing behaviors have been associated with increases in maladaptive child behaviors, suggesting direct
effects whereby parenting can “dampen” or amplify the effects of heritable risks. Other longitudinal,
cross-lagged work has shown that negative parenting behaviors are related to child outcomes many
years later via environmentally mediated pathways, whereas child behaviors are linked to later parent-
ing via heritable pathways. Finally, research in molecular genetics explores child behaviors as media-
tors between children’s genetic variants and the parenting they receive.
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Gene x Environment Interaction

Evidence of GxE using family-based genetic designs in infancy and early childhood has almost exclusively
come from the EGDS. First, there are instances of inherited risk (as measured by birth parent characteris-
tics) related to child characteristics but only in the context of parenting. Birth mother sociability was only
related to adopted child social competence in middle childhood when adoptive mother responsiveness
during toddlerhood was low (Van Ryzin et al., 2015). In the same vein, only at low levels of observer-
rated adoptive mother positive reinforcement during toddlerhood was birth mother antisocial behavior
related to adopted child callous-unemotional traits measured 9 months later (Hyde et al., 2016). In other
words, inherited risk indexed by birth mother antisocial behavior is linked to child callous-unemotional
traits only for children who experience low levels of adoptive mother positive reinforcement.
The EGDS has also found evidence that some adoptive parenting characteristics are related to child
outcomes only in the context of inherited risk. Infants who experienced low adoptive mother respon-
sivity exhibited higher levels of toddler fussiness, but only if their birth mother had a history of major
depression (Natsuaki et al., 2010). Children whose birth mother reported higher negative affect exhib-
ited greater levels of negative emotionality as toddlers when they experienced low adoptive parent
overreactive parenting in infancy (Lipscomb et al., 2012). Fathers’ child-centered parenting in infancy
is related to higher levels of toddler social inhibition, but only for children whose birth parents experi-
ence more anxiety symptoms (Brooker et al., 2016). For children whose birth mother had a history
of social phobia, adoptive mother responsiveness in toddlerhood was linked to lower levels of toddler
anxiety symptoms measured 9 months later, when the children were 27 months old (Natsuaki et al.,
2013). Finally, Leve and colleagues (2009) found evidence of an interaction in which structured parent-
ing during infancy buffered the effects of birth mother psychopathology on toddler behavior problems
but amplified child behavior problems in the context of low levels of birth mother psychopathology.
cGxE work in young children has largely focused on genetic markers purportedly associated
with children’s sensitivity to environmental inputs. One of those markers, DRD2, has received such
attention as a moderator of the relation between parenting and child outcomes. For instance, children
who carried the DRD2 A+ allele and had mothers who were more sensitive exhibited fewer affec-
tive problems (Mills-Koonce et al., 2007). The same pattern was found for the link between parental
intrusiveness and preschoolers’ clinical symptoms (e.g., anxiety, depression; Hayden et al., 2010).
DRD2 has also been shown to moderate the association between parenting and the development
of children’s physiologic responses. Propper and colleagues (2008) assessed infants’ respiratory sinus
arrhythmia (RSA) withdrawal—an indicator of stress regulation—at three occasions over their first
year of life. For A+ allele carriers only, high maternal sensitivity was linked to more change in RSA
withdrawal over time, suggesting better regulation (Propper et al., 2008).
Summary of Gene x Environment Interaction During Infancy

and Early Childhood
For this age range, family-based GxE work has primarily used the EGDS. Findings from that study
have supported models in which parenting can activate or amplify inherited risks as well as models
in which parenting is linked to maladaptive child behaviors but only for children with particular
inherited vulnerabilities. From a molecular genetic viewpoint, certain genetic variants are associated
with child sensitivity to parenting “inputs”.
Summary of Parenting During Infancy and Early Childhood

Family-based genetic studies of parenting during infancy and early childhood have primarily used
child-twin or adoption designs. Evidence has accumulated to show that many aspects of parenting
141
are influenced by their own heritable characteristics and that children’s heritable characteristics also
evoke certain parenting behaviors, together underscoring the importance of controlling for family-
level heritable confounds when studying the effects of parenting. However, parenting and child
characteristics are also linked via nonheritable pathways, strengthening the case of key assumptions
of developmental theories that parenting causally impacts child development. A separate line of work
has shown that certain combinations of inherited risk and experienced parenting characteristics lead
to different developmental outcomes, highlighting that parenting has the potential to amplify inher-
ited risk and that inherited risk may make children particularly susceptible to negative parenting.
Molecular genetic studies have taken a few different approaches to examining links between
genetic variants and parenting. Researchers have begun to explore mothers’ genetic variants that
are associated with their own parenting sensitivity and have identified some candidate genes that
may warrant further study (e.g., COMT, DRD4, OXTR, and 5-HTTLPR). Other work has exam-
ined children’s genetic variants (e.g., 5-HTTLPR) that may underlie behavioral qualities (e.g., self-
control) that in turn evoke particular parental responses. Finally, cGxE work with young children has
examined DRD2 as a marker of children’s sensitivity to their environment and has shown that A+
allele carriers show adaptive outcomes in the presence of sensitive parenting compared to noncar-
riers. At present, polygenic risk scores have not been incorporated into studies of parenting during
this developmental period.
Parenting During Middle Childhood

For parents of children in middle childhood (roughly ages 6–10 years), developmentalists have high-
lighted qualities like warmth and control as relevant for children’s development. Much like infancy
and early childhood, most studies of parenting have used child-based designs and have given more
attention to negative parenting behaviors. In one of the few parent-twin studies of parenting during
middle childhood, twins who were parents of children under the age of 8 reported on their own
positive and negative parenting (Losoya, Callor, Rowe, and Goldsmith, 1997). Positive support was
mostly explained by heritable influences with the remaining variance due to nonshared environmental
influences, while negative control was explained by moderate heritable influences and modest shared
environmental influences. Heritability estimates in a parent-based design are evidence of passive rGE
or of parents’ heritable characteristics influencing the way they parent. The Twins, Adoptees, Peers,
and Siblings (TAPS) study takes a unique approach to understanding parenting in middle childhood
by including “virtual twins”—same-aged siblings who are biologically unrelated (McGuire, Segal, and
Hershberger, 2012). In this extension of the family-based design, five types of dyads were included
(MZ, DZ, virtual twins, full siblings, and unrelated friend pairs), and both child-rated and parent-rated
warmth were found to be significantly heritable, indicating evocative rGE. In contrast, a subset of
adoptive and nonadoptive families from CAP who participated in the Colorado Sibling Study (CSS)
with a school-aged child and a younger sibling were videotaped interacting with and without their
mothers. The authors found primarily shared environmental influences on maternal affection, atten-
tion, and responsiveness, and heritable influences on observed maternal control of children, suggesting
evocative rGE for maternal control (Rende, Slomkowski, Stocker, Fulker, and Plomin, 1992).
In this age range, evocative rGE seems to be particularly relevant for negative parenting char-
acteristics. In the TEDS sample, parental negativity at age 7 was explained by heritable shared and
nonshared environmental influences, whereas at age 9 parental negativity was only explained by
heritable and shared environmental influences (Larsson et al., 2008; Oliver, Trzaskowski, and Plomin,
2014). Moreover, at age 9 nearly half the variance in parental negativity was heritable, whereas herit-
ability was only modest for parental warmth (Oliver et al., 2014). For a subset of 8-year-old twins
from the TEDS using observational coding of an one-on-one cooperative task with their mothers,
large heritable influences were found on observed maternal “extreme” control (coded when mothers
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interfered with their child’s turn in the task; Eley et al., 2010). A similar interaction procedure was
used in the Twin Study of Behavioral and Emotional Development in Children (TBED-C; Burt and
Klump, 2013), which consists of 6- to 10-year-old twins and their families. Maternal warmth rated
continuously throughout the interaction was explained only by shared and nonshared environmental
influences, whereas maternal control was explained by heritable, shared, and nonshared environmen-
tal influences (Klahr, Thomas, Hopwood, Klump, and Burt, 2013).
Summary of Genetic and Environmental Influences

on Parenting During Middle Childhood
Much like in infancy and early childhood, most parenting constructs measured during middle child-
hood show heritable influences. Because primarily child-based designs have been used during this
developmental period, there is substantial evidence for evocative rGE. Shared environmental influ-
ences appear to play a larger role for positive parenting characteristics, such as warmth, than for nega-
tive parenting characteristics, such as control, indicating that positive parenting may also be subject
to passive rGE or is influenced by other, nonheritable characteristics that are shared among families.
Associations Between Parenting and Child Characteristics

Child-based twin designs have found that evocative rGE accounts, in part, for the association
between negative parenting and child characteristics during middle childhood. For example, using
data from the TBED-C children, the correlation between negative parenting and child aggression
was explained by heritable influences (Klahr, Klump, and Burt, 2014). In other words, for negative
parenting, evocative rGE was specific to child aggression, not rule breaking. In contrast, the covari-
ation between negative parenting and child rule breaking was explained by shared environmental
influences, indicating that there was no role for rGE in this association. Also within this sample,
observed maternal control and child control during a cooperative interaction was explained by over-
lapping heritable influences, suggesting evocative rGE (Klahr et al., 2013). Evocative rGE was also
suggested for the association between observed maternal extreme control during a cooperative task
and child anxiety in 8-year-old twins participating in TEDS (Eley et al., 2010).
Studies using the TEDS have systematically explored associations between negative parenting and
child characteristics using a variety of approaches to clarify the role of heritable and environmental
influences on associations between parenting and child behavior. Common heritable effects were
found between negative parenting and children’s antisocial behavior when the TEDS twins were
7 years old (Larsson et al., 2008). A cross-lagged approach in TEDS compared MZ twins within
families and found that differences in maladaptive parenting were related to differences in child self-
control longitudinally (Cecil, Barker, Jaffee, and Viding, 2012). In other words, because only MZ
twins were included, the authors controlled for shared genetics and shared environmental effects,
thereby examining only differences in parenting across the twins (nonshared environment). Moreo-
ver, longitudinal associations extended beyond middle childhood; differences in harsh parenting (but
not differences in negative parental feelings) in middle childhood were related to within-twin pair
differences in adolescent conduct problems. Finally, parental negativity (rated both by parents and
children in TEDS) during middle childhood was related to child depressive symptoms in adoles-
cence, and this was explained by common heritable influences, indicating evocative rGE (Wilkinson,
Trzaskowski, Haworth, and Eley, 2013).
Adoption designs have also examined associations between parenting and child behavioral out-
comes during middle childhood. For families in the EGDS, marital hostility was associated with child
externalizing problems via hostile parenting (Harold, Leve, Elam, et al., 2013). Because the adoptive
parents and their child in the EGDS are not biologically related, this association can be interpreted as
143
evidence of social transmission from parenting to child behavior problems. Across middle childhood
and into adolescence, adopted children in CAP who were classified as “at risk” (based on birth parent
antisocial behavior) experienced more negative control from adoptive parents compared to children
with no birth parent antisocial behavior “risk”. This was the case only for negative parenting, not for
warmth or inconsistency, and was interpreted as evocative rGE (O’Connor, Deater-Deckard, Fulker,
Rutter, and Plomin, 1998).
Summary of Associations Between Parenting and

Middle Childhood Characteristics
Child-based twin designs during middle childhood have shown that children’s heritable characteris-
tics, such as aggression and anxiety, can evoke negative parenting. However, this is not the case for all
child characteristics, and certain parenting qualities, such as hostility, appear to impact child behaviors
via nonheritable pathways. Notably, very little attention has been paid to whether positive parenting
is related to child characteristics via rGE or nonheritable processes.

Only a few studies have examined parenting within the context of GxE during middle childhood.
One such study using the EGDS found that birth mother processing speed moderated adoptive
mother uninvolved parenting. Specifically, uninvolved parenting was associated with school-aged
children’s membership in an internalizing-only symptom group (as opposed to low-symptom group)
only for children who lacked an inherited “risk” (birth mother slow processing speed; Roos et al.,
2016). A second study using the TEDS sample found that negative parenting in middle child-
hood moderated heritable effects on adolescent depressive symptoms with larger heritable effects
for children who experienced more negative parenting during middle childhood (Wilkinson et al.,
2013). Finally, hyperactivity and inattention problems moderated the shared environmental influ-
ences between child conduct problems and negative parenting in 6-year-old twins assessed through
the Tokyo Twin Cohort Project and the Tokyo Twin Cross-Sectional Survey (Fujisawa, Yama-
gata, Ozaki, and Ando, 2012). Shared environmental influences explained a larger proportion of the
covariation between conduct problems and negative parenting for children with high levels of hyper-
activity and inattention problems as compared to children with low levels. A possible interpretation
of this finding is that within families, parents respond consistently to conduct problems when both
children are highly hyperactive or inattentive, whereas parents respond less consistently to conduct
problems when both children exhibit low levels of hyperactivity or inattention.
Summary of Gene x Environment Interactions During Middle Childhood

GxE during middle childhood remains an area ready for investigation. To date, there is some evi-
dence suggesting that negative parenting during this time may amplify inherited risk for psycho-
pathology symptoms but may also lead to symptoms in the absence of inherited risk. One study
suggests that parenting during middle childhood may have effects on child behavior longitudinally,
at least into adolescence. Thus, middle childhood may be a particularly important developmental
period for identifying early GxE that may have additional implications for adolescent functioning.
Summary of Parenting During Middle Childhood

Middle childhood has received the least attention from behavioral geneticists who study parent-
ing, but the literature in this developmental period strongly indicates that children are not passive
144
recipients of parenting. Rather, most parenting constructs studied during this middle childhood
show heritable influences in child-based designs, suggesting that children’s heritable characteristics
evoke the parenting they receive. Moreover, while many findings from studies examining covaria-
tion between parenting and child characteristics support the importance of evocative rGE, there is
specificity in which child characteristics operate via evocative rGE processes. Notably, during middle
childhood, children can report on the parenting they experience with some studies including child
reports of parenting and of child behaviors. As we review in the next section, estimates of heritable
and environmental influences often vary by reporter, and thus including middle childhood reports of
parenting may be one avenue for researchers to further pursue. Additionally, parenting during mid-
dle childhood has not yet received much attention from researchers interested in GxE and has been
largely omitted from molecular genetics research.
Parenting During Adolescence

In addition to the parenting qualities studied in other developmental periods, such as warmth and
conflict, qualities that are relevant to adolescents’ increasing agency and the social world, such as
autonomy granting and parental monitoring, have been examined in genetically informed designs.
A few studies using parent-based designs have examined various parenting characteristics during
adolescence and generally find significant heritable and nonshared environmental influences. For
instance, adult twins reported on their own use of physical discipline and limit setting towards their
children. There were modest heritable influences and large nonshared environmental influences on
both physical discipline and limit setting, with no evidence of shared environmental influences (Wade
and Kendler, 2000). German twin parents reported on their overprotective, authoritarian, support-
ive/indulgent, and rejecting parenting with all domains showing significant nonshared environmen-
tal influences (Spinath and O’Connor, 2003). In addition, individual differences in overprotective,
authoritarian, and supportive/indulgent parenting were also due to moderate heritable influences,
while the remaining variance in rejecting parenting was due only to shared environmental influences.
Findings of significant shared environmental influences are notable because they reflect experiences
common to family members, such as shared rearing experiences, which for adult twin parents are
likely distal or diffuse experiences. In other words, shared environmental influences that continue to
be important for adult twins could include family-of-origin rearing environment during childhood
or less distal influences like current contact with their co-twin (or among family-of-origin members
more generally). A separate study in which twin parents reported on their own care and overprotec-
tion behaviors found differences for mothers and fathers, with heritable influences on both parenting
constructs greater for mothers than for fathers, whereas nonshared environmental influences were
greater for fathers than for mothers (Pérusse, Neale, Heath, and Eaves, 1994). Shared environmental
influences did not significantly explain variance in parenting in this report. Finally, parenting has
been explored in the Twin/Offspring Study in Sweden (TOSS; Neiderhiser and Lichtenstein, 2008),
a parent-based twin design that consists of families in which parents are twins and offspring are
adolescents range from 11 to 22 years. Findings from this study are discussed in more detail below.
Child-based designs have consistently found that both positive (e.g., positivity, monitoring) and
negative (e.g., negativity, conflict, punitive discipline) parenting qualities during adolescence are her-
itable (Button et al., 2008; Elkins, McGue, and Iacono, 1997; Latendresse et al., 2010; Plomin, Reiss,
Hetherington, and Howe, 1994; Reiss, Neiderhiser, Hetherington, and Plomin, 2000). For instance,
in the population-based FinnTwin cohort of Finnish MZ and DZ twins, autonomy granting, knowl-
edge, and warmth were all heritable (Latendresse et al., 2010). Likewise, within the MFTS, parent-
ing qualities such as warmth, involvement, parent-child regard, and conflict have been found to be
heritable with estimates greater for older adolescents relative to younger adolescents (Elkins et al.,
1997; McGue, Elkins, Walden, and Iacono, 2005). Relative to positive parenting, negative parenting
145
characteristics tend to show higher estimates of heritability (Button et al., 2008; Neiderhiser et al.,
2004, Neiderhiser, Reiss, Lichtenstein, Spotts, and Ganiban, 2007; Reiss et al., 2000). For example,
in the G1219 cohort of twins and siblings living in the United Kingdom, heritability was higher for
punitive discipline than for constructive discipline (Lau, Rijsdijk, and Eley, 2006). Adolescents in the
E-RISK study reported on their experiences of victimization between the ages of 12 and 18 years.
In contrast to reports from the same twins at earlier ages that found no evocative effects ( Jaffee et al.,
2004), there were substantial heritable influences and smaller but significant nonshared influences on
maltreatment (Fisher et al., 2015). Together, this accumulation of evidence of heritability of parent-
ing in child-based designs can be interpreted as evocative rGE.
Much behavioral genetic research on parenting in adolescence comes from the Nonshared Envi-
ronment in Adolescent Development (NEAD) study (Neiderhiser, Reiss, and Hetherington, 2007;
Reiss et al., 2000). NEAD is a study of nondivorced and stepfamilies with same-sex adolescent twin
or sibling pairs who were within 4 years of age of one another. Nondivorced families included,
MZ, DZ, and full siblings, and stepfamilies included full, half, and genetically unrelated (step) sib-
lings. Data from NEAD show that parents’ positivity, negativity, and monitoring were all explained
by heritable influences, but that environmental estimates varied by reporter or assessment method
(Plomin et al., 1994; Reiss et al., 2000). Parents’ reports of these constructs were also explained by
substantial shared environmental influences, whereas adolescents’ reports were explained by modest
shared environmental influences and large nonshared environmental influences. Observer ratings
were primarily explained by shared and nonshared environmental influences; and heritability esti-
mates were negligible (Neiderhiser et al., 2004). Similar patterns were found in an ethnically diverse
sample of adolescent twins participating in the Texas Twin Project who reported on the parenting
they received (Patterson, Cheung, Mann, Tucker-Drob, and Harden, 2017). For warmth and control,
there were modest heritable effects, moderate shared environmental effects, and substantial nonshared
environmental effects. In contrast to data from NEAD, however, there were no heritable influences
on monitoring, which was primarily attributable to shared and nonshared environmental influences.
One approach to clarifying if passive or evocative rGE (or both) are present is to assess the same
constructs in parent- and child-based twin samples and compare the findings. Using this approach,
data from NEAD (child-based) were matched with data from TOSS (parent-based). This comparison
analysis showed five general patterns of findings that varied based on construct, parent assessed (i.e.,
mother or father), and reporter (i.e., parent, child, or observer) (Neiderhiser et al., 2004; Neiderhiser,
Reiss, Lichtenstein, et al., 2007). First, a single construct, maternal monitoring, showed evidence of
only passive rGE for both parent and child reports. Second, three constructs, maternal negativity,
maternal control, and paternal control, showed evidence of only evocative rGE for both parent and
child reports. Third, three constructs, maternal positivity, paternal monitoring, and paternal negativ-
ity, showed evidence of passive and evocative rGE for both parent and child reports. Fourth, interpre-
tations for paternal positivity differed by reporter; father report showed evidence of only evocative
rGE, whereas adolescent report showed evidence of both passive and evocative rGE. Finally, observer
ratings of maternal positivity, negativity, and control showed no evidence of rGE. Taken together, for
most constructs examined, parent and adolescent perceptions of parenting appear to operate under
the same etiologic processes. Moreover, most parenting characteristics examined in this study suggest
that both parents’ and children’s heritable characteristics impact parenting.
Summary of Genetic and Environmental Influences

on Parenting During Adolescence
Most parenting characteristics studied in adolescence show significant heritable influences, regardless
of whether they were examined in a parent-based design or a child-based design. Because heritable
influences have different interpretations depending on the type of design used, this suggests that for
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many characteristics during adolescence, both passive and evocative rGE are operating. For three
out of four constructs, there was evidence of both passive and evocative rGE when systematically
examined in both types of designs, with a caveat that conclusions depend on reporter and the parent
being assessed.
Associations Between Parenting and Adolescent Characteristics

Findings from NEAD suggest that both mother and father positive parenting are associated with
adolescent outcomes such as antisocial behavior, cognitive agency, sociability, autonomy, social
responsibility, and self-worth (Reiss et al., 2000). Moreover, the associations between these parenting
constructs and adolescent outcomes are best explained by common heritable influences. Compared
to positivity, parental knowledge of the child was less heritable in general and correlations with the
adolescent outcomes listed above were somewhat weaker. However, the majority of the covariance
between parental knowledge and adolescent outcomes was best explained by heritable influences.
Particularly notable was that all of the heritable influences on fathers’ knowledge was also corre-
lated with heritable influences on adolescent antisocial behavior and social responsibility. In contrast,
when adolescent drinking frequency was examined in association with parental knowledge in the
FinnTwin study the association between these constructs was predominantly explained by shared
environmental influences (Latendresse et al., 2010). Differences between FinnTwin and NEAD may
stem from different measurement approaches; the FinnTwin study used a combined measure of
adolescents’ perceptions of both parents, whereas NEAD used a multi-rater approach and considered
mothers’ and fathers’ parenting separately.
Similar to earlier ages, much more attention has been given the role of negative parenting in
adolescent outcomes. There is consistent evidence in multiple samples using a variety of research
designs that negative parenting is associated with heritable adolescent characteristics, such as negative
emotionality, antisocial behaviors, and depressive symptoms (Ganiban et al., 2011; Ge et al., 1996;
Narusyte et al., 2007). Findings from the NEAD study indicate that maternal and paternal negativ-
ity are both associated with adolescent antisocial behavior, depressive symptoms, cognitive agency,
and social responsibility, and that these associations can largely be explained by overlapping herit-
able influences (Reiss et al., 2000). One exception to this overall pattern of findings is adolescent
self-worth; for both maternal and paternal negativity, inverse correlations with self-worth were best
explained by nonshared environmental influences (Reiss et al., 2000). However, the overall pattern
that negative parenting characteristics and adolescent outcomes are due to covarying heritable influ-
ences is consistent; within the G1219 cohort, for instance, heritable influences on maternal negativity
were completely explained by heritable influences on oppositionality, delinquency, depression, and
anxiety (McAdams, Gregory, and Eley, 2013) and overlap between punitive discipline and external-
izing behaviors was attributable to heritable effects (Button et al., 2008).
Generally, longitudinal work has further bolstered the claim that both evocative and passive rGE
are at play in the parental negativity-adolescent outcome association. For instance, NEAD families
were assessed twice during adolescence, 3 years apart. Cross-lagged models indicated that for both
mothers and fathers, after controlling for stability and within-time associations, changes between
parent negativity and adolescent antisocial behavior and depressive symptoms were due to heritable
factors, indicating evocative rGE (Neiderhiser, Reiss, Hetherington, and Plomin, 1999). In a simi-
lar approach, a cross-lagged model using data from the MFTS collected when twins were 11 and
14 years old found bidirectional effects between parent-child conflict and adolescent externalizing
behaviors. Unlike the findings from NEAD, both cross-lagged pathways were explained by approxi-
mately equal estimates of heritable, shared, and nonshared environmental influences (Burt, McGue,
Krueger, and Iacono, 2005). It is possible that these differences are attributable to the shorter age
range of MFTS adolescents. Finally, Moberg, Lichtenstein, Forsman, and Larsson (2011) measured
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parental emotional overinvolvement and adolescent internalizing symptoms repeatedly in the Twin
Study of Child and Adolescent Development (TCHAD; Lichtenstein, Tuvblad, Larsson, and Carl-
ström, 2007), a representative sample of adolescent twins living in Sweden. Using a cross-lagged
design, they found that parental emotional overinvolvement was not linked to increases in internal-
izing symptoms later in adolescence, but that girls’ internalizing symptoms increased parental emo-
tional overinvolvement. Moreover, this child-driven effect was attributable to significant heritable
influences, suggesting evocative rGE.
Two studies suggest that passive rGE is also important for associations between parenting and ado-
lescent behavior. In the aforementioned QNTS, when twins were assessed at ages 13 and 14, initial
maternal negativity was related to increased adolescent delinquency within a nongenetic model (i.e.,
one that included one randomly selected co-twin per family), but when these models were re-run
including both twin members, the link between maternal negativity and later adolescent delinquency
became nonsignificant (Guimond et al., 2016). In other words, once child heritable influences were
controlled, maternal negativity was no longer associated with adolescent delinquency, suggesting a
passive rGE process in which mothers who have an inherited predisposition to negativity towards
their children have children who engage in more delinquent behavior. Similarly, a cross-lagged
approach with MZ twins from the MFTS found that overall increases in parent-child conflict over
time were not related to concomitant increases in adolescent externalizing problems, but for a subset
of twin pairs with the most extreme differences in parent-child conflict, the co-twin with the worse
parent-child conflict also had higher externalizing behaviors in mid-adolescence. This link was not
due to evocative factors—in other words, those co-twins with the worse parent-child conflict did
not have more initial externalizing behaviors. Therefore the authors concluded that there is evidence
of an environmental effect between initial parent-child conflict and later externalizing behaviors
(Burt, McGue, Iacono, and Krueger, 2006).
The few COT and ECOT studies of parenting of adolescents confirm what the studies sum-
marized above suggest—that parenting during adolescence can be evoked by adolescents’ herit-
able characteristics—but also support the assumption that some parenting characteristics are causally
related to child characteristics for environmental reasons. For instance, a COT design using twin
parents participating in the Australian Twin Registry examined whether twin parents who were
discordant on their use of harsh punishment (i.e., use of more extreme physical discipline) had
children who exhibited more behavioral problems (Lynch et al., 2006). This design controlled for
family-level confounding factors and found that use of harsh punishment was associated with greater
externalizing and substance use problems, strengthening conclusions of causality within the rearing
environment. All other COT and ECOT studies of parenting have exclusively relied on data from
the parent-based TOSS and either NEAD or TCHAD. In the first published ECOT study, Narusyte
and colleagues (2008) combined TOSS and TCHAD data and found that adolescent internalizing
symptoms evoked maternal emotional over-involvement and that this effect was via heritable, not
environmental, pathways. Similarly, an ECOT design using TOSS and NEAD data found that ado-
lescent externalizing behavior evokes parental negativity, indicating evocative rGE (Marceau et al.,
2013). These findings can both be interpreted as evocative rGE.
Different parenting constructs examined in those same studies appear to be causally related to ado-
lescent characteristics. TOSS and TCHAD data, for instance, have been combined in an ECOT design
that found differential effects for mothers’ and fathers’ criticism with adolescent externalizing behavior
(Narusyte et al., 2011). For mothers, evocative rGE was present, such that adolescents’ heritable exter-
nalizing behavior evoked maternal criticism; that was not the case for fathers. For fathers, there was
no evidence of rGE, but instead results indicated a direct environmental association between pater-
nal criticism and adolescent externalizing behavior. In a different report (also using TOSS/TCHAD
data), parental criticism was associated with adolescent somatic symptoms via environmental pathways,
rather than rGE for both mothers and fathers (Horwitz et al., 2015). Similarly, an ECOT using TOSS
148
and NEAD demonstrated that parental knowledge was related to decreased adolescent externaliz-
ing behaviors, not via heritable pathways but through mechanisms that suggest social transmission
(Marceau et al., 2014). Finally, using a COT approach with the TOSS and TCHAD cohorts, Hannigan
and colleagues (2017) found that the link between poor parent-child relationship quality and adoles-
cent internalizing problems was not attributable to rGE but to environmental mechanisms.
In summary, COT and ECOT designs are useful tools that demonstrate that different parenting
constructs are subject to different etiological processes. Together, the few studies that have examined
parenting and offspring outcomes using polygenic risk scores have found evidence for both passive
and evocative rGE effects. Data from the Adolescent/Adult Family Development Project (Chassin,
Rogosch, and Barrera, 1991), a multigenerational sample of families of alcoholics, sheds light on rGE
processes at the measured gene level. This sample includes genotypic data from the second (moth-
ers and fathers) and third generation (child) of study participants. Researchers created a polygenic
risk score by combining genotypes (i.e., DRD1, DRD2, ANKK1, DDC, TPH2, CHRM2) that
have been associated with behavioral under-control in prior literature. Both mothers’ and fathers’
polygenic risk scores were included in analyses, so the authors were able to control for passive rGE
while testing for evocative rGE. There was evidence of passive rGE between parental risk score and
parental monitoring but not family conflict (Elam et al., 2016, 2017). Additionally, there was evi-
dence of evocative rGE between child risk score and parental monitoring, via child impulsivity. In the
same sample, a different polygenic risk score composed of genotypes (i.e., DBH, GABA, GABRB1,
PRKCE) that index adolescents’ response inhibition was created based on an independent GWAS of
young adults’ Stroop Colorword performance. Higher scores were related to maternal inconsistency
for boys whose parents met criteria for substance use disorder, which can be interpreted as evocative
rGE (Wang, Chassin, Lee, Haller, and King, 2017). Finally, in a separate study of adolescents, Salvatore
and colleagues (2015) tested whether a polygenic risk score for externalizing behavior that had been
derived from a discovery sample of adults was also related to adolescent externalizing behavior. Even
after controlling for parental history of externalizing symptoms, the risk score was related to ado-
lescent externalizing symptoms. Moreover, adolescents with the highest risk scores who also expe-
rienced low parental monitoring exhibited the most externalizing problems (Salvatore et al., 2015).
Summary of Associations Between Parenting and

Adolescent Characteristics
In this developmental period, many parenting and adolescent characteristics are associated via herit-
able pathways and, particularly, are subject to evocative rGE processes. However, there are excep-
tions; links between some parenting and adolescent characteristics have shown significant shared
and nonshared environmental influences, and longitudinal work has found evidence of passive rGE
and environmental pathways. Moreover, ECOT and COT approaches have found evidence for two
of the possible mechanisms (evocative rGE and environmental) linking parenting and adolescent
characteristics. Externalizing symptoms appear to evoke parental negativity and maternal criticism
via heritable pathways, and internalizing symptoms elicit parental emotional overinvolvement. In
contrast, parental knowledge and paternal criticism are related adolescent externalizing symptoms via
nonheritable pathways, as are poor parent-child relationship and adolescent internalizing symptoms.
At the molecular genetic level, polygenic risk scores have been used to identify the measured genes
purportedly underlying evocative rGE processes.

Parenting moderates the heritability of adolescent externalizing behavior. In the FinnTwin study,
parental monitoring moderated both heritable and shared environmental influences on smoking; at
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higher levels of parental monitoring when twins were 12 years old, heritable influences on smok-
ing at 14 years were lower, whereas shared environmental influences were higher (Dick et al., 2007).
In other words, higher levels of parental monitoring during early adolescence increased the effects
of within-family similarity (shared environment) during middle adolescence at least in regard to
smoking. In tandem, there was a corresponding decrease in the heritable effects on smoking. The
same general pattern was found in NEAD families, but in this case, heritable, shared, and nonshared
environmental influences on adolescent antisocial behavior were all moderated by parental negativity
(Feinberg et al., 2007). Specifically, estimates of heritable, shared, and nonshared influences on adoles-
cent antisocial behavior were all highest at higher levels of parental negativity. Additionally, heritable
and shared environmental influences on adolescent antisocial behavior were moderated by parental
warmth, such that heritability estimates were highest and shared environmental estimates were lowest
at low levels of warmth. These findings indicate that, as the parent-adolescent relationship worsens,
heritable influences on antisocial behavior increase. One way to interpret this moderation is that
a parent-adolescent relationship characterized by low negativity or high warmth may dampen the
genetically influenced propensity to engage in antisocial behavior during adolescence. These findings
must be considered with caution, however, as the Feinberg et al. findings are cross-sectional. A more
mixed pattern of findings was found for the G1219 cohort (Button et al., 2008). At low levels of
maternal punitive discipline, there were higher heritable influences on externalizing, whereas there
was the opposite for paternal punitive discipline—at high levels of paternal punitive discipline, there
were greater heritable influences on externalizing (Button et al., 2008). Taken together these find-
ings suggest that, in some cases, parenting characteristics or behaviors can buffer heritable effects on
adolescent externalizing behaviors.
While these studies conceptualized parenting as the moderator of the heritability of adolescent
phenotypes, two studies to date have done the opposite and conceptualized adolescent phenotypes
(i.e., temperament and personality) as moderators of heritability of parenting. Using the NEAD
study, Ganiban and colleagues (2011) found that certain features of adolescent temperament moder-
ated heritable and shared environmental influences on parental negativity even after taking possible
rGE effects into account (Ganiban et al., 2011). For example, at higher levels of offspring negative
emotionality or sociability, heritable influences on parental negativity were larger than at low levels
of offspring negative emotionality. The opposite pattern was found for adolescent shyness, such
that, at high levels of child shyness, there were diminished heritable influences on father negativity
only. High negativity, emotionality, and shyness in the adolescent were all associated with decreased
shared environmental influences on parental negativity. The authors interpreted these findings as a
whole to indicate that parenting negativity becomes more child-specific when adolescents exhibit
more challenging temperament characteristics. Data from the MFTS used a similar approach with a
somewhat different pattern of findings. For adolescent twins, estimates of heritable, shared, and non-
shared environmental influences on parental regard and conflict were moderated by adolescent per-
sonality factors (South, Krueger, Johnson, and Iacono, 2008). For instance, when adolescent positive
emotionality was high, heritable influences on parental regard were highest and on parental conflict
were lowest. Additionally, for parental regard, shared and nonshared environmental influences were
moderated by positive emotionality, such that both were highest when adolescents exhibited low
positive emotionality. It is possible that these differences are attributable to differences in constructs
(temperament as opposed to personality), or that the NEAD study used parent-report whereas the
MFTS used adolescent self-report and therefore the two studies capture different processes.
Another published report from the NEAD study suggests that broader family characteristics,
such as marital conflict about the adolescent and marital satisfaction, moderate the heritable and
environmental influences on parenting (Ulbricht et al., 2013). Although there were some differ-
ences for mothers and fathers, patterns were generally consistent. When marital conflict was low,
there was more variance in mothers’ negativity attributable to shared environmental influences; as
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levels of conflict increased, there were dramatic decreases in shared environmental influences on
negativity. In contrast, when marital conflict was low, there was more variance in fathers’ negativity
attributable to heritable and shared environmental influences. For both parents, at increasing levels of
conflict, nonshared environmental influences increased. Marital satisfaction models showed that for
mothers, at increasing levels of dissatisfaction, there were increasing heritable influences on parenting
negativity. For fathers, as marital dissatisfaction increased, nonshared environmental influences on
parenting negativity increased, but shared environmental influences decreased. One interpretation of
this moderation is that, at higher levels of marital distress, children’s inherited characteristics impact
mothers’ parenting negativity to a greater degree, while nonshared environmental factors become
more important for fathers’ parenting negativity.
One of the few genetically informed studies to include both parenting and culture combined
data from the adult twins from the TOSS with data from the Keio Twin Project, a study of Japanese
adult and adolescent twins (Shikishima, Hiraishi, Yamagata, Neiderhiser, and Ando, 2013). Twins
recalled their experience of being parented before the age of 16 years for both their mothers and
fathers in domains of warmth, authoritarianism, and protectiveness. Although phenotypic structures
were similar for Japanese and Swedish twins, relative heritable and environmental influences differed
by culture. For Japanese twins, except for maternal protectiveness (which was explained by moderate
shared environmental influences), the covariation of warmth and authoritarianism was attributable to
substantial heritable influences. In contrast, for Swedish twins, heritable influences on parenting were
negligible and the covariation of the three parenting domains was explained by shared environmental
influences. The significant heritability of parenting within the Japanese sample suggests that either
parents were responding to their children’s heritable characteristics or that heritable characteristics
impact the twins’ recall of parenting. Shared environmental influences on Swedish twins’ experiences
of parenting suggest that within those families, parenting was not child-specific. Findings from the
Child Development Project, a community sample assessed longitudinally, have reported GxE interac-
tions at the molecular genetic level using genotypes that have been associated with alcohol depend-
ence in adult samples. Within this cohort, there was an interaction between GABRA2 and parental
monitoring on adolescent externalizing behavior trajectories. As parental monitoring decreased, the
link between genotype and externalizing behavior became stronger (Dick et al., 2009). Similarly,
Dick and colleagues (2011) found an interaction between several CHRM2 and parental monitor-
ing on adolescent externalizing behaviors. Much like the previous study, as parental monitoring
decreased, the link between the CHRM2 “risk” genotype and externalizing behavior increased.
Two studies tested Differential Susceptibility Theory (Belsky and Pluess, 2009), namely that certain
individuals are particularly sensitive to environmental inputs and fare best in positive environments
and worst in negative ones due to genetic variation. Each used data from the National Longitudinal
Study of Adolescent Health (Add Health; Harris et al., 2009), a nationally representative sample of
adolescents in the United States, measured a compilation of purported “plasticity” genes, coded those
genes based on the hypothesized effects (e.g., the genotype is associated with environmental sensitiv-
ity), and combined each genotype into a score indexing each participant’s overall genetic plasticity.
In addition, both studies tested for evocative rGE prior to evaluating GxE by assessing whether each
genotype was correlated with parenting. Belsky and Beaver (2011) found that for males who had
higher plasticity scores across five genes (i.e., DAT1, DRD2, DRD4, 5HTTLPR, MAOA), negative
maternal parenting was associated with poorer adolescent self-regulation, while positive maternal
parenting was associated with better adolescent self-regulation compared to males with lower plas-
ticity scores. The second study found that for adolescents with a high plasticity score based on four
genotypes (i.e., DAT1, DRD2, DRD4, 5HTTLPR), maternal positive parenting was linked to the
lowest levels of adolescent parenting stress when they themselves became parents (Beaver and Belsky,
2012) compared to individuals with low plasticity scores. Conversely, maternal negative parenting
was linked to the highest levels of parental stress for adolescents with high plasticity scores.
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Summary of Gene x Environment Interactions During Adolescence

Within the adolescent period, both family-based and molecular genetic designs have taken a variety
of approaches to study parenting in a GxE framework. Some have conceptualized parenting as a
moderator of the heritability of adolescent phenotypes, finding evidence that monitoring, warmth,
negativity, and punitive discipline moderate heritable and environmental estimates on offspring phe-
notypes. Others have investigated child-level and family-level moderators of the heritability of par-
enting, finding evidence that etiology of parental negativity, regard, and conflict depend on child
characteristics such as temperament. Finally, molecular genetic research has explored singular can-
didate genes as well as composites that interact with parenting to predict adolescent maladjustment.
Summary of Parenting During Adolescence

The literature we have reviewed highlights that adolescents’ inherited characteristics evoke many
types of parenting behaviors, lending further credence to the assertion that developmental work
should take family-level confounds into account when examining links between parenting and off-
spring characteristics. In addition, some parenting constructs show evidence of passive rGE and direct
environmental influences. Notably, innovative approaches, such as ECOT and polygenic risk scores,
have only been employed with data collected within this developmental period. Findings from
ECOT studies are generally consistent with other approaches used during adolescence and have
found evidence of evocative and passive rGE as well as direct environmental effects, and polygenic
risk score approaches have begun to examine rGE at the molecular genetic level, finding evidence of
both passive and evocative rGE.
Aside from some of the adoption work in infancy and early childhood, the bulk of genetically
informed studies that measure fathers’ parenting has been conducted on adolescents. There are many
consistencies across mothers and fathers, but studies that include both parents have elucidated some
differences as well. For instance, in some parent-based designs, heritability estimates for mothers’
parenting are higher than for fathers’ parenting. Another example is evidence of environmentally
mediated links between paternal criticism and adolescent externalizing behaviors, but genetically
mediated links for maternal criticism and adolescent externalizing behaviors. In conjunction with
findings indicating that heritable and environmental influences on some parenting constructs are dif-
ferentially moderated for mothers and fathers, there is accumulating evidence that certain parenting
processes, such as negativity, may differ for mothers and fathers.
Future Directions in Genetics and Parenting

We conclude with a discussion of the mutual benefits that may be derived by a more thorough
integration of family-based genetic research and typical developmental studies and the consequent
implications for both theory and methods. Behavioral genetic designs that control for confounding
genetic influences within families provide an opportunity to test crucial assumptions underlying
many theories of parenting. Simultaneously, this work provides an impetus for a revision of exist-
ing theoretical models of development to incorporate findings from both areas of research (Moore
and Neiderhiser, 2014). For instance, there is no clear theory to explain why there would be both
direct (via child impulsivity) and indirect (via maternal hostility) evocative rGE processes between
genetic predisposition for ADHD and child ADHD symptoms (Harold, Leve, Barrett, et al., 2013).
Such revisions may aid in the clarification of developmental processes and may open new avenues
of exploration.
Critical to the future of family-based genetic research is adoption of some of the measurement
used in developmental and family process research studies. Specifically, measurement of purported
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environmental features, such as parenting, are often assessed only using parent report in behavioral
genetic designs. Studies that have used different approaches, many of them described above, show
that the findings of heritable and environmental influences tend to vary as a function of the way the
construct was measured, which may reflect that measures are capturing different aspects of related
processes (Neiderhiser et al., 2004; Neiderhiser, Reiss, Lichtenstein, et al., 2007). Moving towards
fine-grained observational assessments of parenting, as some genetically informed studies have begun
to do (Klahr et al., 2013; Roben et al., 2015), may make behavioral genetics work more directly
relevant to cutting-edge developmental methodology.
Registry data have proven extremely useful for obtaining the large sample sizes necessary to
understand low-prevalence phenotypes and to study diverse family configurations, thus providing
additional and powerful support for the critical role of GE interplay in the etiology of problematic
behaviors and psychopathology. However, a major drawback of studies that rely on registry data is
that the level of detail one can obtain from registries tends to be limited. For instance, a series of stud-
ies using Swedish registry data has explored the link between parent and offspring psychopathology
(e.g., criminal behavior, drug abuse) using complementary family-based designs (e.g., comparison of
intact, step, and adoptive families) (Kendler, Ohlsson, Morris, Sundquist, and Sundquist, 2015; Kend-
ler, Ohlsson, Sundquist, and Sundquist, 2016a; Kendler, Morris, et al., 2016). Kendler, Morris, et al.
(2016) examined biologically related sibling pairs in which one sibling was reared with a biological
parent at high risk for drug abuse and one was living with adoptive parents to estimate risk for drug
abuse in the offspring. Compared to siblings living with their biological parent, adopted offspring had
a significantly decreased risk for drug abuse. A similar pattern has been found for criminal behavior;
moreover, this protective factor of adoptive environment is diminished if one of the adoptive parents
had their own history of criminal behavior (Kendler et al., 2015; Kendler, Ohlsson, et al., 2016).
Together, these findings suggest the rearing environment buffers a genetic liability for drug abuse or
criminal behavior, but the mechanisms by which this occurs are unknown.
We have limited our discussion of empirical behavioral genetics research to those that expressly
measure parenting. However, parent-offspring designs have produced additional noteworthy findings
relevant to parenting. For instance, Turkheimer, Haley, Waldron, D’Onofrio, and Gottesman (2003)
found that SES moderated heritability estimates on children’s intelligence. For children in low-SES
households, intelligence was explained by large shared environmental estimates and minimal herit-
able influences. For children in high-SES households, intelligence was explained by large heritable
influences and minimal shared environmental influences. A possible interpretation of this is that the
measure of SES is actually capturing meaningful differences in rearing environment and that these
differences result in differences in how the heritable and environmental factors influence child intel-
ligence. Similarly, an innovative and alluring finding from molecular genetics supports the concept of
“genetic nurturing”—that nontransmitted alleles impact offspring via effects on the rearing environ-
ment (Kong et al., 2018). In this case, parent-offspring pairs were genotyped and a polygenic score
was created based on a GWAS of educational attainment. The researchers found that parents’ non-
transmitted alleles explained a significant proportion of variance in offspring educational attainment.
While studies such as these offer exciting possibilities about the influence of parents on children,
further work must explicitly measure parenting in order to conclude that such links are attributable
to parenting. Moreover, for genetically informed designs to inform basic and applied developmental
work, they need to include high-quality assessments of family processes that map onto what is being
studied by developmentalists (Leve et al., 2017; Moore and Neiderhiser, 2014). Importantly, a number
of studies described above have begun to do this (Boivin et al., 2005; Burt and Klump, 2013; Leve
et al., 2013; Oliver and Plomin, 2007; Reiss et al., 2000).
One aim for integrating the fields of behavioral genetics and developmental science is to use
behavioral genetics findings to inform prevention and intervention research on child socioemotional
problems (Harold et al., 2017; Leve et al., 2017). Heritability is not synonymous with immutability,
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and mean-level changes can occur within a phenotype without changing estimates of heritable,
shared, or nonshared environmental influences (Maccoby, 2000). Applied work often relies on the
assumptions that parenting is the catalyst for problematic child outcomes and that improving par-
enting will result in improved child outcomes. Yet, as discussed above, links between parenting and
child behaviors could be under the influence of passive rGE, which would mean that interventions
may target the wrong mechanism. By removing family-level confounds, it is possible to identify
mechanisms to target that are more likely to be amenable to change. For instance, if evocative rGE
is operating, parents could be taught skills to effectively respond to specific, relevant child charac-
teristics (Harold et al., 2017). Leve and colleagues propose genetically informed interventions, akin
to personalized medicine, that capitalize on identified GxE (Harold et al., 2017; Leve et al., 2017).
Namely, interventions could be tailored to specific combinations of inherited and environmental
risk (or protective) factors. In addition, by clarifying the timing of heritable or environmentally
influenced developmental changes, it may be possible to identify sensitive periods particularly ame-
nable to intervention. At the molecular genetic level, there is preliminary evidence suggesting that
interventions can diminish the effects of heritable risks over time (Brody, Chen, Beach, Philibert, and
Kogan, 2009), and gene-by-intervention research has suggested that genetic variants moderate the
effects of intervention (Albert et al., 2015; Musci and Schlomer, 2017). Taken together, GxE findings
and research can help to provide information about the characteristics that can diminish the impact
of one’s risk factors or enhance the effects of one’s existing protective factors (Leve et al., 2017).
While genetically informed intervention work is in its nascent stages, in practice, such an approach
could involve choosing or modifying interventions based on an individual’s temperamental charac-
teristics or genetic makeup.
At present, the field of behavioral genetics could benefit from expanding the participant ages
and ethnicities studied. Genetically informed studies of parenting are focused on early childhood
or adolescence; GE processes specific to middle childhood are less well understood and work is just
beginning to examine parenting beyond adolescence. Moreover, most of the samples comprising
genetically informed designs rely on White samples (Fullerton, Yu, Crouch, Fryer-Edwards, and
Burke, 2010; Knerr, Wayman, and Bonham, 2011). Because Scandinavian countries have a long
history of excellent population-based record keeping, they have been the source of data for many
registry-based analyses and by definition primarily include European-ancestry individuals. Further-
more, minority populations often endorse a distrust of genetic research rooted in historical mistreat-
ment (Scharff et al., 2010). Although studies examining other psychosocial constructs have not found
evidence of etiological differences based on ethnic heritage (Rowe, Vazsonyi, and Flannery, 1995),
cultural contexts impact parenting and child development (Bornstein and Lansford, 2010; Kotchick
and Forehand, 2002) and minority families may have systematically different sociocultural experi-
ences that impact family functioning (Mcloyd, Cauce, Takeuchi, and Wilson, 2000) and warrant
inclusion in family-based genetic research.
Conclusions
In conclusion, we echo statements made by many prominent scholars about the importance of
behavioral genetics for developmental science (Plomin, Owen, and McGuffin, 1994; Rutter, 2005;
Scarr and McCartney, 1983). Findings from behavioral genetics have been misapplied in the past to
support claims that what parents do does not matter for their children’s’ development (Harris, 1998),
but we have summarized a number of studies above that highlight the bidirectional links between
parents and children that exist beyond inherited characteristics. Importantly, genetically informed
designs are a tool that can be used by developmental researchers to test crucial assumptions underly-
ing many parenting theories. In other words, to conclude that parents do matter, we must adequately
test the competing alternative hypothesis that they do not (Rutter, 2005). To do that, behavioral
154
geneticists must collect measurements of parenting that are compatible with the high-quality devel-
opmental and family process approaches relevant to parenting.
Acknowledgments
Research reported in this publication was supported by the Environmental influences on Child
Health Outcomes (ECHO) program, Office of the Director, National Institutes of Health, under
award number UG3 OD023389.
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5
PRENATAL PARENTING
David A. Coall, Anna C. Callan, Julie Sartori, and James S. Chisholm
Introduction
Because the factors that contribute to the development of a healthy newborn are so complex,
understanding them requires insights from many disciplines. Crucial to this understanding is the role
parents play. Through their behavior, psychology, and biology, parents influence the development of
their children well before birth (Glover and Capron, 2017). In this chapter we review insights from
evolutionary ecology that we believe help marshal what is known about prenatal influences on preg-
nancy, fetal development, and child outcomes into a more coherent whole. We focus in particular
on the concept of maternal effects.
Maternal effects are particularly influential across mammalian species where the interactions
between the mother and offspring are close and over an extended duration. Humans are an extreme
example due to our haemochorial placenta (Mossman, 1987). Through internal fertilization and ges-
tation, the prenatal environment is the maternal domain, and mothers have the most profound influ-
ence on their developing offspring’s phenotype (Clutton-Brock, 1991). An individual’s phenotype is
partly established through the contribution of genes from mother and father and the environment of
development. Therefore, it makes sense that the paternal phenotype also influences prenatal develop-
ment. This is the case in animals where the influence of the father’s phenotype is strong (Smiseth,
Kölliker, and Royle, 2012). In humans the strength and consistency of paternal effects are still being
established. Indeed, many of these paternal effects may work indirectly through the maternal social
environment and phenotype (Figure 5.1). Therefore, in this chapter maternal effects provide a use-
ful perspective for exploring influences of the maternal environment, parental behavior, maternal
physiology, and placental and fetal growth.
Maternal effects are the effects of the mother’s phenotype on her unborn child’s phenotype
independent of the effects of her genotype (Figure 5.1). The mechanisms by which the maternal
phenotype can affect the fetus’s phenotype include her behavior and hormone levels, nutrition,
mental and physical health, and size. For example, across mammalian species smaller mothers tend
to produce smaller offspring, not necessarily because of genetic inheritance but because of other
constraints such as a smaller pelvic outlet or reduced nutrient supply to the fetus. Maternal effects
are important for understanding fetal development and evolution, for they provide a mechanism
for the nongenetic intergenerational transmission of phenotypes (Maestripieri and Mateo, 2009a,
2009b; Smiseth et al., 2012).
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Prenatal Parenting
Local
environment
Maternal
Maternal
development
phenotype
Maternal
genotype
Fetal Fetal Fetal
genotype development phenotype
Paternal
genotype
Paternal
Paternal
phenotype
development
Local
environment
Figure 5.1 Maternal effects are the effects of the mother’s phenotype on her unborn child’s phenotype inde-
pendent of the effects of her genotype. This schematic incorporates the life history theory frame-
work broadening the factors that contribute to maternal effects. The black represents the maternal
effects are particularly influential in mammalian species.
Source: Adapted from Cheverud and Wolf (2009).
Life history theory is the branch of evolutionary ecology devoted to the study of life cycles and
how they change in response to environmental variability. Here we are concerned with the begin-
ning of the life cycle and how the prenatal environment affects fetal growth and development and
the subsequent neonatal phenotype. Like all life history traits, these prenatal traits are manifest in a
“reaction norm”—the range of normal phenotypes that can develop from the same genotype in dif-
ferent environments. In what follows we show how a combination of maternal effects and life history
theory makes sense of the abundant empirical evidence for prenatal influences. We focus on mater-
nal psychosocial stress and nutrition, environmental toxins, and the crucial role of placental physiol-
ogy and epigenetic processes in transducing these maternal effects to the fetus, subsequent postnatal
growth, development, health, and potential for intergenerational, nongenetic inheritance. We aim for
a multi-level synthesis, utilizing a broad evolutionary perspective to examine how organisms respond
to their environment with specific physiological examples and their epigenetic regulation.
We focus on one mechanism of prenatal development that embodies information about a broad
range of maternal environmental factors in the fetus: the hypothalamic-pituitary-adrenal (HPA) axis
(see Gunnar, Doom, and Esposito, 2015; Spencer, 2017). The HPA axis is highly conserved across
vertebrate species and carefully aligns development with environmental stressors in species as distant
to humans as the spadefoot toad (Denver, 1997). Stress hormones influence the expression of 10% of
the genotype across metabolism, growth, repair, and reproduction (Phillips and Matthews, 2011) and
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David A. Coall et al.
are therefore able to adapt whole life cycles to adversity. During pregnancy, one specific mechanism
involved in regulating the impact of maternal stress on the fetus is the enzyme 11beta-hydroxysteroid
dehydrogenase type 2 (11β-HSD-2). This enzyme is involved in limiting the amount of maternal
cortisol, a stress hormone that crosses the placenta from the mother to the fetus. Cortisol itself is
involved in the epigenetic regulation of 11β-HSD-2. Moreover, during pregnancy the expression of
11β-HSD-2 is influenced by maternal stress, nutrition, hypoxia, reproductive hormones, and some
toxins and pollutants. This mechanism interprets a range of maternal signals in ways that tend to
ensure a more appropriate fit between the fetus and the environment into which it will be born than
would otherwise be the case.
We begin our review by examining the role maternal effects have in the interaction between
development and evolution, culminating in modern epigenetics. The next section provides an over-
view of life history theory and parent-offspring conflict theory, focusing on the evolutionary con-
sequences of environmental effects on the timing of life history stages. Then we review the impact
of maternal stress and nutrition during pregnancy, two prominent maternal effects, on fetal develop-
ment. The following section examines the impact of prenatal exposure to toxins on the growing
fetus with focus on pollutants of emerging concern. Finally, we highlight future empirical and theo-
retical pathways suggested by our synthesis.
An Evolutionary-Developmental Framework:
Maternal Effects, Evolution, and Epigenetics
In this section we introduce evolutionary-developmental (“evo-devo”) thinking to frame our discus-
sion of prenatal parenting and development. This perspective combines maternal effects, evolution,
and phenotypic development. The evolutionary-developmental nature of prenatal parenting is illus-
trated through the epigenetic effects of maternal nutrition.
Maternal Effects
Maternal effects are independent of the effect of maternal genes (Figure 5.1), but because they
increase the intergenerational transmission of traits, and thus the fit between the offspring’s phe-
notype and its environment, they can provide an evolutionary advantage (Maestripieri and Mateo,
2009b; Smiseth et al., 2012). Maternal effects are particularly influential in mammalian species
where the interactions between mother and offspring are close and prolonged, especially in humans.
Pleistocene environments (our “environment of evolutionary adaptedness”; Bowlby, 1969, p. 50)
were highly variable (Potts, 1996), but the prenatal environments our ancestral mothers provided
children remained relatively constant; mothers are their unborn child’s environment (Caldji, Diorio,
and Meaney, 2000; Coall, Callan, Dickins, and Chisholm, 2015). Selection is therefore expected
to have favored fetal sensitivity to the effects of the maternal environment on her behavior and
physiology—maternal effects. To be sure, paternal effects exist, but most operate indirectly, through
the father’s effects on the mother. Some direct paternal effects have been reported, but need to be
replicated with paternal age emerging as one of the best documented (Alio et al., 2012; Janecka
et al., 2017).
Although maternal effects have only recently been studied in humans (Bjorklund, 2006), their role
in human evolution should not be underestimated. It is likely that critical characteristics of human
social life evolved from maternal-offspring interactions, possibly beginning before birth, and were
subsequently extended to a general system of prosocial behavior (Brazelton, Koslowski, and Main,
1974; Brown, Brown, and Preston, 2011; Cheverud and Wolf, 2009; Chisholm, 2017; Chisholm,
Coall, and Atkinson, 2016; Hilbrand, Coall, Gerstorf, and Hertwig, 2017).
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Evolution
Evolution results in biological change over time. Charles Darwin (1859) detailed this process with
much empirical evidence and some abstract description, transforming a basic idea of gradual change
over time into something very much more specific. As Godfrey-Smith (2010) noted, many scholars
have tried to continue the abstraction that Darwin began in the later stages of On the Origin of the
Species by Natural Selection. Here is a now-classic example:
Darwin’s scheme embodies three principles:
1. Different individuals within a population have different morphologies, physiologies, and behav-
iors (phenotypic variation).
2. Different phenotypes have different rates of survival and reproduction in different environments
(differential fitness).
3. There is a correlation between parental fitness and offspring fitness (fitness is heritable).
These three principles embody the process of evolution by natural selection. While they hold,
a population will undergo evolutionary change (Lewontin, 1970). Darwin’s own focus fell on evo-
lutionary change within biological systems. Therefore, in terms of maternal effects we can see that
maternal characteristics and behaviors that increase her offspring’s survival and reproduction would
be selected for and increase in frequency in future generations.
In the example above, the term differential fitness is used. From this point, the term inclusive fit-
ness rather than differential fitness is adopted. Inclusive fitness is the sum of direct fitness, achieved
through reproduction, and indirect fitness, achieved through the reproduction of genetic relatives.
Modern evolutionary theory adopts this mid-level theoretical concept to capture life-history dynam-
ics for individuals. The assumption is that organisms act to maximize their average lifetime inclusive
fitness through a series of trade-off decisions across time. These decisions happen at all levels of
biological organization and are captured by formal life history theory, which we discuss in the next
section.
Phenotype From Genotype

Waddington (2012) introduced the notion of the epigenotype and epigenetics to distinguish the com-
plex set of processes that allow the formation of a phenotype from a genotype. “Epigenetics” is
derived from epigenesis, and Waddington used the term to refer directly to developmental processes,
as Pythagoras had before him. Waddington understood these developmental processes as being, to
some extent, under genetic control. The amount of regulatory control could be referred to as the
degree of buffering of an organism’s development relative to the perturbations of genetic and envi-
ronmental variation. More famously, Waddington referred to developmental trajectories as more
or less “canalized” as a consequence of such regulation. The discussion of development within the
context of natural selection made Waddington a pioneer of evolutionary developmental biology
(Haig, 2004).
The information represented in a zygote’s DNA is derived from or is about the environments of
its ancestors, not its current environment. Because environments change more-or-less constantly, this
“old” information is not necessarily a good predictor of the zygote’s future environment. As Wad-
dington (1969) put it, “The main issue in evolution is how populations deal with unknown futures”
(p. 122). Development “buffers” the organism against unknown futures, the vicissitudes of changing
environments, by enabling the developing phenotype to be affected by or to embody “new” infor-
mation about the environment in which it develops. Because selection operates only on phenotypes,
169
not genotypes, environmental effects on phenotypes can create novel phenotypes as targets of selec-
tion. Through canalization, genetic assimilation, epigenetic inheritance, and the many kinds of learn-
ing, novel phenotypes can become heritable. Indeed, developmental plasticity, the capacity of the
phenotype to be affected by the environment in potentially adaptive ways, is an important driver of
evolution (West-Eberhard, 2003).
Modern Epigenetics
Evolutionary developmental biology has re-established itself within biology and more broadly within
behavioral biology (Bateson and Gluckman, 2011; Jablonka and Lamb, 2005; West-Eberhard, 2003).
Use of the term “epigenetics” has markedly increased (Haig, 2012). Histone modification is one
mechanism that allows transcription regulation. The direct addition of a methyl group to cytosine
on the DNA is another—and is most closely associated with the modern concept of epigenetics.
Methylation of cytosine can prevent the binding of transcription factor and suppress transcription.
Ultimately, methylation acts to silence gene expression.
Specifically in terms of maternal effects, there is increasing evidence of perinatal methylation and
a role for this form of regulation in parenting effects and environmental modification of behavio-
ral phenotypes in animal models (Champagne, 2013; Youngson and Whitelaw, 2008). In this way,
characteristics can be acquired or developmentally induced, and their effects transmitted to the next
generation. These effects are the result of proximate epigenetic mechanisms acting to modify gene
expression. In many ways, these mechanisms appear to be adaptations designed to calibrate organ-
isms to their environment (Dickins and Rahman, 2012). There follows an illustration of epigenetic
mediated calibration of an organism to the nutritional environment.
Epigenetic Effects of Maternal Nutrition

Kuzawa (2005) introduced the concept of “intergenerational phenotypic inertia”. On the basis of
extensive evidence that a mother’s birth weight is among the strongest predictors of her offspring’s
birth weight (even more so after controlling for mothers’ gestational age at birth), Kuzawa revisited
the hypothesis that the nutritional experiences of a mother when she was a fetus can affect the
intrauterine nutritional environment that she provides her own offspring, especially her daughters
(Ounsted, Scott, and Ounsted, 1986; Wells, Sharp, Steer, and Leon, 2013).
Kuzawa noted that, among survivors of the Dutch Hunger Winter of 1944–1945 (Stein and
Lumey, 2000), daughters who were exposed to their mother’s undernutrition were significantly
lighter at birth than Dutch girls born between 1944 and 1946 whose mothers were not exposed to
the Hunger Winter. Furthermore, individuals who were prenatally exposed to the famine showed—
six decades later—less DNA methylation of a gene that regulates growth (IGF2) than their unex-
posed, same-sex siblings (Heijmans et al., 2008). Kuzawa’s intergenerational phenotypic inertia
model provided an adaptationist rationale for expecting the effects of prenatal malnutrition or stress
to last more than one generation: When environments are stochastic over time scales greater than
a generation, 9 months of gestation cannot provide the fetus with enough information on which
to “predict” its own, within-generation optimal growth and development (Nettle, Frankenhuis, and
Rickard, 2013). Physiological adaptations to pregnancy that likely match future environments are
encompassed within the “predictive adaptive response” (PAR) model. Therefore, Kuzawa argued,
intergenerational phenotypic inertia provides the fetus with information, not only about the envi-
ronment into which it will be born, but the environment into which its mother was born, and
perhaps even its mother’s mother, and so on, back an unknown number of generations. Intergenera-
tional phenotypic inertia “has the effect of limiting changes in growth rate in response to short-term
ecologic fluctuations, and thus may allow the fetus to cut through the ecologic ‘noise’ of seasonal
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or other stochastic influences to read the ‘signal’ of any longer term nutritional trends in the local
ecology” (Kuzawa, 2005, p. 17).
Conclusion
A likely mechanism through which maternal effects work is Waddington’s view of epigenesis, now
grounded in modern epigenetics. Histone modification and methylation allow a complex and varied
array of alterations to gene expression, which in turn affect the phenotype. As the maternal behavior
example demonstrated, methylation can occur in response to external inputs (in this case the mater-
nal environment), not only affecting the current organism but also future generations. Moreover, the
reversible nature of parental effects make them candidates for facultative (e.g., developmental) adap-
tations. In the following section, we present life history theory and facultative reproductive strategies
as developmental trajectories that may help us organize our understanding of factors that influence
prenatal development.
Evolutionary Theory and the Maternal Environment: Life History

Theory and Parent-Offspring Conflict Theory
A multitude of factors impact maternal and fetal phenotypes during pregnancy. No one discipline
can encompass this diversity. In this section we use two evolutionary perspectives to broaden and
organize our investigation of maternal effects (see Figure 5.1).
Life History Theory

Life history theory is the branch of evolutionary theory devoted to the study of life cycles. It seeks to
understand the evolution of the developmental processes that produce life history traits (e.g., length
of gestation, number and size of offspring, interbirth interval, length of lactation, age and size at
maturity, postreproductive lifespan, total lifespan; see Table 5.1). It views development as an adapta-
tion for reproduction and life cycles as reproductive strategies—naturally selected patterns of growth
Table 5.1 The minimax and maximin reproductive strategies
Reproductive strategy Minimax: Maximin:

(r-strategy) (K-strategy)
maximize current reproduction maximize future reproduction
Ecology more variable and/or unpredictable more constant and/or predictable
Mortality rates often catastrophic, nondirected, more constant, directed, density
density independent dependent
Survivorship low in early life high in early life
Population size more variable, nonequilibrium more constant, equilibrium
Intra- and interspecific competition more variable, lax more constant, intense
Traits favored by selection 1. rapid development 1. slow development
2. early reproduction 2. delayed reproduction
3. high reproductive rate 3. low reproductive rate
4. low parental investment 4. high parental investment
5. small body size 5. large body size
6. semelparity (large litters) 6. iteroparity (small litters)
7. short lifespan 7. long lifespan
Adapted from Pianka (1970) and Reznick, Bryant, and Bashey (2002).
171
and development for maximizing reproductive success under particular ecological conditions (Bon-
ner, 1965; Charnov, 1993; Stearns, 1992; West-Eberhard, 2003). Maternal effects are nongenetic
(Maestripieri and Mateo, 2009b), but the capacity to develop alternative reproductive strategies has
a genetic basis.
Reproductive success is fitness. By definition, fitness is measured in terms of an organism’s reproduc-
tive success (relative to others in its breeding population) but it consists of work—the work required
to stay alive, grow and develop, and reproduce. The latter requires both the production of offspring
(which increases their quantity) and rearing them (i.e., parental investment, which increases their
“quality” [reproductive value; probability of producing grandchildren]). But work requires resources
(e.g., energy, nutrients, time, information), which are sooner or later always limited. Because selec-
tion inexorably favors organisms with greater fitness—but greater fitness requires more resources—
something has to give. Selection is therefore expected to favor organisms’ capacity to allocate their
limited resources to the most pressing adaptive problems posed by their particular ecology—staying
alive, growing and developing, producing offspring, or investing in them. Trade-offs are thus inevita-
ble. Resources allocated to the production of offspring, for example, increases offspring quantity but
reduce the amount parents can invest in each offspring, and thus reduce offspring quality.
The most all-encompassing trade-off is that between current and future reproduction (Stearns,
1992). At issue is whether it would be better for an organism’s lifetime reproductive success to repro-
duce at a given time or to wait for another opportunity in the future. Consider a nursing mother.
Would it be better for her lifetime reproductive success to continue nursing her current child, delay-
ing future reproduction, or to wean now and have another (future) child? Continuing to nurse not
only consumes maternal resources but has diminishing returns for the child’s fitness as it grows. At
some point the lifetime fitness benefits accruing to the mother from continuing to nurse will be
less than those she would receive if she ceased investing in her current child and had another. The
trade-off between current and future reproduction will therefore be optimized according to the local
environment.
The major determinants of the optimal current-future trade-off are (1) the probability of death at
a given age and (2) the availability of energy and other resources that determine parents’ capacity to
invest in offspring. Mathematical modeling and evidence show that when environmental conditions
are risky or uncertain, with high or unpredictable mortality rates and few or uncertain resources,
organisms in general, including humans, tend to reproduce early and often (Coall, Tickner, McAllis-
ter, and Sheppard, 2016). Maximizing current reproduction reduces the chance of lineage extinction
(by maximizing the probability that at least one offspring will survive), but it reduces the amount
parents can invest in each, which only further reduces their probability of producing grandchildren.
However, when conditions have been severe enough, long enough, parents have little to invest any-
way, so downside risk protection against lineage extinction can be evolutionarily rational (Gillespie,
1977; Promislow and Harvey, 1990, 1991; Schaffer, 1983; Stearns, 1992).
At the other end of the continuum, when environmental conditions are safe and predictable, with
low and predictable mortality rates and plentiful, predictable resources, organisms tend to reproduce
later and less often. Minimizing offspring number can be evolutionarily risky, but with low mortality
rates this is not such a problem. By having fewer offspring parents can invest more of their relatively
plentiful resources in each, thereby increasing their quality (growth and development), setting the
stage for the production of grandchildren, great-grandchildren, and so forth in the future (Hill and
Kaplan, 1999; Low, Hazel, Parker, and Welch, 2008; Placek and Quinlan, 2012; Promislow and Har-
vey, 1990, 1991).
In summary, there is no a priori “best” reproductive (life history) strategy. The optimal strategy
in one environment is unlikely to be optimal in another. Humans have a long evolutionary his-
tory of selection for developmental plasticity—the capacity to respond adaptively to environmental
change—due to the rapid fluctuations of Pleistocene environments (Potts, 1996). Our ancestors were
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able to take advantage of good times by maximizing future reproduction (investing more in fewer
offspring) and to cope with bad times by maximizing current reproduction (investing less in more
offspring). None of this, of course, required conscious awareness, and the world’s most disadvantaged
peoples still tend to reproduce early and often (Low et al., 2008).
Parent-Offspring Conflict Theory

The most pressing adaptive problem for unborn mammals has always been to acquire the resources
needed to grow and develop well enough to reach term. Consequently, the placenta (a fetal organ)
has been under more intense selection than perhaps any other organ since it appeared in Eutherian
mammals over 100 million years ago (Power and Schulkin, 2012). The consensus now is that the
intensity of this selection was due to parent-offspring conflict. Fetuses would have been selected to
extract more maternal resources and mothers would have been selected to resist, reserving resources
for future offspring, resulting in positive feedback and a maternal-fetal “arms race” (Haig, 1993, 1996;
Power and Schulkin, 2012; Wildman et al., 2006).
Parent-offspring conflict theory (Trivers, 1974) holds that mothers and offspring—including
unborn offspring—are naturally conflicted. Mothers share 50% of their genes with each fetus, but
each fetus also shares 50% of its genes with its father, making mother-fetus conflict inevitable (i.e.,
the fetus shares 100% of its genes with itself ). The fetus is thus expected to seek more parental invest-
ment than would be optimal for mothers to provide because they seek to benefit themselves (copies
of both parents’ genes). At the same time, mothers are expected to provide fewer resources than would
be optimal for the fetus because mothers, too, seek to benefit themselves through copies of their
genes in current or future offspring.
Haig (1993, 1996) used parent-offspring conflict theory to model maternal-fetal interactions
(see Figure 5.2). When conditions are good, mother-fetus conflict is minimal because each can
afford to give a little. But when maternal resources are limited by environmental risk and uncer-
tainty, any maternal investment in the fetus entails a trade-off. For every benefit a fetus gains from
maternal resources (B in Figure 5.2), there will be a correlated cost to its existing or future siblings
(C in Figure 5.2). Because the mother is equally related to all of her offspring, current and future,
it is evolutionarily rational for her to invest equally in each according to their capacity to benefit
from the resources invested. Therefore, mothers are expected to seek the best possible balance
between the benefit to the existing fetus and the cost to its existing or future siblings. In other
words, at some point (x1 in Figure 5.2), the increased benefit to the current fetus from mother’s
continuing investment, will be outweighed by reducing the resources she could invest in existing
or future offspring.
Maternal-fetal conflict and the trade-off between offspring quantity and quality mean that fetal
adaptations to the effects of environmental stress on the mother can have evolutionarily adaptive con-
sequences for her (future reproduction) but developmentally disadvantageous effects on fetal growth
and development and thus postnatal health. The intergenerational consequences may be related to
the fact that, despite our species’ extraordinary reproductive success, human gestation is sensitive to
the environment and characterized by high rates of early miscarriage, nausea and vomiting, and con-
ditions such as preeclampsia, maternal hypertension, gestational diabetes, and preterm birth.
Prenatal Stress, Biological Mechanisms, and Developmental

Outcomes in Offspring
The prenatal period is a time of rapid cell division and differentiation. As with other periods of rapid
growth, environmental insults can disrupt development. The uterus is an adaptation for support-
ing fetal growth and buffering the fetus against challenges that affect the mother. Here, we use the
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Cost
Cost to Siblings (C)

Benefit to fetus (B)
Benefit
x1 x0
Maternal investment in fetus (x)
Figure 5.2 Representation of the trade-off between maternal investment in the current fetus versus existing or
future siblings (x1 maternal optimum, x0 current fetus’s optimum).
Source: Adapted from Haig (1996).
perspective of human evolutionary ecology to examine the impact of a range of stressors on devel-
opment, including stressors in the maternal psychosocial environment, nutritional stressors, disease
pressures, and exposure to toxicants in the local environment. We focus on these changes as poten-
tially adaptive responses to environmental change that in some cases produce pathology.
Developmental Plasticity
Plants and animals that develop in heterogeneous environments typically have the ability to alter
their phenotypes in response to those environments (Bateson, 2017; Via and Lande, 1985). The
response of an individual’s phenotype to this environmental variability is referred to as phenotypic or
developmental plasticity (Stearns, 1982; West-Eberhard, 2003). Human developmental plasticity ena-
bles children to make unconscious “bio-assays” of their physical environments (Ellison, 1990, 1996)
and “socio-assays” of their social environments (Chisholm, 1999a; Draper and Harpending, 1982). If
these assays accurately predict the adult environment, they are likely to provide a fitness advantage
(Bateson et al., 2004; Hill and Kaplan, 1999).
Phenotypic changes (e.g., physiology, growth, mental health) induced by the environment are
often presented as being programmed (Whimbey and Denenberg, 1967). We will use such terminol-
ogy, but with caution. It is important to keep in mind that these changes are latent potentials, part
of the individual’s norm of reaction, and are elicited by the environment, not created by it (Bateson,
2007). Moreover, these phenotypic changes, even in sensitive periods of development, are often
modifiable (Bornstein, 1989), which ultimately keeps them open to intervention.
Across multi-cellular species, an organism’s developmental environment can have consequences
for its own growth, development, and reproduction and for those of its descendants (Bateson et al.,
2004; Coall and Chisholm, 2010; Kuzawa, 2005; West-Eberhard, 2003). Evolutionists have focused on
the intrauterine environment, the broad range of challenges that influence the uterine environment,
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Prenatal Parenting
and their consequences for fetal development and subsequent health: maternal effects on offspring
phenotype (Wells, 2007).
Reduced fetal growth is associated with higher rates of perinatal morbidity and mortality (Kramer,
1987) and an increased risk of developing coronary heart disease, hypertension, and diabetes in adult
life (Barker, 1994, 2004). Experimentally inducing fetal growth restriction in animal models (via the
administration of synthetic glucocorticoids, inducing maternal stress, controlling the maternal diet, or
reducing placental function) results in cardiovascular, metabolic, and endocrine changes throughout
childhood and even into adulthood (McMillen and Robinson, 2005). This burgeoning focus on the
intrauterine environment has culminated in a new paradigm, the Developmental Origins of Health
and Disease (DOHaD). Within the DOHaD paradigm, there has been a particular focus on birth
weight, placental weight, and placental weight to fetal weight ratio (hereafter placental ratio) as indi-
cators of the quality of the uterine environment. We examine these below.
Fetal Growth
It is well known that reduced fetal growth predicts subsequent morbidity and mortality. In a review
of the epidemiological literature, Kramer (1987) identified a number of maternal factors associated
with intrauterine growth restriction (IUGR), including demographic and psychosocial elements of
the mother’s environment, obstetric factors, nutritional status, morbidity during pregnancy, toxic
exposures, and prenatal care. Placental efficiency plays an integral role in two of the three main causes
of reduced fetal growth, which are (1) abnormal placental structure and/or function; (2) inadequate
maternal supply of oxygen and/or nutrients due to either maternal factors or placental factors; and
(3) decreased ability of the fetus to use the available supply (Brodsky and Christou, 2004).
Increasingly, the role of maternal constraints on fetal growth—and even that of future generations—
are receiving attention (Gluckman and Hanson, 2008; Kuzawa, 2005; Lewis, Cleal, and Hanson,
2012; Ounsted et al., 1986; Wells, 2003). For example, in a study of 513 low-risk pregnancies, Coall,
Charles, and Salafia (2009) found that maternal birth weight was the only factor that consistently
predicted children’s fetal and placental growth, affecting outcomes including birth weight, placental
weight, placental ratio, placental surface area, and placental thickness.
Birth weight is a crude measure of the quality of intrauterine environment (Salafia et al., 2008;
Wilcox, 2001). From the perspective of life history theory, fetal growth is a measure of resource flow
to the fetus (parental investment). A more direct measure of resource flow is provided by propor-
tionate birth weight. Proportionate birth weight is the neonate’s birth weight as a proportion of the
mother’s weight and represents the amount of parental investment made during gestation (May and
Rubenstein, 1985). This characteristic remains remarkably constant at 5% across all placental mam-
mals (Land, 1985). Proportionate birth weight has been used as a direct measure of resource flow to
the fetus and, therefore, of parental investment throughout gestation (Coall and Chisholm, 2010).
A range of insults have a measurable impact on the uterine environment and health in the next gen-
eration. However, much of the uterine environment is buffered against the maternal environment
through placental adaptation.
Placental Weight and Placental Ratio

The placenta is the common filter through which maternal effects pass. Placental growth and devel-
opment are strongly correlated with the growth trajectory of the fetus and have a major influence on
birth weight (Salafia et al., 2005, 2008). The placenta has evolved to provide a protected environment
to support fetal growth, buffering the developing organism from maternal effects, environmental
insults (Lewis et al., 2012; Wooding and Burton, 2008), and preparing it for the postnatal environ-
ment it is likely to inhabit (Gluckman, Hanson, Spencer, and Bateson, 2005).
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The functional efficiency of the placenta may be challenged by disturbances in the maternal envi-
ronment, such as poor nutrition, obesity, illness (e.g., diabetes, hypertension), smoking, medications,
and maternal stress (Sibley, Brownbill, Dilworth, and Glazier, 2010; Tegethoff, Greene, Olsen, Meyer,
and Meinlschmidt, 2010). Other known modulators include maternal factors, such as country of
origin, weight at birth, height, age, and parity (Kiserud et al., 2017). Although beyond the scope of
this chapter, male and female placentae consistently show different physiological responses to changes
in the maternal environment (Terrade, Panchenko, Juneir, and Gabory, 2015; Rosenfeld, 2015). The
effect of such disturbances may create adverse intrauterine conditions resulting in adaptations (that
are often under endocrine control) by the placenta (Coan et al., 2010; Myatt, 2006). The placenta, as
a metabolically active tissue, also plays a role in sensing the flow of nutrients, modulating the alloca-
tion of resources between the mother and fetus to maintain pregnancy (Dimasuay, Boeuf, Powell,
and Jansson, 2016). Facultative metabolism provides the placenta with the capacity to adapt to chal-
lenges, for example by increasing surface area and signaling pathways to extract more nutrients when
maternal nutrient levels fall (Sibley et al., 2010). In the long term this adaptation may be associated
with poorer adult health outcomes (Dimasuay et al., 2016); however, when the placenta is not able
to adapt in the short term, adverse outcomes for the fetus can result (Sandman and Davis, 2010).
In epidemiological studies, the role of the placenta is often extrapolated from placental weight.
Placental weight is generally associated with placental function (Robinson, Owens, de Barro, Lok,
and Chidzanja, 1994). Like birth weight (Godfrey, Breier, and Cooper, 1999; Harding, 2001; Wilcox,
2001), placental weight is the end product of a chain of events that include the growth and develop-
ment of the placenta throughout gestation (Barker and Thornburg, 2013; Coall et al., 2009; Salafia
et al., 2005). As with other aspects of biology, early beneficial adaptations that improve survival can
have later costs in morbidity and shortened lives (Kirkwood and Austad, 2000). DOHaD research
has shown that a high placental ratio is associated with higher blood pressure throughout childhood
(Moore et al., 1996) and into adulthood (Moore, Cockington, Ryan, and Robinson, 1999) and may
predispose individuals to adult illnesses, such as hypertension, glucose intolerance, blood coagulation
disorders, and coronary heart disease (Barker, 1997; Barker, Bull, Osmond, and Simmonds, 1990;
Forsén, Eriksson, Tuomilehto, Osmond, and Barker, 1999; Forsén et al., 1997; Law, Barker, Bull, and
Osmond, 1991; Phipps et al., 1993). Some researchers, however, have found a U-shaped association
between placental ratio and coronary heart disease in men (Martyn, Barker, and Osmond, 1996).
Others have found an association between placental weight, but not placental ratio, and subse-
quent blood pressure (Blake et al., 2001; Whincup, Cook, Papacosta, and Walker, 1995; Williams,
St George, and Silva, 1992), some have found sex differences (Taylor, Whincup, Cook, Papacosta,
and Walker, 1997), and others have found no effects of gender (Burke et al., 2004; Leon et al., 1996;
Martyn et al., 1995; Matthews, Lewis, and Bethel, 1994). It may be that a relatively large placenta does
not directly cause adult disease but reflects the quality of the fetal supply line throughout gestation
(Robinson et al., 1995). However, animal and human studies have both shown that placental size and
placental ratio play important roles in fetal adaptation to the maternal environment (Faichney and
White, 1987; Godfrey, 2002; Kingdom, 1998; Robinson et al., 2001; Steyn et al., 2001).
In remarkable accord with the theoretical predictions from Haig’s (1993) application of parent-
offspring conflict theory to the uterine environment, the DOHaD perspective proposed that
increased placental ratio was an adaptive placental response to intrauterine growth restriction (Barker
et al., 1990). Initial studies involving sheep showed that moderate dietary restrictions at various times
during pregnancy were associated with increased placental weight (Faichney and White, 1987). The
placental growth was interpreted as “an attempt by the fetus to compensate for the reduced supply of
nutrients in the maternal blood” (Faichney and White, 1987, p. 373). This compensatory mechanism
maintained fetal growth in the ewes that had their diets restricted late in pregnancy, and may actually
have increased the resource flow to the fetus when the dietary restriction was removed. It must be
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Prenatal Parenting
recognized, however, that, although similar responses have been documented in humans, the picture
is over-simplified. Importantly, the timing of dietary restriction and the previous nutritional state of
the ewes are known to influence placental responses (Barker et al., 1993; Robinson et al., 2001). In
a natural experiment, Lumley (1998) reported that women conceiving or in the first trimester of
pregnancy during the Dutch Famine Winter of 1944–1945 had relatively heavy placentae, while
birth weights remained static. Lumley interpreted this finding as compensatory placental growth in
response to reduced maternal nutrition. Both animal (Robinson et al., 1994) and human models
(Wheeler et al., 1994) supported this interpretation. These findings reflect the importance of the
timing of maternal effects.
Environmentally Appropriate Adaptation, Not Pathology

This application of evolutionary theory challenges the common perception that increased placen-
tal weight is pathological (Lao and Wong, 1999, 2001). For example, infants born at high altitudes
have reduced birth weights (Mayhew, Jackson, and Haas, 1990), higher placental weights (Kruger
and Arias-Stella, 1970), and show structural changes that improve the efficiency of oxygen transfer
between mothers and fetuses (Mayhew et al., 1990; see review by Zamudio, 2003). This placental
adaptation can improve fetal growth (Mayhew, Jackson, and Boyd, 1993). Moreover, there is evidence
that increased placental size can improve fetal survival. In appropriate-for-gestational-age infants of
insulin-dependent diabetic mothers, a higher placental ratio is associated with an increased likelihood
of offspring survival (Evers, Nikkels, Sikkema, and Visser, 2003). Increased placental size in insulin-
dependent diabetic pregnancies are likely to reflect the larger placenta necessary to maintain a fetus
when the placenta is less efficient (Lao and Ho, 2002). More generally, placental adaptation to the
uterine environment, which maintains fetal growth, is a normal process of pregnancy (Pardi, Mar-
coni, and Cetin, 2002). Therefore, pregnancies in which the placentae show evidence of compensa-
tory growth resulting in increased placental ratios may actually represent appropriate responses to
suboptimal maternal/uterine environments of diverse origin in the short term, rather than pathology
itself (Adair and Thelander, 1925; Fox, 2000; Kaplan, 2008).
DOHaD, as the name suggests, is interested in this developmental plasticity because of its impact
on health. Differential reproductive success, however, is what drives evolution and placental func-
tion is crucial for successful reproduction (Lewis et al., 2012). Postnatal pathology arises when the
trade-offs with other components of fitness entrain developmental trajectories that increase short-
term survival at the cost of ill-health and premature mortality in the future. These costs include
low birth weight, preterm birth, altered stress reactivity, rapid childhood growth, early reproductive
development, and short lifespans. If these costs minimize the chance of lineage extinction—that is,
if they increase the chance of reproducing at all in the harsh or unpredictable environments that
cause them—they may also be evolutionarily rational (Borgerhoff Mulder, 1992; Chisholm, 1999a;
Harpending, Draper, and Pennington, 1990; Stearns, 1992). In summary, life history theory views
many of these changes as adaptations to environmental challenges, including maternal effects that
maximize survival and, ultimately, reproduction.
We turn now to the impact, first, of maternal stress on early development, second, the role of
maternal nutrition, and, last, the increasingly apparent role of toxicants in the environment. We begin
by discussing the meaning of “stress”.
Stress, Stressors, and Adaptive and Maladaptive Coping

Environmental stress affects development and does so in potentially adaptive ways by entraining
alternative reproductive strategies (Belsky, Steinberg, and Draper, 1991; Chisholm, 1993; Worthman,
177
1999). “Stress” has long been a slippery concept, with considerable disagreement about its mean-
ing (Cohen, Kessler, and Gordon, 1995; McEwen, 1995). We view stress generally as a broad range
of environmental challenges, including social-emotional or psychosocial, inadequate nutrition, and
toxicants. We focus first on the psychosocial stress entrained by the perception of negative life events,
a fairly objective measure of environmental risk and uncertainty (Cohen et al., 1995). We refer to
the other stressors as biological stressors. These are factors such as malnutrition, disease, toxicants, and
material poverty and include the physiological and energetic stressors associated with them (Ellis,
2004; Ellison, 1990; Thayer and Kuzawa, 2011). Therefore, biological stress represents situations of
reduced biological resource availability. It must be emphasized that, although we treat psychosocial
and biological stressors as independent factors, they should not be seen as mutually exclusive because
there is considerable interplay between them and they are often inseparable (Chisholm, 1999a; Coall
and Chisholm, 2010). The survival, growth and development, and reproduction of all organisms
always depend on biological resources (e.g., nutrition, disease), but because of our species’ reliance
on cooperation, social exchange, and sharing to gain access to these resources, our life history, as with
most other primates, depends on the availability of psychosocial resources.
Any analysis of psychosocial stress must consider individual differences in the ability to cope with
or adapt to stressors (Ellis, 2004). Paramount for our intensely social species is the social-emotional
support that can dramatically enhance individual coping styles, life satisfaction, and sense of control
(Compas, Slavin, Wagner, and Vannatta, 1986; Steptoe and Marmot, 2003; Suldo and Huebner, 2004;
Thoits, 1982). These psychosocial resources serve as buffers against the cumulative cost of constantly
adapting to environmental challenge (allostatic load; McEwen, 1995) and are known to reduce the
impact of stressful life events on adult health (Compas et al., 1986; Greenberg, Seltzer, Krauss, Chou,
and Hong, 2004; Kaplan, Cassel, and Gore, 1977; Patterson and McCubbin, 1984). Stress has a direct
effect on adult outcomes, such as mental health, and an indirect influence via the impact it has on
relationships that constitute these psychosocial resources (McEwen, 2000; Thoits, 1982). Everything
else being equal, an increase in the level of psychosocial stress results in a reduced availability of psy-
chosocial resources (Ellis, 2004; Taylor and Seeman, 1999). Therefore, psychosocial stress not only
puts wear and tear on an individual’s physiology but also consumes valuable psychosocial resources.
From an evolutionary perspective, coping with psychosocial stress is work, and like any other work,
it requires resources, in this case, social-emotional resources.
In adulthood, the social support provided by family and friends is important for adjustment to life
events (Runtz and Schallow, 1997). Across cultures, social support and social networks are associated
with less depressive symptomatology during pregnancy and after childbirth (Byrd-Craven and Mas-
sey, 2013; Surkan, Peterson, Hughes, and Gottlieb, 2006). Although the evidence is not conclusive
(Bryce and Stanley, 1991), social support interventions during pregnancy can reduce the rate of low
birth weight births (Norbeck, DeJoseph, and Smith, 1996). The association between social support
and birth weight appears to operate through improved fetal growth rather than longer gestation
(Feldman, Dunkel-Schetter, Sandman, and Wadhwa, 2000; Rothberg and Lits, 1991). In a study of
3,073 low-income women receiving general psychosocial services during pregnancy, social support
was associated with a reduced risk of delivering low birth weight babies (Zimmer-Gembeck and
Helfand, 1996). It is clear that people employ both adaptive (e.g., social support) and maladaptive
(e.g., alcohol and nicotine) coping mechanisms. The interacting influence of prenatal stress and social
support on fetal growth and development is mediated by poor maternal coping mechanisms that
include risky health behaviors (Orr et al., 1996; Zuckerman, Amaro, Bauchner, and Cabral, 1989).
Lack of social support is an important determinant of smoking (McCormick et al., 1990; Rodriguez,
Bohlin, and Lindmark, 2000) and prenatal weight gain (Hickey, 2000). Maladaptive coping strategies
elevate stress hormones, influence nutrition and exposure to toxins during pregnancy, and lead to
poor fetal development (Sandman and Davis, 2010).
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Maternal Stress During Pregnancy and Early Development

Before Barker’s work in the late 1980s, the question was whether stress during pregnancy could
affect fetal or child development (Bryce and Stanley, 1991). However, because of his work and rapid
progress in DOHaD, the questions now are: how does maternal stress affect the developing fetus or
child, what are the mechanisms whereby these early effects might predispose poor adult health, and to
what extent do these adult effects constitute evolutionarily adaptive “programming” of the fetus’ or
child’s developing physiology and organ systems by the maternal environment? (Sandman, Davis,
Buss, and Glynn, 2012). It is now established that across mammalian species prenatal stress affects off-
spring development in response to changes in maternal resource provision (Berghänel, Heistermann,
Schülke, and Ostner, 2017).
There is now abundant evidence that maternal psychosocial stress during pregnancy is a risk
factor for abnormal pregnancy/delivery, fetal abnormalities, and poor adult health. The effects of
stress on the mother include the premature rupture of membranes, preeclampsia, excessive weight
gain, and preterm labor and birth (Dole et al., 2003; Newton, Webster, Binus, Maskrey, and Phillips,
1979); those on the fetus include reduced fetal growth and low birth weight (Newton and Hunt,
1984; Wadhwa, Sandman, Porto, Dunkel-Schetter, and Garite, 1993), which in turn are associated
with increased infant and child morbidity and mortality, impaired psychological development, and
increased morbidity in adulthood (Istvan, 1986; Tegethoff et al., 2010).
Most research into the effects of maternal stress during pregnancy has used either subjective
self-ratings of anxiety/depression or more objective measures of stressful life events (e.g., cortisol).
Because the two measures do not predict the same outcomes, different pathophysiology pathways
seem to be involved (Richardson, Zorrilla, Mandyam, and Rivier, 2006; Tegethoff et al., 2010). For
this reason alone, research on prenatal influences must be multivariate, incorporating (1) objective
indices of stressful life events, (2) individuals’ subjective impressions of their stressors and their mean-
ing, (3) their own and others’ responses to these stressors, and (4) the biochemical, physiological,
neuroendocrine, and immunological mechanisms involved.
Biological Mechanisms of Maternal Stress

The steroid hormone cortisol is a key component of the HPA axis, the “fight or flight” response
of all vertebrates. It has two evolutionary functions, one short term, the other long term. Its short-
term function is to maximize the probability of survival in the face of some immediate threat. For
example, negative life events typically (but not always, as is commonly thought) activate the HPA
axis that stimulates the adrenal cortex to release glucocorticoids (cortisol). This response is adaptive
in the short term in that it helps to maintain homeostasis in the face of environmental (including
social-emotional) challenges (Cohen et al., 1995; McEwen, 1995; Repetti, Taylor, and Seeman, 2002;
Selye, 1957; Tsigos and Chrousos, 2002; Wingfield and Sapolsky, 2003). Its long-term function is
developmental: Cortisol is essential for immune function, glucose metabolism, and fetal (e.g., brain)
development and the maturation of fetal organs (e.g., lungs; Lupien, McEwen, Gunnar, and Heim,
2009; Reynolds, 2013). Growing up in a chronically risky or uncertain environment entails chronic
threats, chronic psychosocial stress, and chronic HPA activation. In chronically risky and uncertain
environments the optimal reproductive strategy is to maximize current reproduction through accel-
erated development and early childbearing. Chronic psychosocial stress and high levels of cortisol
“predict” risky or uncertain environments and may help to entrain evolutionarily adaptive develop-
mental responses (Finch and Rose, 1995; Worthman, 1999; Worthman and Kuzara, 2005).
The stress response begins with the hypothalamus, which integrates the nervous and endocrine
systems. Stress, defined as any challenge to the body’s homeostasis (McEwen, 1995; Selye, 1957),
179
activates the HPA axis. In response to a stressor, the hypothalamus releases corticotrophin-releasing
hormone (CRH), a peptide hormone that travels through local blood vessels and binds to receptors
on the plasma membranes of cells in the anterior pituitary gland. This stimulates the production of
adrenocorticotropic hormone (ACTH), which enters the blood and travels around the body, hav-
ing its primary influence on the cells of the adrenal gland. ACTH acts on the cortex of the adrenal
gland to stimulate production of glucocorticoids (GCs), steroid hormones that include cortisol and
corticosterone (Gunnar et al., 2015; Nakamura, Sheps, and Arck, 2008).
The hypothalamus also directly influences the adrenal medulla via the sympathetic nervous
system. Neurons of the sympathetic nervous system from the hypothalamus synapse in the adrenal
medulla and release the hormones epinephrine and norepinephrine. These hormones stimulate
rapid, organism-wide responses to a stressor in the form of an elevation in heart rate, respiratory
rate, and release of energy reserves in preparation for quick responses to environmental threats
(Sapolsky, 1994).
Because steroid hormones can cross the plasma membranes of cells, the effects of glucocorticoids
are mediated by glucocorticoid receptors (GRs), which have been identified in the cells of almost
all tissues within the body. GCs regulate a variety of important functions and are essential for life in
adult mammals. Excessive levels of GCs, however, potentially affect many internal systems by dys-
regulating biological homeostasis (Nakamura et al., 2008).
Fetal exposure to GCs and CRH is essential for fetal growth and development, but excess levels
can be detrimental. During pregnancy, due to the elevation of maternal CRH levels, maternal GC
production and secretion are increased. High levels of GC hormones are necessary due to their
potent and long-term effects on cellular function in almost all organ systems, particularly in regard
to cellular differentiation and homeostasis (Burton and Waddell, 1999). Later in pregnancy, exposure
to GCs is essential for the maturation of fetal organs and systems (Ellman, Dunkel Schetter, Hobel,
Glynn, and Sandman, 2008). Throughout pregnancy, while fetal and maternal neuroendocrine func-
tions are largely independent, with placental barriers to most maternal hormones, they still interact
(Lester, Marsit, Conradt, Bromer, and Padbury, 2012). Approximately 10%–20% of maternal GCs
cross the placenta to the fetus (Seckl and Meaney, 2006).
Gitau, Cameron, Fisk, and Glover (1998), examining the relation between maternal and fetal cor-
tisol levels, showed that, while fetal levels are lower than maternal levels (indicating the metabolism
of 80%–90% of maternal cortisol by the placenta), the two are correlated, but not with gestational
age. Even so, a 10%–20% contribution of maternal cortisol to the fetus could still double fetal con-
centrations. This study also measured the amount of unmetabolized cortisol crossing the placenta
at 15%. Although most cortisol is metabolized by the placenta, high maternal concentrations can
affect fetal concentrations because 40%–50% of fetal cortisol is derived from the mother (Gitau et al.,
1998). Cortisol also plays a central role in fetal programming of adult disease (Harris and Seckl, 2011;
Reynolds, 2013; Seckl and Holmes, 2007).
Circulating GCs that potentially influence the tissues of the body are regulated by the 11β-HSD
enzymes: 11β-HSD-2 inactivates cortisol, converting it to the inactive cortisone; conversely, 11β-
HSD-1 activates cortisone back into cortisol, increasing local CG levels in tissues. In the placenta,
circulating maternal GCs, including cortisol, are metabolized by 11β-HSD-2 into its inactive form,
cortisone (Benediktsson, Calder, Edwards, and Seckl, 1997; Brown et al., 1996). These placental
enzymes catalyze and modulate ligand access to the glucocorticoid receptor (GR). Therefore, the
11β-HSD enzymes are referred to as the feto-placental barrier to maternal GCs. Because GCs are
potent and have long-lasting effects, changes in the maternal environment that increase GC exposure
may persist long after the original insult, thus acting on tissue accretion, differentiation and program-
ming of the fetal HPA axis (Fowden, Giussani, and Forhead, 2005), changing baseline levels for
physiology, metabolism, and behavior (Lester et al., 2012; Seckl and Meaney, 2006).
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Edwards, Benediktsson, Lindsay, and Seckl (1993) proposed that low levels of 11β-HSD-2, and
concomitant over-exposure to maternal cortisol, reduced fetal growth and programmed adult dis-
ease risk. Since then, it has been shown that overexposure of GCs to the fetus reduces fetal growth,
increases the risk of developing hypertension and metabolic disorders, and inhibits fetal HPA axis
development (Burton and Waddell, 1999; Seckl, 1997). Moreover, Stewart, Rogerson, and Mason
(1995) showed a positive association between placental 11β-HSD-2 activity and birth weight. In
humans, 11β-HSD-2 activity is inhibited by medical disorders involving increased fetal exposure
to cortisol, such as pre-eclampsia, maternal asthma, and intrauterine growth restriction (O’Donnell,
O’Connor, and Glover, 2009). Moreover, a deficiency in 11β-HSD-2 is associated with fetal growth
restriction and increased risk of hypertension in adulthood (White, 2001). Maternal stress down
regulates 11β-HSD-2 in both animal and human studies (Glover, Bergman, Sarker, and O’Connor,
2009; Mairesse et al., 2007). Thus, elevated levels of maternal stress can down regulate 11β-HSD-2
activity, allowing more maternal cortisol to cross the placenta and reduce fetal growth.
Maternal environments that activate the HPA axis, thereby raising cortisol levels, can also affect
CRH levels and lead to preterm birth. Questionnaire studies linking measures of perceived stress,
stressful life events, racism, and domestic violence with preterm birth suggest that maternal stress
shortens gestation (Istvan, 1986). Increased levels of stress hormones have also been associated with
preterm birth (Wadhwa et al., 1993). Rather than involving GCs suppressing CRH, as in the hypo-
thalamus of nonpregnant individuals, the relation between GCs and CRH during pregnancy is
stimulatory, with levels of both rising throughout gestation (Ellman et al., 2008; Reynolds, 2013;
Sandman and Davis, 2010). In response to maternal cortisol, the feto-placental unit activates the pla-
cental CRH gene, increasing placental CRH synthesis and secretion to the fetus. This stimulates the
maternal pituitary to release ACTH, and thus cortisol from the maternal adrenals. Maternal cortisol
again stimulates CRH, creating a positive feedback loop and potentially high fetal exposure to GCs.
By the end of pregnancy, cortisol levels are up to three times higher than nonpregnant levels.
High levels of CRH are also involved in the cascade of events resulting in uterine muscle contrac-
tion and parturition, thus preterm birth. Preterm birth is directly correlated with low birth weight
and suboptimal development of the brain and other tissues and organs. Resulting adverse outcomes
include structural changes in the brain, negative temperament, impaired cognition, and the devel-
opmental programming of adult pathophysiologies such as metabolic and cardiovascular diseases
(Mairesse et al., 2007; Reynolds, 2013; Sandman and Davis, 2010).
From our evolutionary perspective, however, maternal stress-induced preterm birth, low birth
weight, and their sequelae may not represent pathology so much as the adaptive response by mother
and fetus to their respective risky or uncertain environments (Ellman et al., 2008), for three rea-
sons. First, elevated maternal stress hormones, whether due to psychosocial stress, undernutrition, or
exposure to environmental toxins, may accelerate maturation of fetal organs and systems in prepara-
tion for a “predicted” shorter gestation. Second, in high-risk environments, where the probability
of survival for mother and fetus is reduced, maternal-fetal competition for resources increases and
maternal stress hormones may allocate resources away from the fetus to the mother, resulting in fetal
growth restriction but increasing the mother’s chance for “predicted” future reproduction (Haig,
1993; Stearns, 2005). Third, mother and fetus may “versusadapt” to each other as a unit, compromis-
ing on the allocation of maternal resources (Kölliker, Royle, and Smiseth, 2012) so that nutrient flow
to the fetus simply reflects maternal condition and access to resources (Coall and Chisholm, 2003;
Dimasuay et al., 2016; Jansson and Powell, 2013). In all of the above, apparently detrimental mater-
nal stress effects on the fetus might represent adaptive fetal programming for survival and growth
and development in the risky and uncertain environment that caused the mother’s stress in the first
place—and therefore “predict” that such risk and uncertainty will continue in the future (Gluckman
et al., 2005).
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Stress During Pregnancy and Developmental Outcomes

The human capacity for developmental plasticity means that maternal prenatal stress can have del-
eterious effects on fetal development and future health and/or evolutionarily adaptive programming
effects. The nature and duration of the maternal stress, together with the timing of impact during
gestation, determines the severity of fetal developmental outcomes, depending on the stages of devel-
opment and the number of organs and systems affected (Fowden et al., 2005). We review below a
number of factors associated with stress during pregnancy to illustrate the diverse impacts prenatal
stress can have.
Fetal brain development commences during the third week post-conception with structural
development of the central nervous system (CNS) extending to week 20 of pregnancy, providing
ample opportunity to incur insults that may result in impaired development (Kurjack, Medic, and
Salihagic-Kadic, 2004; Moore, 1989). There is strong evidence that stress has an impact on fetal
development of the brain and CNS, including effects on brain morphology, receptor density and
sensitivity, CNS function, and the activity of the autonomic and neuroendocrine systems (Stanton,
Lobel, Sears, and DeLuca, 2002). GCs play an important role in the survival and maturation of
CNS neurons as different regions of the brain express GRs at different stages of development, ena-
bling selective effects of stress throughout gestation (Lupien et al., 2009; Nyirenda, 2006; Seckl and
Meaney, 2006).
The fetal brain also appears to silence 11β-HSD-2 expression and activity during gestational
weeks 19 to 26, indicating the role of GCs in aspects of brain development (Nyirenda, 2006). Animal
studies show that excessive GC exposure in utero influences hippocampal and related complex func-
tions, including cognition, behavior, memory, versusordination of autonomic activity, and impor-
tantly, the regulation of the endocrine systems (Seckl and Meaney, 2006).
These programming effects on the hippocampus include fetal HPA axis deregulation. The HPA
axis is sensitive to excessive GC exposure in utero, with repeated animal studies showing that exces-
sive GC exposure during development permanently alters HPA axis function, accounting for exag-
gerated HPA axis reactivity in adulthood (Nyirenda, 2006). The hippocampus expresses both types
of corticosteroid receptors (GRs). There is also evidence that increased levels of GCs are associated
with decreased receptor expression in the hippocampus, resulting in hypersensitivity and increased
reactivity of the HPA axis (Nyirenda, 2006; Seckl and Meaney, 2006). We turn now to the evidence
that the impact of GCs on the CNS and HPA axis reactivity translates into changes in postnatal
behavior, cognition, and mental health.
Behavior, Cognition, and Mental Health

Mounting evidence shows that overexposure to GCs in utero leads to modifications in adult behavior
in a number of species (Seckl and Meaney, 2006). In humans, prenatal exposure to elevated maternal
stress hormones is associated with behavioral and emotional disturbances during both infancy and
childhood (Sandman and Davis, 2010). In one of the first studies of the impact of prenatal stress
on the subsequent behavioral adjustment of children, O’Connor, Heron, Golding, Beveridge, and
Glover (2002) found, in a longitudinal sample of 7,448, that women who were in the highest 15%
of the sample for anxiety at 18 or 32 weeks of pregnancy had children who were 2 to 3 times more
likely to be 2 standard deviations above the sample mean for total behavioral and emotional problems
at 4 years of age. The association between prenatal maternal anxiety and filial behavioral problems
remained after controls for birth weight, potential maternal confounders (age, socioeconomic sta-
tus), pregnancy related anxiety, behavioral mediators (smoking and alcohol use), and postnatal (8
weeks) depression and anxiety. This adjusted analysis suggests that, barring the potential influence
of maternal weight and postnatal parenting practices, the impact of prenatal anxiety was direct (i.e.,
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not through fetal growth, maternal behavioral modifications, or postnatal anxiety and depression).
Prenatal anxiety and postnatal depression both independently predicted childhood behavioral and
emotional adjustment problems, and this effect persisted to 7 years of age (O’Conner et al., 2002;
O’Conner, Heron, Golding, and Glover, 2003). Unfortunately, the authors did not control for birth
length, which (in a later study with the same sample) was shown to be negatively associated with
total behavior difficulties, hyperactivity, and conduct problems at 7 years of age (Wiles et al., 2006).
Subsequent to these studies, a veritable feast of investigations has examined these associations in
detail and analyzed the physiological mechanisms involved. In a review of 32 studies, Korja and
colleagues (2017) found a consistent association between maternal prenatal stress (or anxiety) and a
child’s negative emotional reactivity or self-regulation before 2 years of age. A likely mechanism is
re-programming of the fetal HPA axis in response to maternal stress.
Maternal prenatal stress and cortisol may re-program the postnatal HPA axis, influencing adult
phenotypes. Maternal cortisol levels, and by association maternal stress and nutrition levels, increase
the activation of the HPA axis postnatally. In mothers and their full-term infants, Davis, Glynn,
Waffarn, and Sandman (2011) found an increased cortisol reaction to the newborn heel prick test
(blood draw) in the neonates of women who had higher cortisol levels during the second half of
pregnancy. This association remained even after adjustment for maternal medical history, socio-
economic status, ethnicity, and sex of the child. Similarly, maternal cortisol and prenatal anxiety
levels mid-pregnancy were associated with elevated cortisol levels in response to the first day of a
new school year (Gutteling, de Weerth, and Buitelaar, 2005). Such resetting of the HPA axis has
consequences for postnatal health; we next examine a specific example: attention deficit hyperac-
tivity disorder (ADHD).
A multitude of pre- and postnatal risk factors (e.g., low birth weight, smoking, alcohol use, stress
during pregnancy, and insensitive or abusive parenting) increase the risk of ADHD and disruptive
behavior disorders (DBDs; Latimer et al., 2012). Some studies have managed to examine the inde-
pendent impact of maternal stress on ADHD. One of the most striking examples involves a twofold
increase in ADHD symptoms in children whose mothers were pregnant during, but not exposed
to, the Chernobyl disaster in 1986 (Huizink et al., 2007). Similarly, in the Avon Longitudinal Study
of Parents and Children (ALSPAC), prenatal anxiety at 32 weeks of gestation was associated with
an increased risk of inattention/hyperactivity at 4 years of age (O’Conner et al., 2002). And in a
more recent prospective study of 1,247 Finnish children, maternal depression at 10 and 28 weeks of
gestation are associated with externalizing problems on the Child Behavior Check List, while mater-
nal postnatal factors (illness, tiredness, and anxiety) were associated with internalizing problems at
12 years of age (Pihlakoski et al., 2013). As with several other programming effects, there is intriguing
evidence of sexual dimorphism in the impact of the maternal environment. For example, Rodriguez
and Bohlin (2005) found that maternal stress at 10 weeks of gestation was associated with ADHD
symptoms at 7 years, but only in boys. Likewise, examination of a Finnish birth cohort revealed that
several measures of placental size were associated with ADHD symptoms in boys, but not girls, at 8
and 16 years (Khalife et al., 2012). These sex differences are intriguing and are consistent with animal
and other human studies and may be underpinned by epigenetic mechanisms.
A consistent finding in the DOHaD literature is that boys are more susceptible to environmental
insults during pregnancy than girls (Barker, 2003). Male-biased effects of prenatal exposures have
been found for schizophrenia, autism spectrum disorders, and anxiety (Bale, 2011). Perhaps, as the
faster-growing sex, males are more vulnerable. A possible mechanism underlying the differing impact
of maternal stress on the male and female postnatal phenotype is altered sensitivity to GCs. Across
several species, females appear to be more resistant and males appear to be more sensitive to GC
exposure (Phillips and Matthews, 2011; Seckl and Holmes, 2007). Evidence of sex differences in the
activation of the HPA axis has been found in animal studies (Liu, Li, and Matthews, 2001). Detailed
analyses in mice show that the male vulnerability to prenatal stress, resulting in increased emotionality
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postnatally, may be attributable to increased placental responsivity with an up-regulation of placen-

tal gene expression found in the placentae of male but not female mice (Mueller and Bale, 2008).
Moreover, the prenatal stress-influenced placental epigenetic processes with increased expression of
DNA methyl transferases and methyl-binding proteins in the male but not female placentae. Thus,
the sex-specific effects of maternal stress on postnatal phenotypes may be transduced via epigenetic
processes in the placenta.
What, if anything, do evolutionary models say regarding sex-specific effects? An evolutionary
model, sex allocation theory (Charnov, 1982), proposed that, when maternal condition is poor,
investment of resources should be allocated to the offspring sex that requires the least investment.
As sons grow faster and require more resources to produce, under conditions of prenatal stress the
maternal physiology may preferentially allocate resources away from male fetuses and thus increase
their sensitivity to environmental insults. Crucially, for this argument, stress hormones appear to
provide the link between the maternal condition and the allocation of resources. In a study of
European starlings, Love, Chin, Wynne-Edwards, and Williams (2005) found that elevated maternal
corticosterone levels were associated with biasing allocation of resources towards females, with males
experiencing higher mortality and being lighter at hatching. Therefore, maternal condition during
pregnancy may have sex-specific effects on the postnatal phenotype, including mental and physical
health.
Prenatal Nutrition, Fetal Development, and the Postnatal Phenotype

No discussion of prenatal development is complete without examining the impact of maternal nutri-
tion on the fetus and its postnatal phenotype. For the vast majority of human history, the primary
concern has been undernutrition. Now, with the emergence of the obesity epidemic, overnutrition
is of increasing concern (Wells, 2012). Unfortunately for the developing fetus, both maternal under-
and overnutrition provide suboptimal environments that can affect early development and lifelong
health (Ojha, Robinson, Symonds, and Budge, 2013). Even in “well nourished” populations, an
imbalanced diet can affect fetal development, and, if not in the form of reduced growth during preg-
nancy, it can still influence postnatal health (Barker, 2003; Kind, Moore, and Davies, 2006). Below, we
briefly examine two important components of the maternal diet, iron and folate, before looking at
the consequences of maternal nutrition for postnatal phenotypes.
Iron
Iron deficiency is the most common nutrient deficiency among pregnant women (Guilbert, 2003).
The prevalence of anemia among pregnant women worldwide is estimated to be 41.8%, with iron
deficiency the leading cause (McLean, Cogswell, Egli, Wojdyla, and da Benoist, 2009). Iron demands
during pregnancy increase rapidly, especially late in gestation. Peak gastrointestinal iron absorption
in pregnant women occurs in the third trimester (Barrett, Whittaker, Williams, and Lind, 1994),
concurrent with the increased iron transfer to the fetus. In the absence of adequate maternal iron
status, the placenta increases the number of transferrin receptors in order to enhance placental iron
absorption in an attempt to maintain adequate iron transfer to the fetus (Cetin, Berti, Mandò, and
Parisi, 2011).
Maternal iron deficiency early in pregnancy is associated with an increased risk of low birth
weight and preterm delivery (Chang, O’Brien, Nathanson, Mancini, and Witter, 2003; Scanlon, Yip,
Schieve, and Cogswell, 2000). The relation, however, appears to be U-shaped, with high maternal
hemoglobin concentrations in early to mid-gestation also associated with increased risk of adverse
birth outcomes, including preterm birth and low birth weight (Chang et al., 2003; Scanlon et al.,
2000).
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Iron is required for a number of physiological processes crucial to development, including oxy-
gen transport, energy production, and, notably, brain development. Dietary iron deficiency during
prenatal development in animal models leads to an alteration in the composition and quantity of
neuronal myelin sheaths (Ortiz et al., 2004). Similarly, iron is required for the synthesis of the neuro-
transmitters serotonin (Martinez, Knappskog, and Haavik, 2001) and dopamine and norepinephrine
(Nagatsu, 1995).
Iron deficiency during infancy is associated with long-lasting effects on neurocognitive outcomes,
including decreased executive function at the ages of 5, 10, and 19 (Lozoff et al., 2006). Moreover,
interventions improve neurocognitive outcomes. For example, a study in rural Nepal with a high
prevalence of anemia observed improved general intellectual functioning, executive function, and
motor skills in 7- to 9-year-old children following maternal supplementation with iron and folic acid
compared with a control group (Christian et al., 2010). While there is a paucity of data regarding the
mechanism through which maternal iron deficiency impacts cognitive function in children, impaired
myelination is a potential cause.
Folate
Folate (folic acid) is an important factor in fetal development that is involved in a number of biologi-
cal processes, including acting as a cofactor for many essential cellular reactions, such as amino acid
metabolism, DNA synthesis, DNA methylation, and red blood cell production. Fetal development is
a time of rapid and sustained cell division and thus the demand for folate increases during this period
(McPartlin, Halligan, Scott, Darling, and Weir, 1993). In the absence of adequate dietary folate intake
and/or folic acid supplementation, maternal serum and erythrocyte folate concentrations decrease
from mid-pregnancy onwards and continue to decline postpartum (Qvist, Abdulla, Jagerstad, and
Svensson, 1986).
The implications of maternal folate nutrition on fetal growth have been investigated in a num-
ber of studies worldwide, with the clearest finding that peri-conceptual folic acid supplement use
reduces the risk of neural tube defects (e.g., spina bifida) by almost three quarters (Lumley, Wat-
son, Watson, and Bower, 2001). Based on animal models, which show the importance of adequate
methyl donors, such as folate, during neural tube closure, it has been hypothesized that inadequate
folate availability may increase the risk of neural tube defects in humans through reduced DNA
methylation (Dunlevy et al., 2006). In a hyperhomocysteinemia rat model dietary folate intake was
found to correlate with the methylation of DNA in the placenta (Kim, Hong, Lee, Lee, and Chang,
2009). Folic acid over supplementation in pregnant rats was found to modify the methylation and
expression of the placental gene encoding 11β-HSD-2 in a sex-dependent manner, suggesting that
maternal diet can induce sex-specific differences in physiology that may have long-term health con-
sequences (Penailillo, Guajardo, Llanos, Hirsch, and Ronco, 2015).
Nutrition and Fetal Programming

Nutrition is a key determinant of fetal growth and size at term. The initial findings by Barker, linking
small size at birth and chronic adult diseases such as hypertension, diabetes, and cardiovascular disease,
were thought to result from an imbalance between the maternal supply and fetal demand for nutri-
ents during pregnancy (Phillips, 2004). This hypothesis is now supported by evidence from animal
studies (Eriksson, Forsen, Tuomilehto, Osmond, and Barker, 2003) and longitudinal epidemiological
investigations of diet during pregnancy in humans (Ojha et al., 2013; Phillips, 2004).
Prenatal brain development is dependent on adequate nutrition (Walker, 2005). It is now appar-
ent that the impact of maternal nutrition also extends to mental health outcomes. Prenatal exposure
to the Dutch hunger winter (1944–1945) in the second or third trimester was associated with an
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increased risk of affective disorders in both males and females (Brown, van Os, Driessens, Hoek, and
Susser, 2000). In a study of 23,020 Norwegian women, Jacka et al. (2013) found that higher intake
of unhealthy foods during pregnancy was associated with increased levels of externalizing problems
at 1.5, 3, and 5 years of age. This relation held after adjustment for a range of confounds, including
socioeconomic status, depression during pregnancy, and children’s postnatal diet. Similarly, in a non-
human primate model, maternal nutrient restriction during pregnancy was associated with juvenile
attentional and behavioral problems (Keenan et al., 2013). Together, these studies highlight a poten-
tial role of nutrition during pregnancy for mental health disorders.
The evidence that small size at birth is associated with poorer adult mental and physical health
outcomes is substantial. The proximate mechanisms by which these effects could be transmitted
throughout a lifespan, however, are not completely understood. A likely candidate is that maternal
undernutrition, like maternal prenatal stress (see above), re-programs the HPA axis and has conse-
quences for postnatal health (Phillips and Matthews, 2011). Undernutrition as a stressor makes sense
because glucocorticoids are involved in both the stress response and the liberation of glucose for the
cells of the body. In a meta-analysis of 11 studies (N = 2,311), lower birth weight was significantly
associated with higher levels of cortisol during childhood (van Montfoort, Finken, le Cessie, Dek-
ker, and Wit, 2005). Increased stress reactivity to the Stroop Test has also been detected in a study
of 721 people who were exposed to the Dutch hunger winter prenatally (Painter et al., 2006). In
animal models, maternal undernutrition during pregnancy has been associated with the increased
expression of glucocorticoid receptor 11β-HSD-1 and the decreased expression of 11β-HSD-2
suggesting programming of gene expression related to the HPA axis (Whorwood, Firth, Budge, and
Symonds, 2001). Other emerging areas of interest involve pathways such as the immune process and
inflammation during pregnancy which are also crucial to reproduction, interact with the HPA axis,
and contribute to human life history trade-offs (Clancy, 2013; Entringer et al., 2012; McDade, 2005).
Researchers rarely consider that stress hormones may also have a behavioral effect on maternal
nutrition. In humans, stressful events or administration of CRH increase the eating of comfort
food, but there has been no study examining the impact of maternal stress on nutritional intake
during pregnancy and child outcomes (Entringer et al., 2012). Thus, evidence supports the hypoth-
esis that maternal nutrition influences childhood mental and physical health by re-programming
the HPA axis.
Epigenetics as a Potential Organizing Mechanism

Epidemiological data and evolutionary theory suggest that insults during one generation can have
consequences for the growth and development of second and third generation descendants (Bateson
et al., 2004; Coall and Chisholm, 2003; Kuzawa, 2005; Ounsted et al., 1986). During development
in mammals, there is extensive DNA methylation. The majority of methylation occurs during the
development of the germ cells (future sperm and ovum) and in the early embryonic cells forming
the blastocyst (Reik, Dean, and Walter, 2001; Schaefer, Ooi, Bestor, and Bourc’his, 2007). This meth-
ylation follows periods of demethylation where the epigenetic markers are thought to be “wiped
clean” and reset.
The mechanisms responsible for fetal growth and intrauterine programming are regulated by
placental nutrient transport. Placental inefficiency is also associated with epigenetic modifications.
Jansson and Powell (2013) reported that restriction of protein in the maternal diet of rats induced
epigenetic changes in specific genes, the glucocorticoid receptors in the liver of the offspring. Fur-
thermore, genes expressed in the placenta may undergo epigenetic modification in response to dis-
turbances in the maternal compartment due to direct exposure to maternal blood. The methylation
of certain genes, such as the trophoblast genes, also results in altered placental structure and function.
The effects of methylation due to maternal stressors have direct effects on placental morphology
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and fetal programming. In addition to modifications in methylation, epigenetic mechanisms include

noncoding RNA molecules, such as microRNAs (miRNA). These short RNA fragments can pro-
vide post-translational gene expression, through the inhibition of protein translation.
Prenatal Toxins, Teratogens, and Offspring Development

Research on prenatal development in human evolutionary ecology, life-course epidemiology, and the
DOHaD paradigm predominately focuses on maternal stress and nutrition. Stressors often ignored
in these analyses, possibly because of the disruptive rather than adaptive nature of exposure, are tox-
ins and pollutants. Many of these substances persist in the environment and accumulate over time.
Human exposure arises through multiple pathways including direct contact with soils, inhalation
of particulate matter and household dust, dermal contact, and the consumption of contaminated
food or drinking water. In this section we highlight some pollutants and their influence on prenatal
development.
Tobacco Smoke
Maternal smoking during gestation is known to result in reduced fetal growth, with a decrease of
approximately 200g in infants born to smoking mothers relative to those born to nonsmokers (Rog-
ers, 2009). Tobacco smoke contains more than 4,000 chemicals, one of which is cadmium, and hence
cadmium may play a role in the reduction in birth weight.
Prenatal tobacco exposure was significantly associated with ADHD in a study of 4,704 U.S. chil-
dren (Braun, Kahn, Froehlich, Auinger, and Lanphear, 2006). Analysis of a nationally representative
sample of 8- to 15-year-olds revealed a greater than eightfold increased risk of ADHD for children
who had both prenatal environmental tobacco smoke exposure and postnatal lead exposure (Froe-
hlich et al., 2009), demonstrating the importance of environmental exposures for this prevalent
health condition.
A meta-analysis of 14 studies conducted worldwide with a total of over 84,000 children aged
2 years and above revealed that maternal smoking during pregnancy was associated with an elevated
risk of the offspring being overweight at the ages of 3–33 years (Oken, Levitan, and Gillman, 2008).
Notably, the odd ratios were almost unchanged when adjusted for a range of factors including paren-
tal sociodemographic factors, signifying that social and behavior differences between smokers and
nonsmokers were unlikely to account for the increased risk (Oken et al., 2008). Maternal smoking
during pregnancy has also been associated with an increased risk of early onset of type 2 diabetes
in young adults (Montgomery and Ekbom, 2002) and early onset of puberty in males (Fried, James,
and Watkinson, 2001).
Although discussions of prenatal exposures tend to focus on maternal exposures, onset of paternal
smoking prior to puberty has been associated with an increase in body mass index (BMI) in male
offspring at the age of 9 years, with the effect most pronounced if fathers started smoking when
they were aged 10 years or under (Pembrey et al., 2006). The relation with early paternal smoking
provides compelling evidence of a male-line transgenerational effect.
Bisphenol A
Bisphenol A (BPA) is a ubiquitous chemical used in the manufacture of polycarbonate plastics and
epoxy resins. It is found in a number of types of food packaging, including in the lining of tins.
Biomonitoring in developed countries has indicated that exposure is widespread. BPA has been
shown to transfer across the human placenta, mainly in the active unconjugated form (Balakrishnan,
Henare, Thorstensen, Ponnampalam, and Mitchell, 2010; Mørck et al., 2010). BPA is known to
187
have detrimental effects on placental cells, with low, environmentally relevant, doses of the chemi-
cal inducing apoptosis in placental trophoblasts via increased expression of tumor-necrosis factor α
(TNF-α) (Benachour and Aris, 2009) as well as altering the microRNA expression in cells derived
from the first trimester placenta. Therefore, BPA may alter the DNA repair capabilities of these cells
(Avissar-Whiting et al., 2010).
Consistent with the effects of BPA on placental cells, some studies have shown an association
between prenatal BPA exposure and reduced head circumference and weight (Philippat et al., 2012;
Snijder et al., 2013; Wolff et al., 2008). Because BPA concentrations in maternal urine are known to
vary during pregnancy (Braun, Kalkbrenner, Calafat, Bernert, et al., 2011), exposure misclassification
is possible when samples are collected at a single time point. Equally, the specific window of vulner-
ability to BPA exposure in terms of fetal growth outcomes is unknown.
The potential relation between BPA exposure and neurobehavioral outcomes has been examined
in a number of studies, with sex-dependent effects reported. In boys, prenatal BPA exposure has been
associated with increased emotionally reactive and aggressive behavior at ages 3–5 years (Perera et al.,
2012) and internalizing behaviors, including anxiety and depression, at 7 years (Harley et al., 2011;
Roen et al., 2015), as well as a range of parent-reported problem behaviors aged 6–10 years (Evans
et al., 2014). Gestational exposure to BPA has also been associated with increased risk of ADHD-
related behaviors at 4 years of age, again with stronger associations in boys (Casas et al., 2015). Con-
versely, one study identified increased externalizing behaviors at age 2 and higher scores for measures
of anxiety, hyperactivity, emotional control, and behavioral inhibition at age 3, with stronger effects
in girls at both ages (Braun et al., 2009; Braun, Kalkbrenner, Calafat, Yolton, et al., 2011). The rea-
sons for the discrepancies in findings are unclear and may reflect methodological differences in the
studies. Nonetheless, the ability for prenatal BPA exposure to modify child behavior in a sex-specific
manner is apparent.
In vitro experiments have shown that BPA can inhibit adiponectin release from human adipose
tissue (Hugo et al., 2008), suggesting adverse affects on metabolic homeostasis. Similarly, prenatal
exposure to BPA has also been associated with BMI and waist circumference in children at 4 years
(Valvi et al., 2013) and fat mass index, percentage body fat, and waist circumference at age 7 (Hoe-
pner et al., 2016).
Phthalates
Phthalates are used widely in a range of consumer products. For example, the high molecular weight
di(2-ethylhexyl) phthalate (DEHP) is used as a plasticizer in the manufacture of polyvinyl chloride,
whereas low molecular weight phthalates, such as diethyl phthalate (DEP) and dibutyl phthalate
(DBP), are used in cosmetics and other personal care products. Animal models have demonstrated
that prenatal exposure to DEHP, DBP, or benzyl butyl phthalate (BzBP) has a marked effect on
fetal Leydig cells, reducing the production of fetal testosterone and insulin-like growth factor 3
(Insl-3; Foster, 2006). These effects are responsible for a syndrome of male reproductive abnormali-
ties, including shortened anogenital distance, hypospadias, cryptorchidism, and malformations of the
epididymis, vas deferens, seminal vesicles, and prostate (Foster, 2006). In rats, the programming win-
dow in which androgen action is required for the normal development of the reproductive tract has
been identified, corresponding to 8–14 weeks’ gestation in humans (Welsh et al., 2008).
In human studies, DEHP metabolites measured in maternal urine collected during gestation
(mean week 28.6) were associated with reduced anogenital distance in males (median age at assess-
ment 12.8 months), decreased penile width and impaired testicular descent (Bustamante-Montes
et al., 2013; Suzuki et al., 2010; Swan, 2008). A larger study was established to examine the association
between first trimester phthalate exposures and anogenital distance in children, with the timing of
sample collection matching the window of vulnerability identified in animal models (Welsh et al.,
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2008). This study confirmed that, despite low levels of exposure, metabolites of the anti-androgenic
DEHP measured in first trimester urine samples were significantly negatively associated with ano-
genital distance and penile width in newborn boys (Swan et al., 2015). Anogenital distance has been
identified as a strong predictor of multiple semen parameters in adult men, which suggests that this
measure is a good indicator of male reproductive function (Mendiola, Stahlhut, Jorgensen, Liu, and
Swan, 2011). Therefore, there are possible lifelong impacts of prenatal phthalate exposure on male
reproductive development at the population level, which is of concern given the ubiquitous nature
of exposure.
Maternal occupational exposure to phthalates (estimated by questionnaire only) was associated
with decreased placental weight and also decreased fetal growth in a Dutch cohort (Snijder et al.,
2012). Consistent with this indication of impaired placental development, maternal urinary phtha-
late concentrations were associated with placental gene expression in trophoblast differentiation and
steroidogenesis pathways (Adibi et al., 2010). Increased maternal phthalate exposure in the early third
trimester was correlated with decreased placental gene expression, with results more pronounced for
genes of the trophoblast differentiation pathway (Adibi et al., 2010).
The effect of prenatal exposure to anti-androgenic phthalates on sexual differentiation of the
brain is also being explored. Concentrations of two metabolites of DBP and two metabolites of
DEHP in maternal urine (mean 28.6 weeks gestation) were associated with reduced masculine play
in boys aged 3–6 years (Swan et al., 2010). Prenatal phthalate exposure also influences neurobehavior
in infants and children, including increased ADHD (Engel et al., 2009; Kobrosly et al., 2014; Lien
et al., 2015; Whyatt et al., 2012) and autistic behaviors (Miodovnik et al., 2011), impaired mental
and psychomotor development (Kim et al., 2011; Whyatt et al., 2012), and increased emotional dif-
ficulties (Whyatt et al., 2012). Prenatal phthalate exposure measured late in gestation has also been
associated with decrements in children’s IQ at the age of 7 (Factor-Litvak et al., 2014). Despite some
variations in the findings between studies, which may reflect differences in the gestational timing
of exposure assessment and/or differences in the age of children or method of neurodevelopmental
assessment, overall the literature supports the notion that prenatal phthalate exposure is associated
with behavioral and cognitive impacts in children.
Perfluoroalkyl Substances
Perfluoroalkyl substances (PFAS) are used extensively in a range of commercial and industrial appli-
cations. Perfluorooctanesulfonate (PFOS) and perfluorooctanoate (PFOA) are PFAS that are pro-
duced either directly or from the metabolism of other PFAS, with half-lives between 4 and 5 years
(Olsen et al., 2007). These chemicals are used as industrial surfactants and may be present in a
number of consumer products including nonstick pans, soft furnishings, clothes, and food packaging.
PFAS have been extensively used for several decades and persist for long periods in the environment,
but human exposure has only been assessed relatively recently, in part due to our previous inability
to detect these chemicals with sufficient sensitivity in biological media. Although the use of PFAS
was phased out in most countries in 2000, human exposure to perfluorinated compounds remains
widespread with biomonitoring studies indicating that four perfluoroalkyl acids (perfluorooctanoic
acid [PFOA], perfluorooctane sulfonate [PFOS], perfluorononanoic acid [PFNA], and perfluorohex-
ane sulfonate [PFHxS]), are detectable in most blood samples. These compounds are known to cross
the placental barrier and are prevalent in cord blood samples.
The evidence to date regarding prenatal PFAS exposure and fetal growth is largely consistent,
with decreased fetal growth reported to be associated with PFOS, PFOA, or both, in most studies
(Apelberg et al., 2007; Chen et al., 2012; Fei, McLaughlin, Tarone, and Olsen, 2007; Washino et al.,
2009; Whitworth et al., 2012). Prenatal exposure to PFHxS has also been associated with reduced
fetal growth (Callan et al., 2016; Maisonet et al., 2012).
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In British girls from the ALSPAC study, weight at 20 months increased with increased prenatal
exposure to PFOS, with those in the highest tertile for PFOS on average 580 g heavier than those
with prenatal exposure in the lowest tertile (Maisonet et al., 2012). Prenatal PFOA concentrations
have also been associated with increased BMI, waist circumference and body fat in children at 8 years
of age, as well as increased gains in child BMI between the ages of 2 and 8 years (Braun et al., 2016).
Most studies find that decreased birth weight is associated with prenatal PFOS exposure, suggesting
a rapid early growth catch-up that may have implications for long-term child health.
Furthermore, there is evidence that the effects of prenatal PFAS exposure on anthropometry
persist into adulthood, with positive associations between prenatal exposure to PFOA and the preva-
lence of overweight and high waist circumference (>88 cm) in females, but not males, at 20 years
of age in a Danish cohort (Halldorsson et al., 2012). Maternal PFOA concentrations in the third
trimester were also associated with biomarkers of adiposity in female adult offspring, with positive
associations with insulin, leptin, and leptin-adiponectin ratio, and negative associations with adi-
ponectin (Halldorsson et al., 2012).
To date the potential associations between prenatal PFAS exposure and the male human repro-
ductive system have received limited attention in epidemiological studies. In a Danish study,
increased maternal PFOA serum concentrations measured at week 30 in gestation were associated
with lower sperm concentration and reduced total sperm count in male offspring at 19–21 years of
age, with higher concentrations of luteinizing hormone and follicle-stimulating hormone detected
in the blood samples of those with higher in utero PFOA exposure (Vested et al., 2013). No asso-
ciations were observed between prenatal PFOS exposure and the outcomes measured. Given the
widespread nature of exposure to PFOA, these decreases in semen quality in adult men following
prenatal exposure could pose a significant population health issue and warrant further investi-
gation. Of equal concern is the suggestion that prenatal exposure to PFOS is associated with
reduced antibody-mediated immune responses to childhood immunizations at the age of 5 years
and increased odds of diphtheria antibody concentrations below the protective level (Grandjean
et al., 2012). Consistent with this potential immunotoxic effect, prenatal exposure to PFOS and
PFHxS has been associated with increased prevalence of infectious diseases in children in the first
4 years of life (Goudarzi et al., 2017).
Conclusion and Perspectives for Future Studies

Investigations into the potential health effects of prenatal exposures to environmental toxicants are
essential. The main challenge in this field is to deal with a number of methodological issues to
improve the accuracy of conclusions that can be made. Epidemiological, in vivo, and in vitro studies
frequently examine the effects of exposure to a single toxicant, but environmental exposure involves
chronic simultaneous low-level exposure to multiple toxins which may have synergistic or cumula-
tive effects. The ability to accurately assess the health implications of exposure to multiple ubiq-
uitous environmental pollutants and identify the pollutants most strongly associated with adverse
outcomes remains a challenge. Longitudinal birth cohort studies are expensive and need to follow
up participants for many years to accurately ascertain the health effects of prenatal exposure, some of
which may not be manifest until the offspring reach adulthood. In addition, for most adverse health
effects associated with environmental exposures a discrete window of vulnerability in the prenatal
period is yet to be identified, which means that ideally such studies must collect biological samples
at multiple time points throughout pregnancy. Furthermore, as increasing evidence emerges of sex-
specific health implications of exposure to environmental substances, sample sizes in studies will need
to increase to accommodate the statistical analysis of sexually dimorphic associations. Finally, many
health outcomes assessed by longitudinal birth cohorts involve a complex interplay among multiple
elements, including genetic susceptibility, maternal diet, and prenatal (and postnatal) environmental
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exposures. For example, an association between prenatal exposure to pesticides and autism spectrum
disorders was attenuated by high folic acid supplementation early in pregnancy (Schmidt et al.,
2017). If indeed such a protective effect of folic acid represents a causal relation, the mechanisms by
which it could occur are yet to be elucidated, although a role for DNA methylation pathways has
been speculated. Nonetheless, this study highlights the importance of including full consideration of
all aspects of the prenatal environment when assessing maternal influences on offspring phenotype.
Prenatal Parenting: Future Applied, Empirical,

and Theoretical Perspectives
Prenatal parenting and the prenatal environment more broadly is a burgeoning field of enquiry with
increasing significance for health. Our understanding of and thus potential to improve aspects of the
maternal environment is progressing rapidly. In this section we explore specific issues that we believe
are pertinent to future applied, empirical, and theoretical work in prenatal parenting.
Prenatal Bonding
Early parent-child relationships are known to have a strong psychological influence that lasts a life-
time (Bowlby, 1969). Consequently, early relationships with parents are crucial throughout devel-
opment and into adulthood. Moreover, these early experiences have a profound influence on an
individual’s health and emotional well-being (DeKlyen and Greenberg, 2008; Dozier and Rutter,
2008). Insensitive, unresponsive parenting is commonly associated with more insecure attachment,
which has consequences for children’s relationships, development, parenting, and the intergenera-
tional transmission of insecure attachment (Belsky et al., 1984; Berthelot et al., 2015; Chisholm,
Quinlivan, Peterson, and Coall, 2005). Prenatal mother-fetal bonding appears to be a reliable pre-
dictor of early postnatal bonding. In an Australian study, Rossen and colleagues (2016) found that
prenatal bonding, as measured via the Maternal Antenatal Attachment Scale in the first, second and
third trimester, predicted bonding 8 weeks postnatally. These associations remained after adjustment
for a wide range of factors including age, socioeconomic status, birth weight, crying, and pregnancy
complications. Moreover, a systematic review found that lower levels of prenatal bonding were asso-
ciated with poorer infant developmental outcomes, such as a difficult temperament, an increased risk
of colic, and delayed developmental outcomes (Branjerdporn, Meredith, Strong, and Garcia, 2017).
Therefore, prenatal parenting in the form of mother-fetal bonding appears to have consequences for
the postnatal phenotype.
Social support buffers individuals against stressful life events and may play a role in adjustment to
stressors experienced during pregnancy and neonatal outcomes. Indeed, across cultures, having two
or more friends or family members available in your social network and higher levels of social sup-
port were both independently associated with lower levels of depression postnatally (Surkan et al.,
2006). Consistent with this social support perspective and the proposition that humans are coopera-
tive breeders (Hrdy, 1999), a Japanese study found that having fewer support people available during
pregnancy was associated with poorer mother-fetal bonding before 25 weeks of pregnancy and 1
month postnatally (Ohara et al., 2017). Thus, consistent with life history theory interpretations of
attachment (Chisholm, 1996), mounting evidence suggests the impact of a mother’s social environ-
ment is transmitted to her offspring through maternal-fetal bonding.
Higher quality maternal-fetal bonding is associated with improved neonatal and infant outcomes
(Branjerdporn et al., 2017) suggesting bonding is a potential intervention target. Moreover, interven-
tions exist that have been shown to promote prenatal bonding. For example, in a study of a prenatal
education course consisting of five 1-hour sessions that focused on fetal physiology, development,
and perceptions through singing, dance, and massage were associated with higher levels of prenatal
191
attachment in the third trimester (Bellieni et al., 2007). As with other species (Colombelli-Negrel
and Kleindorfer, 2017), during the neonatal period the newborn seems to prefer its mother’s voice
(DeCasper and Fifer, 1980) which may provide a mechanism for intervention. Therefore, antenatal
care can play an important role by integrating activities into maternal care that promote maternal-
fetal bonding during pregnancy (Rossen et al., 2016), such as the use of singing, ultrasound, and
memories of parental relationships (Branjerdporn et al., 2017).
Antenatal Care
Maternal health and behavior during pregnancy clearly have consequences for offspring phenotype.
Some aspects of maternal health can be modified and will benefit from antenatal care, whereas others
cannot (Kramer, 1987). Maternal health is unequally distributed within and between populations and,
like parenting, is a mechanism for the intergenerational transfer of risk factors for poor health that
may perpetuate health inequalities across generations (Fonagy and Higgitt, 2000; Kelly et al., 2017).
In developing countries antenatal care is crucial for improving maternal and child health out-
comes (Lincetto, Mothebesoane-Anoh, Gomez, and Munjanja, 2006). Moreover, the introduction
of or additional support for antenatal care has positive effects (Choe, Min, and Cho, 2017). In cases
where antenatal care is well established, the health impact which the timing of antenatal care has
(when it begins) and the number of antenatal visits are difficult to assess. Moreover, study design is
difficult because women with pre-existing health conditions may be more likely to initiate antenatal
care earlier and attend more often. However, the quality of antenatal care, particularly for high-risk
pregnancies, appears to improve outcomes (Kramer, 1987). In an audit of maternal mortality (2009–
2012) in the Republic of Ireland and the United Kingdom, the Confidential Enquiry into Maternal
Deaths and Morbidity, of the 357 women who died during pregnancy or within 6 weeks of the end
of their pregnancy, only 29% had the recommended level of antenatal care and 10% received no
antenatal care (Shakespeare and Knight, 2015), suggesting that antenatal care is especially important
for high-risk pregnancies.
Maternal care is crucial for removing or reducing any modifiable risk factors which ultimately pro-
motes healthy pregnancy, placental development, and outcomes (Kramer, 1987). In turn, minimizing
risk factors is associated with the general health of the broader population (Kloosterman, 1970). Ensur-
ing sustainable, equal, and relatively easy access to antenatal care is the issue of concern. Thus, it is likely
antenatal care that can support existing health conditions (e.g., diabetes, asthma), help make informed
decisions about care (e.g., morning sickness, mental health), and modify behavior (e.g., smoking, diet)
will produce favorable outcomes for maternal and fetal phenotypes (Kramer, 1987).
Resource Flow to the Fetus—Not Maternal Nutrition

Our evolutionary approach to reduced fetal growth suggests that the current emphasis of the DOHaD
model on maternal under- or malnutrition as the main cause of reduced fetal growth and subsequent
increased risk of adult disease may be misplaced. Adverse prenatal environments definitely contribute
to reduced placental function and restricted fetal growth, but the impact of maternal nutrition on
fetal and placental growth is complicated (Robinson et al., 1994, 2001). The influence of maternal
dietary supplementation on birth weight is small (Harding, 2001), and the outcomes depend heav-
ily on the existing nutritional state of the population (Kind et al., 2006). Investigations of mater-
nal nutrient intake and subsequent fetal and placental weights at birth have yielded mixed results
(Godfrey, Robinson, Barker, Osmond, and Cox, 1996; Kuzawa, 2005; Mathews, Yudkin, and Neil,
1999; Moore, Davies, Willson, Woesley, and Robinson, 2004). Our model suggests that the effect of
maternal nutrition on placental ratio is a consequence not only of maternal nutrient intake, but also
that the flow of resources from the uterus to the fetus is “negotiated” by the mother and fetus. The
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model we are working with suggests that the long and vulnerable fetal supply line, not measurable
maternal nutrition per se, determines the ultimate flow of resources to the fetus (Godfrey et al., 1999;
Harding, 2001).
As resources from the maternal diet travel along the fetal supply line, what ultimately becomes
fetal nutrition is mediated by the mother’s metabolism and endocrinology, uterine blood flow, pla-
cental transport and metabolism, umbilical blood flow, and the fetus’ metabolism and endocrinology
(Harding, 2001). Therefore, the allocation of resources between competing demands during preg-
nancy mean that maternal nutrition and fetal nutrition are not the same thing (Gillman, 2002; Hard-
ing, 2001). This allocation of resources may also partly explain the finding that maternal insults that
do not necessarily affect fetal growth can still have consequences for a child’s subsequent postnatal
development and health. Our evolutionary synthesis highlights an interaction between the maternal
reproductive strategy and diet as a possible explanation for some of the inconsistencies in studies of
maternal nutrient supplementation and fetal growth and the fetal antecedents of adult disease (Coall
and Chisholm, 2010; Kramer, 2000; Kramer and Joseph, 1996).
Morning Sickness: Nutrition, Stress, Hormones, and Long-Term Outcomes

A common characteristic of the maternal phenotype that includes nutritional and stress compo-
nents is morning sickness. Morning sickness (nausea and vomiting in pregnancy [NVP]) is a suite of
symptoms affecting up to 90% of pregnant women (Einarson, Piwko, and Koren, 2013). Symptoms
may include nausea, vomiting, retching, and food avoidance or aversion (Patil et al., 2012). Typically,
symptoms begin around 4 to 6 weeks of pregnancy and continue until the 16th week of pregnancy.
Given its frequency, it is reasonable to say that morning sickness is a “normal” part of pregnancy that
exists as a continuum, with expectant mothers experiencing no symptoms at one end and those with
exacerbated symptomology at the other. Hyperemesis gravidarum (HG) occurs in up to 3% of all
pregnancies (reported numbers only) and is characterized by persistent nausea, vomiting, retching,
dehydration, maternal weight loss of up to 3%–5%, and electrolyte imbalances that often result in
hospitalization (Goodwin, 2002; Koren, 2014). Many women find NVP stressful and debilitating
which can affect physical and mental health for the entire pregnancy (Bustos, Venkataramanan, and
Caritis, 2017; Lee and Saha, 2011).
Morning sickness in the first trimester of pregnancy is believed to be associated with a protective
role and positive birth outcomes (Profet, 1992). Food aversion in early pregnancy (in animal and
human models) stimulates the growth of the placenta and reduces the risk of miscarriage, preterm
birth, and low birth weight babies (Huxley, 2000). Compensatory placental growth mechanisms are
believed to be responsible for the redirection of maternal resources in response to nutrient restric-
tion. Evidence suggests morning sickness may be a direct result of adaptive placental functioning
( Jauniaux, Poston, and Burton, 2006). Women who experience no symptoms of morning sickness
are reported to have larger placentas and low birth weight outcomes. Nutritive insult in the second
and/or third trimester (seen with severe HG) can result in fetal growth restriction and consequent
shorter gestation and lower birth weight offspring (Vandraas et al., 2013; Veenendaal et al., 2011).
The impact of morning sickness extends well beyond the antenatal period (Veenendaal et al.,
2011). Indeed, HG is associated with a range of mental and physical issues for the mother both before
and after pregnancy (Tian, MacGibbon, Martin, Mullin, and Fejzo, 2017). Moreover, HG impacts
the entire family with expectant fathers who, as principal carers, reported elevated feelings of anxi-
ety (Sartori et al., in press). Large population-based cohort studies have investigated the maternal,
birth, and neonatal outcomes after pregnancies complicated with HG. In a cohort of more than
eight million pregnancies from England (1997–2012; Fiaschi et al., 2018), HG was associated with an
increased risk of diseases of pregnancy (e.g., anemia, eclampsia), poor birth outcomes (e.g., preterm
birth, low birth weight), and increased neonatal morbidity (e.g., neonatal intensive care). Results
193
from a Swedish birth cohort find these risks are higher when first admission occurs in the second
trimester, suggesting there may be a relation between abnormal placentation and HG (Bolin, Akerud,
Cnattingius, Stephansson, and Wikström, 2013).
HG also is associated with subsequent maternal morbidity and mortality, including placen-
tal dysfunction, inflammation, and autoimmune diseases (Bolin et al., 2013; Jørgensen, Nielsen,
Pedersen, Jacobsen, and Frisch, 2012). A population-based cohort from Norway, however, found
no increase in subsequent maternal mortality (all-cause, cardiovascular disease, external cause,
or mental or behavioral disorder cause) in women who experienced HG (Fossum et al., 2017).
A decreased risk of cancer, particularly tobacco-related cancers, was noted. However, the lower
rates of cancers may be confounded by the fact that nicotine receptors suppress nausea via decreas-
ing levels of estradiol and human chorionic gonadotropin (hCG) resulting in a low rate of HG in
smokers (Bernstein et al., 1989).
Consistent with the DOHaD paradigm, it is likely the nutritional and psychological impact of
severe nausea and vomiting during pregnancy may reduce the resource supply to the fetus and result
in lifelong consequences for health. To date, however, few studies have examined the long-term
outcomes of children from HG pregnancies (Ayyavoo, Derraik, Hofman, and Cutfield, 2014). In a
study of healthy New Zealand children (4–11 years of age), 36 exposed to HG during pregnancy,
compared to 42 controls, showed reduced insulin sensitivity, higher fasting insulin levels, and lower
insulin-like growth factor binding-protein 1, which are associated with an increased lifelong risk
of diabetes mellitus (Ayyavoo et al., 2013). These children also had basal cortisol levels that were
22% higher than controls. Inadequate nutrition or increased prenatal stress may have reset the HPA
axis, influencing postnatal development. Therefore, as a maternal effect, HG may lead to long-term
adverse metabolic outcomes in exposed offspring (but see Finnish birth cohort; Koot et al., 2017).
Assisted Reproductive Technologies

Thus far we have considered maternal adaptations to relatively familiar environmental challenges and
their health consequences. Assisted reproductive technologies (ART), however, constitute a novel but
increasingly common environmental challenge that may be usefully explored from the perspective
of maternal effects (Golombok, 2019). Since the first IVF baby, Louise Brown, was born in 1978
(Steptoe and Edwards, 1978), an estimated five million children have been born worldwide through
ART (Hansen, Kurinczuk, Milne, de Klerk, and Bower, 2013). The most common ART treatment
involves in vitro fertilization (IVF). In industrialized societies, people who plan to have children often
delay reproduction in order to accumulate resources (Hammarberg and Clarke, 2005). Unfortu-
nately, delayed reproduction is associated with decreased fertility, leading to greater demand for ART
(Oakley, Doyle, and Maconochie, 2008), to a level that most industrialized countries cannot meet
(Hoorens, Gallo, Cave, and Grant, 2007).
While the benefits of ART treatments to couples who are unable to conceive are patent, there are
also increased risks of poorer outcomes for children conceived via ART (Fisher, Hammarberg, and
Baker, 2005; Kalra and Molinaro, 2008). The higher rate of multiple births in ART pregnancies was
thought to account for these differences, but studies examining singleton births confirm that babies
conceived by ART are more likely to be born preterm or have low birth weight than children from
unassisted conceptions (Squires and Kaplan, 2007). Placentae from ART pregnancies have not been
studied extensively ( Joy, Gannon, McClure, and Cooke, 2012), but, perhaps in response to reduced
fetal growth, placentae from ART pregnancies are heavier and have higher placental ratios (Daniel
et al., 1999; Haavaldsen, Tanbo, and Eskild, 2012). Other placental pathologies, including low-lying
placenta, placenta previa, placental abruption, and poor implantation, are more common in ART
pregnancies ( Jauniaux, Englert, Vanesse, Hiden, and Wilkin, 1990). Confirming these concerns, the
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Prenatal Parenting
British Scientific Advisory Committee of the Royal College of Obstetricians and Gynecologists
(2012) stated that the babies who emerge from IVF pregnancies are at an increased risk of adverse
outcomes, such as low birth weight, small for gestational age, and premature birth.
Maternal prenatal stress is another factor that may be associated with poorer pregnancy outcomes
in ART. During successful IVF pregnancies, couples report higher levels of stress than in successful
nonassisted pregnancies (Eugster and Vingerhoets, 1999). Moreover, the difference between ART
and non-ART pregnancies is likely to be underestimated as there is some, albeit mixed, evidence
that pre-existing stress is associated with the increased risk of unsuccessful IVF pregnancies (Mat-
thiesen, Frederiksen, Ingerslev, and Zachariae, 2011; Sanders and Bruce, 1999). In a study of 837
Danish women, higher levels of negative life events in the preceding 12 months were associated with
a reduced chance of falling pregnant in the first IVF cycle, independent of current perceived stress
and depressive symptoms (Ebbesen et al., 2009).
The associations between ART and fetal and placental growth suggest the possibility of fetal
programming in these pregnancies. However, the range of factors influencing babies conceived via
ART are extensive (inheritance of infertility, ART procedures, stress during pregnancy, nutrition,
multiple births, preterm birth, fetal growth restriction) and must be carefully considered before the
consequences of ART for child and adult health become clear (Hediger, Bell, Druschel, and Louis,
2013). There is increasing evidence that programming effects, associated particularly with adult car-
diac and metabolic heath, occur in children born from ART pregnancies (see reviews by Rinaudo
and Wang, 2012; Yeung and Druschel, 2013). Possibly because of the positive parenting practices
found in ART families, there is no evidence that children born from ARTs have poorer cognitive or
socioemotional development (see Golombok, 2019). Here, postnatal attachment relationships may
moderate the impact of prenatal cortisol exposure on postnatal cognitive development (Bergman,
Sarkar, Glover, and O’Connor, 2010).
It is clear that a full understanding of the developmental effects of ARTs cannot be assessed until
ART children become parents and grandparents themselves. Perhaps the most important considera-
tion will be the research designs developed to examine these associations. As for all pregnancies,
prenatal maternal stress and nutrition must be considered, but many factors unique to ART preg-
nancies must be considered as well: the underlying cause of infertility itself, the ART procedures
themselves (including delayed fertilization and freezing and thawing of embryos), the medications
used to induce ovulation or maintain pregnancy (Hansen, Bower, Milne, de Klerk, and Kurinczuk,
2005), and the couples’ emotional reactions to these procedures. In addition, factors known to influ-
ence ART outcomes, including the culture media used to store, grow, and protect the eggs and
sperm used in some treatments, must all be taken into account (Barnes and Sato, 1980; Trounson and
Gardner, 2000). As the number of babies conceived through ART continues to grow, and research-
ers and clinicians strive to improve maternal and neonatal outcomes, the answers to these questions
become increasingly crucial.
Perceived Stress, Objective Stress, and the Biology of Stress

There has been substantial debate about how subjective stress during pregnancy affects maternal
physiology such that fetal growth is impaired. The problem is that, while there are many reliable
and valid instruments for measuring subjective psychosocial stress, their results do not correlate
very well with biological measures of stress during pregnancy (e.g., free cortisol in saliva; Voegt-
line et al., 2013). However, both pathways—perceived stress and maternal glucocorticoids during
pregnancy—can influence maternal physiology and pregnancy outcomes. Experiences during preg-
nancy as diverse as pregnancy-specific anxiety, perceived stress, nutrient restriction, and cadmium
exposure can affect maternal and placental stress hormone levels. Because both the mother and fetus
195
are influenced by psychosocial and physiological stressors, the correlation between perceived stress
and stress hormones is often low (Sandman et al., 2012). Similarly, the commonly found association
between perceived stress and preterm birth is not always reflected in hormonal measures. For exam-
ple, Kramer and colleagues (2013) found that, while maternal ACTH was associated with cortisol,
which in turn enhanced placental CRH, neither maternal cortisol or CRH were associated with
perceived stress, maternal distress, or preterm birth. Research is beginning to establish the independ-
ent effects of perceived stress, objective stress, and the biology of stress, suggesting that several at least
partially independent pathways might have been operating on maternal physiology.
Co-adaptation: Adaptive Maternal and Fetal Programming

The evidence that prenatal adversity programs fetal physiology and affects subsequent infant, child,
and adult phenotypes is well-established. The focus throughout this literature is strongly on the fetus:
What impact does adversity in utero have on the fetus, and how does this prepare it for a harsher
postnatal environment? (Barker, 1994). A new area of research suggests that these events also program
maternal physiology, potentially leading to co-adaptation of maternal and offspring physiology. Our
argument throughout is that the prenatal environment is as much a maternal adaptation to her life
cycle as it is for the fetus (see Berghänel et al., 2017). Although parent-offspring conflict theory is
a useful perspective, there is good evidence that this conflict is tempered by parents and offspring
monitoring each other and adjusting their demands according to resource availability: Co-adaptation
(Bateson, 1994, 2017; Kölliker et al., 2012). Sandman et al. (2012) highlighted animal and human evi-
dence that the structure and function of the maternal brain are also programmed in response to stress
during pregnancy and that these changes persist over time. They proposed that changes in maternal
stress reactivity, cognitive function, and the risk of psychopathology in response to endocrine changes
throughout pregnancy prepare the mother for pregnancy, birth, and postnatal life.
In the general population, CRH levels have been associated with mental health disorders, par-
ticularly depression and anxiety (Risbrough and Stein, 2006). Yim et al. (2010) examined the
relation between placental CRH (pCRH) and postnatal depressive symptoms, finding that women
with elevated pCRH levels at 25 weeks of pregnancy were more likely to develop depressive symp-
toms postnatally. This may be why problems during pregnancy, anxiety, depression, stressful life
events, and low levels of social support are closely associated with postnatal depression (Beck, 2001;
O’Hara and Swain, 1996; Robertson, Grace, Wallington, and Stewart, 2004). Moreover, mothers
suffering from postnatal depression reduce their investment in their new baby (Hagen, 1999). Thus,
maternal stress during pregnancy may signal a risky or uncertain environment to which both the
mother’s and fetus’ physiology respond (Sandman et al., 2012). This dual programming ultimately
improves the fit between mother, infant, and environment. Since postnatal depression affects 10%
to 15% of new mothers (Hagen, 1999), this adaptive interpretation of maternal stress may have
clinical significance.
In early postnatal development there is a remarkable synchrony between maternal behavior and
neonatal sensory perception (Brazelton et al., 1974; Trevarthen, 2012; Tronick, 2007). During the long
period of slow development that is human childhood this ensures a coordinated mother-infant rela-
tionship and transduction of social environmental cues via attachment relationships, among other sys-
tems (Chisholm, 1999b, 2017). Ultimately, these “synchronized capabilities” translate to the improved
probability of survival and fitness benefits for the mother and infant (Simpson and Belsky, 2008).
Therefore, it makes sense that the mother’s, not only the fetus’s, behavior and physiology are adapted
to the environment they share. Future research must examine the interaction between cues in different
life stages, as well as intergenerational inheritance, as it is likely the best fit between an organism and
its ecology is affected by many environmental cues across different time frames (Nettle et al., 2013).
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Childhood Psychosocial Stress: A Developmental

Predictor of Maternal Environment
The critical influence of childhood experience on subsequent growth, development, and adult health
and behavior is central to many disciplines, including evolutionary ecology (Henry and Ulijaszek,
1996), evolutionary psychology (Belsky et al., 1991), ethology (Bateson and Martin, 1999), develop-
mental health (Keating and Hertzman, 1999), developmental psychology (Repetti et al., 2002), and
epidemiology (Kuh and Ben-Shlomo, 1997). In humans, as with other animal species (Meaney and
Szyf, 2005), the parental developmental environment and parenting behavior have particularly strong
influences on offspring physical and mental phenotypes.
Life history theory predicts that the flow of resources to the fetus (parental investment) dur-
ing pregnancy will vary according to the mother’s developmental environment, her reproductive
strategy, her currently available material and socioemotional resources, and the resources likely to
be available in the future (Chisholm, 1993, 1999b; Chisholm and Coall, 2008; Coall and Chisholm,
2003; Ellison, 2005; Gluckman, Hanson, and Beedle, 2007; Jones, 2005; Kuzawa, 2007; Walker et al.,
2006; Wells, 2003; Worthman, 1999; Worthman and Kuzara, 2005). Evidence is emerging, however,
that the developmental trajectories entrained by early stress may reduce the maternal allocation of
resources to the fetus during pregnancy, thus limiting fetal development and increasing the risk of
poor health outcomes (Coall and Chisholm, 2003, 2010). Consistent with this interpretation, women
who experienced two types of childhood trauma (e.g., maltreatment, abuse, neglect) had placental
CRH concentrations 25% higher than a no trauma group (Moog et al., 2015). The authors interpret
this as suggesting early maternal stress may influence the offspring phenotype through placental/fetal
stress reactivity.
Research with the Adverse Childhood Experiences (ACE) Study demonstrates that early stress
is associated with higher adult disease and mortality rates (Anda et al., 1999; Felitti et al., 1998).
After adjustment for age, ethnicity, gender, and education, adverse childhood experiences (e.g., sexual
abuse, domestic violence, imprisonment of a family member) are risk factors for adult diseases, such
as severe obesity, smoking, physical inactivity, and poor self-rated health. Compared to those with
none, individuals who experienced four or more adverse experiences are more likely to develop
diseases, such as ischemic heart disease, stroke, diabetes, cancer, and chronic bronchitis (Felitti et al.,
1998). Moreover, these results have been replicated in four birth cohorts from 1900 to 1978 (Dube,
Anda, Felitti, Edwards, and Williamson, 2002), with the risk increasing in a dose-dependent manner
in association with the number of childhood stressors experienced (Felitti et al., 1998).
There is convincing (albeit retrospective) evidence that the number of adverse childhood experi-
ences predicts increased adult mortality rates, but some early stressors seem to matter more than oth-
ers. This is probably because they reflect or are symptomatic of a child’s generally risky or uncertain
family environment. In particular, childhood sexual, emotional or physical abuse, and witnessing
physical violence between parents, tend to be associated with other adverse experiences during
childhood (Dube et al., 2002; Gladstone, Parker, Wilhelm, Mitchell, and Austen, 1999). For example,
while witnessing parental violence is associated with higher levels of psychological distress and lower
levels of social adjustment in adulthood, this association is mediated by the amount of parental care
and warmth experienced (Henning, Leitenberg, Coffey, Bennett, and Jankowski, 1997; Henning,
Leitenberg, Coffey, Turner, and Bennett, 1996). Moreover, individuals who experience adverse child-
hood events appear to have more negative life events and experience more anxiety and depression
during pregnancy (Benedict, Paine, Paine, Brandt, and Stallings, 1999; Grimstad and Schei, 1999).
The early psychosocial environment may also affect fetal development via maternal body com-
position. Psychosocial stressors, such as neglect, abuse, and unsupportive home environments, have
been associated with childhood obesity (Strauss, 1999) and rapid weight gain during childhood. In a
197
study of 5,399 Swedish school children (2,661 girls), girls with the most rapid weight gain between
7 and 10 years of age also had the highest prevalence of social, behavioral, and learning problems
(Mellbin and Vuille, 1989). In a study of 756 Danish school children, those who were perceived by
their teachers to receive little parental support had higher rates of obesity and were seven times more
likely to be obese at 20 years of age (Lissau and Sorensen, 1994). Thus, early stress may ultimately
influence fetal growth through its impact on childhood weight gain and, subsequently, maternal body
composition during pregnancy.
Whether the mechanism is via chronic stress or increased weight gain, it is likely that these
childhood experiences are embodied (Chisholm, Burbank, Coall, and Gemmiti, 2005; Hertzman,
1999; Hertzman, Power, Matthews, and Manor, 2001; Krieger, 2001) in the form of altered stress
reactivity (Bremner et al., 2003). Consistent with this evolutionary synthesis, increased stress reac-
tivity is associated with lower birth weight (Clark et al., 1996; Ward et al., 2004), lower parental
responsiveness (Haley and Stansbury, 2003; Repetti et al., 2002), earlier menarche (Boyce and Ellis,
2005; Ellis, 2004), and earlier first sexual intercourse (Brody, 2002). The resultant increased levels
of glucocorticoids can reduce insulin sensitivity and increase fat deposition (Brindley and Rolland,
1989; Tsigos and Chrousos, 2002). Furthermore, psychosocial stress is a risk factor for developing the
metabolic syndrome in childhood, adolescence, and adulthood (Eisenmann, 2003; Hjemdahl, 2002;
Rosmond, 2005). Therefore, via the actions of stress hormones, childhood psychosocial stress may
be associated with weight gain throughout the lifespan and an increased risk of adult diseases, which
in turn are associated with reduced fetal growth in the next generation (Lawlor, Davey Smith, and
Ebrahim, 2003).
Predicting Future Environments: Predictive Adaptive Responses

Earlier, we referred only in passing to the concept of “predictive adaptive responses” (PARs) because
it deserves special discussion. The concept needs clarification and analysis because it has different
names in different fields, its meaning is not always made clear, and there are different ways of inter-
preting the concept. These are significant issues because PARs could have critical implications for
understanding and reducing the prenatal origins of health and disease. As commonly understood, a
PAR (in the context of this chapter) is a developmental response by the fetus to a uterine environ-
mental cue such that—if it uses this cue to correctly anticipate or “predict” the future environment—
the developmental response will be evolutionarily adaptive in its postnatal environment. Otherwise,
if the fetal response does not correctly “predict” the postnatal environment, whether because the sig-
nal was not reliable, or the environment changed, there will be a maladaptive mismatch between the
developmental calibration and postnatal environment (Bateson et al., 2004; Gluckman et al., 2005).
PARs have stimulated extensive discussion and analysis about what information from its prenatal
environment the fetus extracts and responds to. This discussion has provided great stimulus to the
field but is currently unresolved. As we have emphasized throughout this chapter, the fetal environ-
ment includes information not only from the mother’s current environment, but her own develop-
mental environment and even her mother’s and her grandmothers’ as well. Therefore, the question
arises: How well can we expect 9 months of pregnancy to predict the child and adult environments
of the future? ( Jones, 2005; Kuzawa, 2005; Worthman and Kuzara, 2005).
Alternative (although not mutually exclusive) models of how adverse prenatal environments affect
development and postnatal health have emerged. In their original form, PARs supposedly prepare
organisms for future environments and provide “weather forecasts” via “fetal programming”. How-
ever, as we discussed in the context of life history theory, under harsh environments the priority is
simply to survive. In this light, various models portray physiological changes during pregnancy to
a challenging environment as “making the best of a bad start” (Berghänel et al., 2017; Jones, 2005;
198
Prenatal Parenting
Rickard, Frankenhuis, and Nettle, 2014; Vitzthum, 2001). Focusing on survival is a form of down-
side risk protection. Individuals born into an adverse environment must change their physiology to
ensure survival and avoiding lineage extinction rather than to “predict” a future environment that
may never exist (Chisholm and Coall, 2008). Still other models see early adversity as a detrimen-
tal disruption no matter what its environment is like. These models are often referred to as “silver
spoon” (the converse as “leaden/wooden spoon”) models (Grafen, 1988) and suggest that there is
a lifelong fitness advantage or disadvantage associated with the early environments that individuals
embody.
Throughout this chapter, these different PAR models have been discussed in specific examples. At
present, attempts to test the different predictions are underway, with some support for the silver ver-
sus leaden spoon model (Hayward and Lummaa, 2013). It is important to recognize that these models
currently ignore the competing demands of the mother and fetus. Thus, the application of parent-
offspring conflict theory to understanding the adaptive functions of prenatal environment may also
be a fruitful area of research (Del Giudice, 2012; Gangestad, Caldwell Hooper, and Eaton, 2012).
Attempts to rationalize the PAR models into manageable categories are under way (Monaghan,
2008; Nettle et al., 2013; Wells, 2012), but consensus is some way off. Some useful results have come
from modeling comparing the “weather forecasting” and “leaden spoon” PAR models. Nettle et al.
(2013) concluded that their model predicted (1) developmental plasticity would evolve to receive
information from the maximum number of environmental cues available (especially if the cues were
not highly reliable) and (2) that the reliability of cues from year to year would need to be exceptional
to have adaptive value across an individual’s reproductive lifespan or even shorter durations. The
relation between environmental risk and how it is embodied has been developed. Indeed, it may be
useful to study the causal pathways by examining the interaction between both the environmental
risk (external PAR) and how the risk is embodied physiologically (internal PAR; Hartman, Li, Net-
tle, and Belsky, 2017). More empirical and theoretical work is needed to turn the promising field of
PARs into strategies for interventions to reduce the negative prenatal impact on offspring pheno-
type. We are currently at the threshold of an exciting new area of research.
Conclusions
In this chapter, we have tried to make two overarching points: (1) the study of prenatal influences is
no longer a “grey area” of modern science because (2) evolutionary theory—our only scientific the-
ory of life (and development)—provides the “bio-logical” basis for coordinating the multidisciplinary
evidence for prenatal influences and making sense of the patterns that emerge as maternal effects.
We have endeavored to present an interdisciplinary perspective within an evolutionary, life history
theory framework. Maternal stress, nutrition, and exposure to toxins all provide crucial information
about the harshness of the environment into which a baby will be born. These cues provide the
fetus with a guide to its optimal behavioral/physiological phenotype for survival and reproduction
in its postnatal environment. When this adjustment is made during sensitive periods of development,
such as prenatally, the short-term benefit of changes that ensure continued survival may be traded off
against the longer-term costs of lifelong and intergenerational health.
Acknowledgments
We thank Tom Dickins for valuable contributions, Toni Wain for editorial assistance, and Renee
Khan-Passetti, Del Periera, Michelle Cannon, and Ruben Phillips for their invaluable research assis-
tance. We also gratefully acknowledge support from the National Evolutionary Synthesis Center,
Duke University.
199
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6
THE SOCIAL
NEUROENDOCRINOLOGY
OF HUMAN PARENTING
Ruth Feldman
Introduction
Nonhuman mammalian mothering is hormone-dependent; hormonal changes occurring during
pregnancy and labor causally determine the expression of maternal behavior. Studies in animal mod-
els have shown that experimental manipulations on the expression of key hormones markedly alter
or totally eliminate the expression of maternal care (Feldman, 2012b, 2016; Lonstein, Lévy, and
Fleming, 2015; Pryce, 1996; Rosenblatt, 1994; Rosenblatt, 2003). Research in rodents describes the
critical role of oxytocin (OT) and prolactin (PRL), which undergo substantial changes during late
pregnancy (PRL) and surge at birth (OT), for the onset of maternal behavior. In parallel, hormones
associated with the stress response, particularly corticosterone (cortisol in humans), modulate mater-
nal vigilance and active protection of offspring (Brummelte and Galea, 2010; Mann and Bridges,
2001; Pedersen and Prange, 1985). Finally, animal studies point to the involvement of vasopres-
sin (AVP) and testosterone (T) in the emergence of fatherhood and the expression of mammalian
paternal care (Carter, 2014; Wynne-Edwards, 2001). In combination with sex-related hormones
(estradiol, progesterone), these hormones establish the neuroendocrine milieu that enables rodent
mothers—and fathers in the 3%–5% of mammalian species who are biparental (Braun and Cham-
pagne, 2014; Kleiman, 1977)—to parent. The hormones of parenting enable parents to recognize
infants as rewarding stimuli, protect infants from harm, nurse, express species-typical parental behav-
ior, and provide external regulation for the infant’s immature regulatory systems, including sleep
organization, thermoregulation, autonomic functions, attention, and exploration (Feldman, 2016;
Hofer, 1995a,b; Numan and Stolzenberg, 2009). These hormones also help parents usher their young
into the social niche and accommodate its distinct features. Finally, the neuroendocrinology of par-
enting promotes the infant’s ability to manage life in harsh ecologies via mechanisms of endocrine fit
and the effects of parental hormones on the infant’s brain maturation and social fittedness (Feldman,
Monakhov, Pratt, and Ebstein, 2016).
Human parenting is not hormone-dependent; however, hormonal changes during pregnancy,
birth, and the postpartum period prime and accompany the expression of parenting, sculpting the
development of the parent-child attachment and its long-term effects on the infant’s brain and behav-
ior (Apter-Levi et al., 2016; Galbally, Lewis, IJzendoorn, and Permezel, 2011; Feldman, 2016, 2017;
Gordon, Zagoory-Sharon, Leckman, and Feldman, 2010b, 2010c). Humans’ large associative cortex,
neural plasticity, and massive limbic-cortical projections enable bottom-up, behavior-based process-
ing so that committed parental care can trigger the hormones of parenting even without pregnancy
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and childbirth; for instance in primary-caregiving fathers or adoptive parents (Abraham et al., 2014;
Bick, Dozier, Bernard, Grasso, and Simons, 2013). Yet, as parenting is the only social phenomenon
observed across species and taxa, there is no other sociobiological process that can provide a clearer
lens into evolution as it occurs and shed light on the roots of humans’ collaborative, empathic, and
relational abilities (Feldman, 2015a, 2015b; Rilling and Young, 2014). Furthermore, the parent-infant
interface marks the arena where Darwin (1859) has initially proposed structural and functional brain
adaptations take place. The neuroendocrinology of human parental care, addressing the hormonal
changes that accompany parenting and their neural, behavioral, and mental correlates, may thus
afford a unique perspective on the evolution of human sociality, highlighting both its conserved and
human-specific features (Feldman, 2015b, 2016).
In addition to a special viewpoint on human sociality, the neuroendocrinology of parenting
provides a unique angle on neural plasticity not available from other topics in neuroscience. Preg-
nancy and the postpartum mark the period of greatest plasticity in the adult brain (Leuner, Glasper,
and Gould, 2010), and such plasticity is observed not only in the maternal but also in the paternal
brain, with fathers’ investment in childrearing increasing plasticity not only in the father’s brain but
also in the brain of his offspring (Braun and Champagne, 2014). Parenting, therefore, enables the
investigation of endocrine systems and neural networks as they reorganize in the parent’s brain and
research on how successful versus less optimal reorganization directly impacts the infant’s emerging
endocrine systems and neurobiological outcomes. Furthermore, parenting is perhaps the most highly
conserved social phenomenon, accompanied by similar species-typical behaviors that are triggered
by the same neuroendocrine events across mammalian evolution (Feldman, 2015a, 2015b; Rilling
and Young, 2014). This is particularly the case with regards to the ancient oxytocin system, which
supports parenting, group cohesion, and stress management in species ranging from nematodes to
humans, including birds, fish, and Caenorhabditis elegans (Feldman et al., 2016; Goodson, 2013). In
comparison with the neuroscience of parenting, assessing the brain basis of emotions, memory, or
categorization involves a much greater conceptual leap; these constructs are heuristic, much farther
away from their biological underpinnings or evolutionary origins, and are constructed online by
humans’ higher-order representations. When research on the neurobiology of parenting is coupled
with detailed observations of parental behavior in the natural habitat, it provides a closer setting to
that of nonhuman mammals as compared to most other domains in neuroscience. Yet, notwithstand-
ing the similarity of human parental care with that of other mammals, human parenting is also greatly
influenced by humans’ higher-order cognitive abilities and cultural construals. Thus, research on the
neurobiological basis of parenting affords a unique view on the integration of mammalian-general
and human-specific features on key biological processes. Because parenting is triggered by the same
hormones across mammalian species, the hormonal basis of human parenting provides a scientifically
plausible tool for mechanistic research as compared to other domains of inquiry (e.g., the neurobiol-
ogy of psychiatric illness); hence, the neuroendocrine basis of parenting is among the few topics that
offer a uniform trajectory of empirical investigation across the evolutionary ladder.
The social neuroendocrinology of human parental care comprises four main lines of research.
The first assesses the hormonal basis of parenting in healthy parents. Most of this line of research
focuses on the hormones of motherhood; the typical hormonal changes occurring in mothers dur-
ing pregnancy, the postpartum period, and across the early years. Often, these studies examine not
only mean-level changes but also individual differences in hormonal levels and their links with
maternal behavior, attitudes, or personality traits. To date, the hormone receiving the most research
in relation to mothering is cortisol (CT), possibly due to its reliable bioassay in saliva that has been
available for some time. Yet, with the development of more sensitive bioassays, studies have also
looked at OT, PRL, AVP, T, salivary alpha amylase (sAA), beta endorphin, and immune biomarkers
(IL-6, salivary IgA). Another area of research within this global line is the hormones of fatherhood.
Fathering in general, and the neurobiology of fathering in particular, has received much less research
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as compared to mothering in both humans and other mammals, but the hormonal basis of father-
ing is a developing area of research and a growing body of literature is beginning to assess fathers’
hormones in relation to observed paternal behavior.
The second line of research on parental hormones examines “endocrine fit”—the “match” or
synchrony between parent and child hormonal levels. Such biological synchrony is conceptualized
as one mechanism by which the parental affiliative system is transferred to the child or as a way in
which parents signal environmental danger to their offspring (Feldman, 2016, 2017; Pratt et al., 2017;
Bornstein, 2013). Our biobehavioral synchrony model contends that such hormonal synchrony, while
genetically informed, matures in the context of coordinated social behavior and shared parent-child
social experiences (Feldman, 2012c, 2015a, 2016, 2017).
A third line views the neurobiology of human parenting as a global area of research, which
includes the brain networks, hormonal systems, and specific behaviors that activate in mothers or
fathers with the birth of an infant. Several studies in this line test associations between activations of
specific brain areas in the parental brain with parenting-related hormones, including OT, AVP, CT, or
T (for a review see Feldman, 2015b). It is hypothesized that the “mammalian parenting network”, the
brain regions that support mammalian parenting, initiates its activation through sensitization by the
hormones of pregnancy (Numan and Young, 2016), thus indicating that hormonal changes trigger
neural alterations that define the neurobiology of parenting.
The last area of research in the neuroendocrinology of parenting, and by no means the least abun-
dant, addresses parental hormones under high-risk conditions, whether the risk stems from mother-
related conditions (e.g., maternal depression, anxiety), child-related conditions (prematurity, autism
spectrum disorders), or contextual adversities (poverty, abuse, war exposure). Several studies of high-
risk parenting compared a high-risk cohort to a typical group, whereas others use a correlational
design within the high-risk sample. Of note, very few studies examine fathers’ hormones in high-risk
contexts, and a smaller number of studies address the fit between high-risk parents and children.
The following literature reviews the hormonal basis of human parenting keeping in mind its
evolutionary origins. Due to the extensive literature on the topic, this review is by no mean com-
prehensive and addresses mainly parental hormones in the first years of life, with a focus on infancy,
and follows the four lines of research outlined above. Consistent with the comparative approach, only
studies that measure hormones in relation to observed parental behavior are reviewed. This approach
accords with the view that in humans the neurobiology of parenting may trigger, not only through
pregnancy and lactation, but via commitment to caregiving and active involvement in daily interac-
tions with the child. Thus, similar to the long line of “cooperative breeding” in primates, the human
“village” (grandparents, “aunties”, male partners, adoptive parents, godparents) can rear the child
through bottom-up, behavior-based activation of the neurobiology of parenting, consistent with
Hrdy’s (2007) suggestion that in species such as mammals parenting is behavior (Feldman, 2012d).
This conceptualization of a bottom-up activation of the neurobiological systems that support par-
enting via parenting behavior is consistent with findings that mother-child synchrony expresses in
the brain as brain-to-brain synchrony of gamma-band oscillations, with gamma marking a distinct
bottom-up, behavior-based mechanism (Levy, Goldstein, and Feldman, 2017).
Hormonal Basis of Human Mothering and

Fathering in Low-Risk Contexts
This section describes each hormonal system separately and addresses findings related to mother-
ing and fathering for each hormone. Following, a section discusses endocrine fit. Overall, hor-
mones of parenting are divided into the “affiliative hormones” module, which includes mainly OT
but also AVP and PRL, hormones that support the formation of parent-infant bonding, maintain
attachments, and buttress human sociality (Carter, 2014; Fleming, Ruble, Krieger, and Wong, 1997;
222
Numan, 2006). The other group involves stress-related hormones, mainly CT but also salivary alpha
amylase (sAA) or immune biomarkers (IL-6, IgA). A third group considers sex-related hormones
(T, progesterone, estradiol). These three classes of hormones are not independent in their action, and
studies have shown co-dependence and mutual influences of these hormones on each other, mainly
in complex, nonlinear ways that require much further research (Gordon, Zagoory-Sharon, Leckman,
and Feldman, 2010a; Gordon et al., 2017). Research in rodents has further shown that the expres-
sion of maternal behavior functions on both the affiliation and stress neuroendocrine systems, with
maternal licking and grooming building the expression of both oxytocin receptor densities in the
nucleus accumbens (Francis, Champagne, and Meaney, 2000) and glucocorticoid receptors in the
hippocampus of the infant’s brain (Liu et al., 1997).
Figure 6.1 describes OT and CT as the main, most well-research hormones of parenting, their
mutual influences on other hormones, and their behavioral correlates.
Oxytocin
Oxytocin is considered the main neuroendocrine system supporting the formation and main-
tenance of the parent-infant bond and a central trigger for the expression of parental behavior
Figure 6.1 Key hormones of human parenting.
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(Feldman, 2015b, 2016, 2017). It is an integrative system that provides a neuroendocrine milieu for
the functioning of multiple hormones. Numerous hormones operate in concert to support parental
care, but OT maintains crosstalk with other hormones in the context of parenting. We found that
OT links and interacts with a range of hormones in supporting parenting behavior, including AVP
(Apter-Levi, Zagoory-Sharon, and Feldman, 2014), CT (Gordon et al., 2010a), T (Gordon et al.,
2017; Weisman, Zagoory-Sharon, and Feldman, 2014), beta endorphin, and IL-6 (Ulmer-Yaniv
et al., 2016), highlighting the role of OT in integrating the affiliation, reward, stress, and immune
systems in support of parenting.
OT is a nine-amino-acid neuropeptide hormone, which presumably evolved from the ancient
vasotocin molecule via gene duplication in jawed fish approximately 650 million years ago (Feld-
man et al., 2016). OT is implicated in sociality across vertebrate evolution and substantial research in
rodents has pinpointed its role in birth, lactation, and maternal care in mammals (Carter, 2014; Feld-
man et al., 2016; Lee, Macbeth, Pagani, and Young, 2009; Lim and Young, 2006). Studies have exam-
ined peripheral levels of OT—in plasma, saliva, urine, and to a lesser extent in cerebrospinal fluid—in
relation to human parenting, aided by the availability of new and more reliable immunoassay kits.
Associations between brain OT and its peripheral indices are not fully clear, but human studies have
lent support to the use of peripheral OT by demonstrating marked increase in peripheral OT when
individuals inhale OT, which has shown to impact the brain’s OT system (Neumann, Maloumby,
Beiderbeck, Lukas, and Landgraf, 2013; Weisman, Zagoory-Sharon, and Feldman, 2012), associations
between plasma OT and more efficient variants of the oxytocin receptor gene (OXTR; Feldman
et al., 2012), and correlations between plasma and salivary OT with brain activations in areas rich in
oxytocin receptors, including the hypothalamus or the amygdala (Abraham et al., 2014; Atzil, Hen-
dler, and Feldman, 2011; Strathearn, Fonagy, Amico, and Montague, 2009). The distributions of OT
receptors in the brain are species-specific (Stevens, Wiesman, Feldman, Hurley, and Taber, 2013) and
the nature of the relation between central and peripheral OT is a matter of ongoing debate, but an
accumulating body of research has shown that variability in peripheral OT is meaningfully linked
with the expression of maternal and paternal behavior in ways that are consistent with research on
central OT in rodents.
In the first longitudinal study of OT and parenting behavior, we followed healthy women across
pregnancy and the postpartum and measured plasma OT and cortisol (CT) at three time-points;
first trimester of pregnancy, third trimester, and the first postpartum month when we also observed
mothers interact with their infant in the home environment. We used an in-depth interview to
measure maternal thoughts, preoccupations, and attachment representations. OT levels increase in
early pregnancy and stay stably high across pregnancy and the early postpartum. OT levels during
the first trimester predict the expression of the human species-typical maternal behavior, suggesting
a priming effect of OT in humans. In addition, OT and CT across pregnancy are unrelated at any
time-point, but CT levels independently predict a decrease in the expression of maternal postpartum
behavior, indicating a joint effect of the two main hormonal systems on maternal postpartum behav-
ior similar to that found in rodents (Feldman, Weller, Zagoory-Sharon, and Levine, 2007). Another
evidence for a priming effect was found when mothers with higher OT during late pregnancy
reported greater bonding to their fetus (Levine, Zagoory-Sharon, Feldman, Lewis, and Weller, 2007).
It has been suggested that mothers develop clear representations of their unborn child during the
last weeks of pregnancy and a failure to do so, due to depression or risk for preterm delivery, impairs
the emerging attachment (Hart and McMahon, 2006; Pisoni et al., 2016). OT in late pregnancy also
predicts increased maternal preoccupations and more positive representations of the infant and the
attachment relationship. Others found similar links between attachment representations and higher
OT levels in pregnancy and the postpartum (Eapen et al., 2014). These findings, therefore, add the
representational component to the priming effect in other mammals and show that in humans the
higher-order cognitive dimension of parenting is similarly triggered by the oxytocinergic system.
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We next followed first-time mothers and fathers from the first postpartum month to 6 months
postpartum and measured plasma OT in relation to maternal and paternal parenting behavior. Post-
partum parents had much higher OT levels as compared to individuals who were not parents or
in a romantic relationship, indicating that OT levels increase when individuals become attached. In
addition, no difference was found between mothers’ and fathers’ baseline OT, even when mothers
were breastfeeding. Similar to the first study, OT levels in individuals were highly stable over time,
and mutual influences between partners’ OT were observed both within and across time-points
(Gordon, Zagoory-Sharon, Leckman, and Feldman, 2010b). This result suggests that plasma OT
may tap a “trait-like” dimension of the individual which is individually stable yet shaped by close
attachment relationships (Schneiderman, Kanat-Maymon, Ebstein, and Feldman, 2014). OT levels
in mother and father were related to the parent-specific behavioral repertoire (Feldman, Gordon,
Schneiderman, Weisman, and Zagoory-Sharon, 2010). Maternal OT was related to the “affiliative
parenting” constellation typical of mothers, including gaze to infant face, expression of positive affect,
“motherese” high-pitched vocalizations, and affectionate touch, the human parallel of “licking-and-
grooming” (Meaney, 2001). In contrast, fathers’ OT was linked with “stimulatory parenting”, a style
typical of mammalian fathering, which included directing attention to the environment, stimulatory
contact, and high, unpredictable positive arousal (Gordon et al., 2010b; Naber, van IJzendoorn, Des-
champs, van Engeland, and Bakermans-Kranenburg, 2010). These findings, the first to test plasma
OT in new fathers, show comparable OT levels in mothers and fathers and indicate that fathers may
be just as biologically prepared to care for infants. In comparison with mothers’, fathers’ parenting
style is expressed via a distinct set of behaviors that prepare infants to explore their physical environ-
ment, rather than focus on the dynamics of face-to-face relationships, and this paternal repertoire is
linked with father’s OT. Finally, observing triadic family interactions between these parents and their
6-month-old infants we found that OT predicted triadic synchrony, the coordination of behavior
among the three family members (Gordon et al., 2010a), highlighting the role of OT as an integrator
of social behavior among affiliative units.
Comparing OT in plasma, saliva, and urine in a group of mothers and fathers (not couples) and
their 4- to 6-month-old infants, we found no differences in baseline OT levels in plasma, saliva, or
urine between mothers and fathers. Of note, OT levels in saliva and plasma showed mid-level cor-
relations, and similar associations were found in several other samples, but no correlations emerged
between these indices and urinary OT, possibly since urinary hormone concentrations travel through
a different bodily route. Both plasma and salivary OT were related to higher parent-infant synchrony
(Feldman, Gordon, and Zagoory-Sharon, 2011). Similar associations obtain between mothers’ plasma
OT response (change from baseline to post-interaction) and gaze coordination and gaze duration
between mothers and their 7-month-old infants (Kim, Fonagy, Koos, Dorsett, and Strathearn, 2014).
In contrast to plasma and salivary OT, urinary OT predicts greater parental stress and attachment
anxiety, highlighting the dual role of OT in linking to both the affiliative and the anxiety/vigilance
components of parenting (Feldman, Gordon, et al., 2011). These findings are consistent with data
from a large cohort of women and men, both parents and nonparents, which showed that in women
plasma OT levels are positively associated with measures of attachment anxiety (Weisman, Zagoory-
Sharon, Schneiderman, Gordon, and Feldman, 2013) as well as with research in animal models
pointing to the role of OT in modulating stress and anxiety (Neumann and Slattery, 2016). Urinary
OT has been linked with infant caregiving behavior in cooperative-breeding marmoset monkeys
(Finkenwirth, Martins, Deschner, and Burkart, 2016), and listening to mother’s voice during a stress
paradigm elevated children’s urinary OT and enabled better stress management (L. J. Seltzer, Ziegler,
and Pollak, 2010), validating urinary OT as a biomarker of parental care.
In the first months of life, parental plasma OT levels are associated with allelic variability on the
OXTR on key SNPs associated with attachment, including OXTR (rs2254298 and rs1042778)
and CD38 (rs3796863), as well as with more parental touch and greater gaze synchrony, the two
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main features of close attachment bonds (Feldman et al., 2012). These findings are consistent with
numerous studies showing associations between more functional OXTR variants with sensitive par-
enting and attachment security throughout life (Bakermans-Kranenburg and van IJzendoorn, 2008;
Feldman et al., 2016; Raby, Cicchetti, Carlson, Egeland, and Collins, 2013).
Consistent with findings on the cross-generational transmission of OT functionality in rodents,
which is mediated by maternal behavior (Champagne, 2008; Champagne, Diorio, Sharma, and
Meaney, 2001), parental OT in the first months of life shapes the infant’s OT system and this cross-
generation transmission is similarly moderated by synchronous parenting; links between parent and
child OT are found only when parents engage in synchronous interactions, but not when minimal
synchrony is observed (Feldman, Gordon, and Zagoory-Sharon, 2010). In another study, following
parents and infants from the first month of life to the preschool stage, we similarly found cross-
generational transmission over time and attachment bonds. Parents’ plasma OT in the postpartum
predicted child salivary OT at preschool as mediated by early parental behavior. Furthermore, paren-
tal OT and synchronous parenting shaped not only the child’s OT but also the degree of reciproc-
ity and positive engagement during interactions with the first best friend at 3–4 years, consistent
with attachment theory’s predictions that parental care shapes children’s ability to enter subsequent
attachments in their lives, with friends, mentors, and romantic partners culminating in their ability
to parent the next generation (Feldman, 2012b; Feldman, Gordon, Influs, Gutbir, and Ebstein, 2013).
The OT response is sensitive to parental touch. Mothers and fathers were tested in the 10-minute
“play and touch” paradigm, where a parent interacts with the infant freely and is instructed to “touch
your infant as you normally do”. Mothers who provided abundant amounts of affectionate touch,
but not those who provided little touch, showed an OT increase following the interaction. In parallel,
fathers who had high levels of stimulatory contact, but not those who showed little touch, increased
their OT levels (Feldman et al., 2010). Furthermore, infants as young as 4 months displayed OT
increases following synchronous interactions with their parents (Feldman, Gordon, and Zagoory-
Sharon, 2010). Like rodents, human parent-specific touch, when provided in abundance, elicits OT
response in parents, which, in turn, elicits a parallel OT response from the infant, priming the infant’s
OT system to respond to pleasurable social touch within future attachment relationships.
Finally, animal studies have suggested that neuroendocrine changes in mothers during pregnancy
and the postpartum provide a template for pair bonding, and, thus, there is continuity between hor-
monal processes implicated in parental and pair bonding (Numan and Young, 2016). We measured
plasma OT, beta endorphin, and IL-6—biomarkers of the affiliation, reward, and immune systems—
in a group of first-time parents and their 3-month-old infants, a group of new romantic partners who
had been together for 3 months, and unattached singles. Synchrony between parents and infants and
among new lovers was microcoded. We found that all hormonal systems underwent changes with
the formation of new attachment bonds. OT increased in parents but was highest in new lovers. In
contrast, both beta endorphin and IL-6 were highest in parents, lowest in singles, and at mid-level in
lovers. In addition to increase, biomarkers of affiliation, reward, and stress management coalesced, and
the correlations between them became tighter during periods of bond formation. Finally, the effects
of beta endorphin and IL-6 on behavioral synchrony were mediated by the oxytocin system (Ulmer-
Yaniv et al., 2016). These findings, combined with the aforementioned continuity between parental
and filial attachment (humans’ attachment to their close friends) highlight the integrative role of OT
across human affiliative bonds, as mediated by sensitive parenting (Feldman, 2012a, 2012c).
Research has also investigated the effects of intranasal OT administration on a plethora of human
social functions. Several studies showed effects of OT administration on increasing fathers’ energetic
and object-focused interactions with their toddlers (Naber et al., 2010) or on brain response to infant
cry and laughter among nonparents (e.g., Riem et al., 2011, 2012).
In the context of parental hormones and behavior, we administered OT to 35 fathers of 5-month-
old infants in a double-blind placebo-controlled within-subject design, examined paternal and infant
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hormones, and microcoded social behavior. As expected, intranasal OT administration markedly

increased fathers’ salivary OT; however, surprisingly, OT administration to parent increased the
infant’s salivary OT by 30-fold, although infants were taken out of the room where fathers inhaled
OT and remained outside for the 45-minute waiting period. This measurable increase in infant OT
may point to mechanisms of chemosignaling between parent and child which require much fur-
ther research. Under the OT condition, subtle differences in parent and child’s social behavior were
observed; fathers touched infants more and gently reoriented infants back to the joint play when
they averted gaze. Infants gazed at their fathers for longer durations and engaged in longer episodes
of joint exploration. Autonomic signs were also higher in the OT condition, and both father and
infant increased cardiac vagal tone, a biomarker of social engagement (Weisman et al., 2012).
OT administration to fathers also impacted the infant’s CT levels as mediated by father-infant
synchrony. Among infants experiencing high synchrony, paternal still-face increased CT production
and elevated infants’ social gaze to the nonattentive parent. However, among infants experiencing
low paternal synchrony, OT reduced the infant’s stress response and decreased social gaze to the
father during the still-face phase (Weisman, Zagoory-Sharon, and Feldman, 2013). As OT increases
the social salience of events (Shamay-Tsoory and Abu-Akel, 2016), it is possible that among infants
who internalized an engaged and available paternal style, the OT condition enhanced their social
attention to father’s communicative failure.
Vasopressin
AVP is a structurally similar neuropeptide to OT, both originating from the ancient vasotocin mol-
ecule, and both implicated in mammalian fathering (Carter, 2014), but little research has examined
peripheral AVP in relation to human parenting. Studies in rodents suggest that AVP is involved in
physiological changes in AVP in fathers may mediate changes in energy balance and stress reactivity
that are required for the onset of fathering (for review: Bales and Saltzman, 2016; Saltzman and Zie-
gler, 2014). AVP is associated with male bonding and defensive and territorial behavior (Bielsky, Hu,
Ren, Terwilliger, and Young, 2005), and AVP promotes social recognition in both rodents (Cald-
well, Lee, Macbeth, and Young, 2008) and human males (Guastella, Kenyon, Unkelbach, Alvares, and
Hickie, 2011). Regions characterized as part of the AVP circuitry are implicated in socio-cognitive
processes in both humans and rodents (Goodson and Thompson, 2010). This AVP-brain associations
may represent elevated AVP-dependent vigilance, which supports father’s ability to read the inten-
tion of others to defend mother and young (Thompson, George, Walton, Orr, and Benson, 2006). In
contrast, AVP supports the mother’s ability to befriend others. Thus, AVP may prompt differential
social strategies in social contexts in women and men (Thompson et al., 2006).
Research on AVP is predominantly male oriented as AVP has been mostly studied in the context
of autism and aggression. Variability on the AVP receptor gene has been associated with observed
parenting in healthy parents (Avinun, Ebstein, and Knafo, 2012) as well as the context of continuous
trauma exposure (Feldman, Vengrober, and Ebstein, 2014). OT administration to males and females
increases both AVP (Weisman, Schneiderman, Zagoory-Sharon, and Feldman, 2013), indicating
affinity between the expression of the two neuropeptides.
Only two studies, to our knowledge, measured plasma AVP levels in parents. In the first, OT
and AVP were measured in relation to neural activations in the maternal and paternal brain (Atzil,
Hendler, Zagoory-Sharon, Winetraub, and Feldman, 2012). AVP correlated with fathers’, but not
mothers’, amygdala response to infant stimuli, supporting the links between fathering and AVP in
humans. In the second study, OT and AVP levels in mothers and fathers of 4-month-old infants
were measured in relation to parent-infant interactions. No mean-level differences emerged in AVP
between mothers and fathers, but plasma OT and AVP were associated with distinct configurations
of parental behavior. Parents with higher OT directed their infants toward a social focus, enhancing
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behaviors such as gaze coordination and affectionate contact, behaviors that were more prevalent in
mothers. In contrast, parents with high AVP engaged in stimulatory contact and tended to increase
object-salience when infants showed bids for social engagement, a behavioral profile more common
in fathers. Thus, synchronous processes with mother and father within the family unit distinctly pre-
pare children to join the larger social world (Apter-Levi et al., 2014). (Another study followed parents
and infants across the first 6 years of parenthood in relation to brain activations and assessed salivary
AVP and is described below in the section on hormones and the parental brain.)
Prolactin
Prolactin (PRL) is a peptide hormone originating mainly in the anterior pituitary lactotroph cells
(Freeman, Kanyicska, Lerant, and Nagy, 2000) that has multiple effects on reproduction and lactation
and is thought to mediate the formation of affiliative bonds (Neumann, 2009). PRL is released within
the hypothalamus and other limbic areas during mother-infant contact in rodents (Torner et al.,
2004) and its administration stimulates maternal care in rats (Bridges, DiBiase, Loundes, and Doherty,
1985). PRL has been examined in relation to fatherhood in a number of animal species (Storey,
Delahunty, McKay, Walsh, and Wilhelm, 2006; Wynne-Edwards, 2001; Ziegler, Wegner, Carlson,
Lazaro-Perea, and Snowdon, 2000). In humans, studies have shown that both men and women exhibit
elevated levels of plasma PRL before childbirth, and fathers who report being more affected by infant
cues show higher PRL (Fleming, Corter, Stallings, and Steiner, 2002; Storey et al., 2006). Experienced
fathers show greater increases in PRL when listening to infant cries as compared to first-time fathers
(Fleming et al., 2002). In contrast to single men, fathers’ PRL does not decline following interac-
tion with their toddler (Gray, Parkin, and Samms-Vaughan, 2007). Among fathers to 6-month-old
children, plasma PRL correlates with OT levels and higher paternal PRL is associated with greater
attention to the environment and joint father-child exploratory play (Gordon et al., 2010c).
In mothers, on the second post-birth day, a rise in PRL was found 20 minutes after infant suckling
( Jonas et al., 2009). Infant stimulation of nipple induces both OT and PRL responses (McNeilly,
Robinson, Houston, and Howie, 1983). Finally, between 4–6 weeks postpartum, higher PRL cor-
relates with lower stress and better mood only among formula-feeding mothers (Groër, 2005), and
following cesarean delivery OT and PRL are related to lower anxiety (Nissen, Gustavsson, Wid-
ström, and Uvnäs-Moberg, 1998). Overall, early animal studies on the neurobiology of maternal
care described the contribution of both OT and PRL, but human studies have placed much greater
emphasis on the role of OT, with less research devoted to the links between PRL and observed
parenting.
Testosterone
Testosterone is an androgenic steroid produced by the hypothalamic-pituitary-gonadal (HPG) axis
that modulates reproductive behavior and plays a key role in human social behavior, particularly in
behaviors associated with social status, at times in combination with aggressive behavior (Eiseneg-
ger, Haushofer, and Fehr, 2011; Mazur and Booth, 1998; Wingfield, Hegner, Dufty, and Ball, 1990).
Testosterone’s involvement in parenting and pair bonding has been described in human and other
mammals (Kuzawa, Gettler, Muller, McDade, and Feranil, 2009; van Anders and Goldey, 2010), and
alterations in T levels in males are thought to reflect a shift between conflicting reproductive strate-
gies, from mating efforts to parenting efforts (Gray and Anderson, 2010). Studies in more than 60 bird
species support the “challenge hypothesis”, which suggests that T levels increase when males compete
for food and territory and decrease when males must care for offspring (Wingfield et al., 1990).
Research in biparental species shows that fathers’ T levels decrease in the presence of a dependent
offspring (Wynne-Edwards, 2001). For example, marmoset males who carried infants the most had
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the lowest urinary T levels (Nunes, Fite, Patera, and French, 2001) as well as the greatest declines
in gonadal steroids (Nunes et al., 2001), and exposure to infant scent lowered serum testosterone
in father common marmosets (Prudom et al., 2008). In the monogamous and biparental California
mouse (Peromyscus californicus), greater T-increase during courtship is associated with paternal cud-
dling and a protective repertoire towards their pups (Gleason and Marler, 2010).
Similar findings have emerged in human fathers. High T was found in single and divorced men,
and low T in men and women within a committed relationship as well as in new fathers (Booth
and Dabbs, 1993; Burnham et al., 2003; Gettler, McDade, Feranil, and Kuzawa, 2011; Mazur and
Michalek, 1998; van Anders and Goldey, 2010). During the transition to fatherhood, men decrease
their T levels (Berg and Wynne-Edwards, 2001; Perini, Ditzen, Fischbacher, and Ehlert, 2012), and
such decrease is associated with positive paternal behavior (Fleming et al., 2002). A study in the
Philippines assessing men before and after becoming fathers showed a decline in T levels in fathers,
which correlated with the degree of father involvement in childcare (Gettler, McDade, Agustin,
Feranil, and Kuzawa, 2013). In our study of intranasal OT administration to fathers, we found
that lower baseline paternal T was associated with more optimal father and infant social behavior,
including gaze, vocalizations, and touch (Weisman et al., 2014). Furthermore, OT-induced changes
in T correlate with more positive affect, social gaze, and synchrony, consistent with the perspective
that neuroendocrine systems in human males evolved to support committed and flexible fathering
(Ziegler, 2000).
Very few studies test T in mothers. An increases in T was found in pregnant women (Edelstein
et al., 2015; Fleming et al., 1997), and mothers’ T levels were associated with infants’ physical and
socioemotional health and lower maternal depression ( J. I. Cho, Carlo, Su, and McCormick, 2012;
J. Cho, Su, Phillips, and Holditch-Davis, 2016). We found that across the first months of parenting,
fathers’ T is associated with lower behavioral synchrony, and mothers’ T is not directly related to
maternal behavior (Gordon et al., 2017). However, in the context of high T, maternal OT predicts
greater mother-infant synchrony, further supporting the mutual influences of OT on T, which
require much further research. Assessing diurnal T in mothers and fathers of two preschool-aged
children, among fathers more diurnal variability in T was associated with more sensitivity and respect
for autonomy, whereas for mothers greater diurnal variability correlated with less sensitivity, further
indicating that T carries differential effects on mothering and fathering (Endendijk et al., 2016).
Higher maternal testosterone and infant cortisol are associated with more positive and more frequent
maternal interactive behaviors ( J. Cho, Su, Phillips, and Holditch-Davis, 2015). It thus appears that
the direct and mediated effects of T on parenting, particularly mothering, requires much further
research both in relation to the effects of T on behavior and the effects of T on other hormones.
Cortisol
Cortisol is a steroid hormone secreted by the hypothalamic-pituitary-adrenal (HPA) axis in condi-
tions of physical and psychological stress (Lupien, McEwen, Gunnar, and Heim, 2009). Cortisol is
a key component of the stress response, and as parenting is a highly stressful evolutionarily adaptive
process, research has pointed to CT’s participation in the vigilant component of parenting. A large
body of research in humans and animal models links CT with the regulation of maternal behavior,
and, to a lesser extent, with paternal caregiving (Fleming et al., 1997; Wynne-Edwards, 2001; Ziegler,
2000). Most research on cortisol in the context of parenting is related to maternal stress, and the vast
majority of studies utilize CT as an index of stressed parenting, associated with maternal early or
current life stress.
The various components of the stress response are indexed in parenting research by multiple CT
indices, including basal cortisol, cortisol reactivity to stressful paradigms, diurnal CT production, and
hair cortisol (Levine et al., 2007). Overall, the stress response involves complex interactions between
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the sympathetic nervous system and the HPA axis, allowing to both prepare for danger and return
to baseline once threat is removed (Laurent, Ablow, and Measelle, 2012; Lupien et al., 2009). The
HPA axis comprises the hormones CRH, ACTH, and cortisol, which interact with contextual fac-
tors to shape both momentary stress reactivity and long-term stress physiology (Ellis and Essex, 2007;
Romeo, 2010). Cortisol plays an important role in the stress response by preventing over-reaction
of the immune system to threats and acting on the hypothalamus and pituitary gland via negative
feedback loops to foster homeostasis once safety is achieved (Kudielka, Hellhammer, and Wüst, 2009;
Miller, Chen, and Zhou, 2007; Smyth, Hucklebridge, Thorn, Evans, and Clow, 2013).
Extant evidence in humans and animals has shown that maternal care provides social buffering
of HPA axis activity in offspring (Hostinar, Sullivan, and Gunnar, 2014; Jessop and Turner-Cobb,
2008; Moriceau and Sullivan, 2006; Shionoya, Moriceau, Bradstock, and Sullivan, 2007). Beginning
in infancy, when the child’s HPA system is labile, and across childhood and adolescence, sensi-
tive parenting attenuates children’s HPA reactivity, expressed in smaller cortisol increases or quicker
returns to baseline following stress (Albers et al., 2008; Blair et al., 2008; Berry et al., 2016; Feldman,
Singer, and Zagoory, 2010). In contrast, insensitive parenting, expressed in intrusive, unavailable, and
fragmented parental style, alters the development of children’s stress response and threat-detection
neurobiological circuits (Hostinar et al., 2014) and correlates with higher CT production (Ahnert
et al., 2004; Berry et al., 2016; Bosquet Enlow et al., 2014; Marceau et al., 2015) or inflexible cortisol
response and reduced variability (Apter-Levi et al., 2016). Thus, a central line in the study of CT
relates to how the nature of parental care shapes the development of children’s HPA axis functioning.
Less research on CT in the context of parenting addresses CT in the parent, and some of these
studies measures parental CT in conjunction with child CT. The bulk of this research focuses on
high-risk conditions (see below), with less research assessing parental CT in low-risk samples. Thus,
from the extant literature and reviews available on the cross-generational transmission of human
stress physiology (for review see Bowers and Yehuda, 2016) less research has focused on CT in rela-
tion to observed parental behavior in low-risk mothers, and even less research has tested paternal CT
in relation to paternal behavior in typically developing families.
Regarding CT and mothering, in the newborn period, reports are mixed on the relations of CT
to maternal behavior. Some found the expression of maternal behavior to correlate with higher CT
(Fleming, Steiner, and Anderson, 1987), but our assessment of plasma CT across pregnancy and the
postpartum month showed that CT increased in late pregnancy and higher CT predicted restricted
maternal behavior (Feldman et al., 2007). Starting at 3–4 months, research has measured CT in
mothers and infants in stressful paradigms, such as the “still-face”, and most studies across infancy,
childhood, and adolescence describe links between higher maternal CT, expressed in higher basal
levels, greater stress reactivity, and slower recovery from stress, and less optimal parenting, expressed
in lower sensitivity, decrease reciprocity, and greater intrusiveness (for review, Gunnar, Talge, and
Herrera, 2009).
Although most studies on CT and stress-inducing paradigms focus on infant CT, few also test
maternal CT. For instance, higher maternal basal cortisol and greater reactivity to the “still-face” at 6
months are related to higher intrusiveness and lower second-by-second synchrony (Feldman, Singer,
and Zagoory, 2010). Similarly, mothers with higher parenting-focused mindfulness show steeper
cortisol recovery slopes following the still-face at 6 months (Laurent, Duncan, Lightcap, and Khan,
2017). Maternal blunted cortisol awakening response (CAR), the typical increase in CT from wak-
ening to 30 minutes post-wakening, during pregnancy predicts lower infant emotion regulation as
mediated by maternal sensitivity (Thomas, Letourneau, Campbell, Tomfohr-Madsen, and Giesbre-
cht, 2017). Similarly, higher diurnal cortisol production is linked with maternal retrospective report
of early life stress and predicts lower sensitivity to their 2–6 months (Gonzalez, Jenkins, Steiner, and
Fleming, 2012). Of note, among mothers and infants aged 6–12 months, those of low socioeconomic
status (SES) had higher diurnal CT production compared to high-SES mothers and infants, as well
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as lower adrenocortical synchrony (Clearfield, Carter-Rodriguez, Merali, and Shober, 2014). Overall,
these studies indicate that across CT biomarkers more attuned parenting and greater maternal abili-
ties to allocate resources to the child are associated with lower CT.
In fathers, CT declines following father-toddler interaction, as does fathers’ PRL, and the decline
in CT is greater in experienced, compared to first-time, fathers (Gettler, McDade, Agustin, and
Kuzawa, 2011). Testing fathers of 22-month-old toddlers following father-child interactions on a
day they spent several hours alone with the child prior to testing versus days without the child, it was
found that CT generally declined following interaction, but a greater decline was observed when
fathers spent time alone with the child (Storey, Noseworthy, Delahunty, Halfyard, and McKay, 2011).
We found greater diurnal CT production in mothers of 6-month-old infants compared to fathers
and both parents’ CT was negatively related to warmth and sensitivity during triadic mother-father-
infant interaction (Gordon et al., 2010a). These studies highlight the stress-reducing function of
positive father-child interaction on the father’s overall cortisol production and CT response.
The use of hair cortisol analysis in humans provides a measure of more chronic aspects of the
stress response (Burnard, Ralph, Hynd, Edwards, and Tilbrook, 2016; Russell, Koren, Rieder, and
Van Uum, 2012; Stalder and Kirschbaum, 2012; Staufenbiel, Penninx, Spijker, Elzinga, and van Ros-
sum, 2013). Each centimeter of hair approximates one month of cortisol secretion, thus measuring
CT in hair presumably integrates free steroids over the time of growth (Russell et al., 2012; Stalder
et al., 2016), and thus hair cortisol concentrations (HCC) are thought to provide a retrospective
month-by-month measure of cumulative cortisol secretion and serve as a reliable biomarker of
chronic stress (Hinkelmann et al., 2013; Ouellette et al., 2015; Simmons et al., 2016; Steudte et al.,
2013; Vanaelst et al., 2012).
HCC has been studied mainly in the context of chronic stress, trauma, and psychiatric illness, and
very few studies have integrated this measure into parenting research. In children, HCC is associ-
ated with lifetime trauma (Simmons et al., 2016), fearfulness upon school entry (Groeneveld et al.,
2013), the number of major childhood traumatic life events (Vanaelst et al., 2012), and lower SES,
which likely involves greater chronic stress (Rippe et al., 2016; Vliegenthart et al., 2016). Higher
parenting stress and greater child socioemotional difficulties are linked with children’s elevated HCC
(Palmer et al., 2013). Similarly, mild perinatal adversity, such as late preterm birth, moderates the
links between maternal harsh parenting and HCC in 6-year-old children (Windhorst et al., 2017).
These studies suggest that, if little research has integrated hair measurement in parents, HCC may
be a unique biomarker of the stress response that requires much further research in the context of
low- and high-risk parenting.
Salivary Alpha Amylase

Salivary alpha amylase (sAA) has been integrated into research on parenting and child outcomes
an index of the sympathetic-adrenal-medullary (SAM) arm of the stress response (Hellhammer,
Wüst, and Kudielka, 2009; Nater and Rohleder, 2009). The stress response involves the coordinated
functioning of two major anatomically distinct systems, the SAM, which initiates the fight-or-flight
response by increasing blood flow, respiration, cardiovascular activity, and the release of catechola-
mines (Nater and Rohleder, 2009), and the HPA system, which has a more gradual onset and is
associated with physiological and behavioral withdrawal (Bauer, Quas, and Boyce, 2002; Tarullo and
Gunnar, 2006). Whereas momentary stress induces immediate changes in each system, chronic stress
exerts a lasting effect and may alter the balance between the functioning of the SAM and HPA sys-
tems (Wolf, Nicholls, and Chen, 2008). Salivary alpha amylase has mainly been tested in children and
less commonly in parents in the context of stress, both alone and in relation to CT (Wolf et al., 2008).
In children, sAA has been tested in relation to physical health (Wolf et al., 2008), negative emo-
tional reactivity (Spinrad et al., 2009), and attachment under stress (Frigerio et al., 2009). Similar to
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CT, the bulk of sAA studies have been conducted in high-risk samples (see below). Lower sAA was
found among maltreating mothers and reduced reactivity to infant crying compared to nonmal-
treating mothers (Reijman et al., 2015). Insecure-avoidant 1-year-old infants had higher sAA levels
following the strange situation paradigm and their mothers, who showed no differences in sAA,
exhibited less vagal withdrawal in the reunion episode (Hill-Soderlund et al., 2008). Parent-child
relationship quality predicts the associations between marital conflict and higher child sAA reactiv-
ity (Lucas-Thompson and Granger, 2014), and smoking mothers have higher salivary CT and lower
sAA compared to nonsmoking mothers (Granger et al., 2007). In fathers of adolescent girls, higher
interparental aggression is related to lower father sAA (Gordis, Margolin, Spies, Susman, and Granger,
2010). Overall, it appears that sAA may be a useful marker of the SAM arm of the stress response,
but much further clarifications are required to integrate it as a measure of optimal versus high-risk
parenting.
Summary
As seen, the hormones of parenting in humans function in a comparable way to those supporting
parental care in nonhuman mammals and enable the expression of the unique evolution of parental
care across human societies. OT, AVP, and PRL appear to ignite the expression of parenting behavior,
CT, sAA, and immune biomarkers to manage the stress involve in parenting, and T plays a special
role in development of fathering.
Endocrine Fit: Synchrony in Parent and Child’s Hormones

Synchrony or attunement between the parent and infant’s physiological and behavioral processes
enables mammalian parents to promote sociality and regulate stress in their young. Hormonal con-
cordance or synchrony, the match between parent and child’s hormones, is a central aspect of such
biobehavioral attunement and a link between parent and child’s hormonal levels has been observed
in most studies reporting on the associations between parent and child’s hormones measured concur-
rently. Hormonal synchrony is thought to stem from both genetic similarity and shared environment;
yet research has tested the degree in such linkage in different contexts and conditions and its associa-
tion to parent-child relational variables. According to our biobehavioral synchrony model (Feldman,
2012c, 2015b, 2016, 2017), biological synchrony is an important mechanism in the development of
mammalian young by which the parent’s mature physiological systems externally regulate the infant’s
immature system through the coordination of biological and behavioral processes during moments
of social contact. We showed in multiple physiological systems, such as heart rate coupling (Feldman,
Magori-Cohen, Galili, Singer, and Louzoun, 2011), brain-to-brain synchrony in neural oscillations
(Levy et al., 2017), and endocrine systems (e.g., Feldman et al., 2010; Pratt et al., 2017), that syn-
chrony in biological processes is anchored in behavioral synchrony and increases during moments of
concordance in parent and child nonverbal behavior in the gaze, affect, vocal, and touch modalities.
Consistent with findings in animal models (Hofer, 1995a), we found that biological synchrony
operates in a system-specific manner. Thus, the following reviews the two main parenting-related
hormonal systems, oxytocin and cortisol, in healthy and high-risk populations, with more empiri-
cal data available for CT compared to OT. The distinction between the two hormones as the main
neuroendocrine systems supporting the affiliation and stress/vigilance components of parenting is
also expressed in a distinction between the developmental goal of hormonal synchrony in OT and
CT. For OT, higher parental sensitivity, synchrony, and reciprocity are associated not only with
higher parental OT and higher infant OT, but in a closer match between their OT levels, which pro-
motes more optimal social-emotional outcomes in children. In contrast, tighter cortisol synchrony
232
is associated with greater child stress physiology and lower parental sensitivity and dyadic reciprocity
(Pratt et al., 2017).
Parent-Child Oxytocin Synchrony

Among 4- to 6-month-old infants, we found hormonal synchrony of oxytocin both prior to and
following a “play and touch” paradigm, and such endocrine synchrony was observed when social
synchrony was high, not when it was low (Feldman et al., 2010). The endocrine fit of parent and
child’s OT only in cases of high behavioral synchrony suggests that the fit between parent and child
is based on parental behavior, consistent with research in animal models.
In three high-risk samples we found OT synchrony between parent and child. In a study com-
paring preschoolers with autism spectrum disorders (ASD) with typically developing children (TD),
OT synchrony in both the ASD and TD group emerged between children with both their moth-
ers and fathers, with no significant differences in the magnitude of OT synchrony between parents
or among the two groups (Feldman, Golan, Hirschler-Guttenberg, Ostfeld-Etzion, and Zagoory-
Sharon, 2014), despite the fact that levels of OT differed between ASD and TD preschoolers but not
among their mothers or fathers (see below).
Following mothers diagnosed with Axis-I depression across the child’s first 6 years of life and their
children, we found that depressed mothers had lower salivary OT as did their children. Fathers in
families of depressed mothers also had lower OT, and low OT was related to the diminished mater-
nal touch and social gaze in depressed mothers (Apter-Levy, Feldman, Vakart, Ebstein, and Feldman,
2013). Lower baseline OT and attenuated OT response to mother-child interaction was also found
in urinary OT in mothers and children whose urinary OT levels were correlated. As stated above,
such urinary OT concordance was sensitive to stressful aspects of the interaction and correlated
with greater maternal intrusiveness and higher child withdrawal. Of note, among depressed mothers,
those who still had higher OT were able to transmit a functional OT system to their child, and their
children’s OT differed from that of controls, highlighting the protective role of the mother’s oxytocin
functionality (Pratt et al., 2015).
Finally, in a group of children exposed to continuous war-related trauma, we found lower OT in
war-exposed mothers and OT synchrony between mother and child. Such OT synchrony mediated
the effects of war on the child so that high maternal OT led to higher mother-child behavioral syn-
chrony leading to reduced child’s anxiety disorders by age 10 (Ulmer-Yaniv et al., 2017).
Parent-Child Adrenocortical Synchrony

The coordination between parent and child cortisol production has been entitled by several terms,
including cortisol coregulation, hormonal concordance, stress contagion, or adrenocortical synchrony
(Atkinson et al., 2013; Mörelius, Örtenstrand, Theodorsson, and Frostell, 2015; Papp, Pendry, and
Adam, 2009; Pratt et al., 2017; Ruttle, Serbin, Stack, Schwartzman, and Shirtcliff, 2011; Saxbe et al.,
2014; Stenius et al., 2008). The vast majority of studies examined the coordination of CT following
stress paradigms and found that, when stress is elevated in mother or child, both partners increase CT
in a coordinated fashion (Atkinson et al., 2013; Hibel, Granger, Blair, and Finegood, 2015; Mörelius,
Broström, Westrup, Sarman, and Örtenstrand, 2012; Mörelius, Theodorsson, and Nelson, 2009; Neu,
Laudenslager, and Robinson, 2009; Ruttle et al., 2011; Sethre-Hofstad, Stansbury, and Rice, 2002).
Much less research has focused on the coordination of diurnal CT patterns between mother and
child (Hibel, Trumbell, and Mercado, 2014; LeMoult, Chen, Foland-Ross, Burley, and Gotlib, 2015;
Papp et al., 2009; Schreiber et al., 2006; Stenius et al., 2008; Williams et al., 2013), a distinct aspect of
HPA axis functioning that is often uncorrelated with CT reactivity to momentary stressors (Edwards,
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Ruth Feldman
Clow, Evans, and Hucklebridge, 2001). Even fewer measured parent-child linkages between hair
cortisol concentrations in mother and child have been examined, yet another dimension of HPA
reactivity, typically unrelated to salivary response to stress (Halevi et al., 2017).
Synchrony of diurnal CT within the family is related to the amount of shared experience (Möre-
lius et al., 2012, 2015; Schreiber et al., 2006; Stenius et al., 2008). For instance, preterm infants placed
in family care and exposed to maternal-infant skin-to-skin contact exhibit cortisol concordance,
whereas no correlations between maternal and infant CT was found among infants placed in stand-
ard incubator care (Mörelius et al., 2012, 2015). Six-month-old infants show greater diurnal adreno-
cortical synchrony with their mothers as compared to their fathers (Stenius et al., 2008). Among
preschool-aged children mother-child morning CT levels show linkage only on nonwork days
(Hibel et al., 2014); and among adolescents, shared environment is a better predictor of afternoon
CT linkage than genetic factors (Schreiber et al., 2006).
Synchrony in diurnal CT was found between mothers and children, above and beyond time
of measurement. Mother-child reciprocity is related to lower adrenocortical synchrony, whereas
father-child tension is marginally predictive of greater adrenocortical synchrony. Higher child diur-
nal CT production predicts a stronger linkage between maternal and child diurnal CT, suggesting
that greater physiological stress may render children more susceptible to the effects of maternal stress
physiology. Maternal depression, although related to attenuated child diurnal CT decline, does not
affect adrenocortical synchrony. Adrenocortical synchrony may tap a unique aspect of HPA axis
functioning, potentially linked with the cross-generation transfer of stress physiology. Results high-
light mothering and fathering family subsystems as moderators of adrenocortical synchrony and
point to the role of parent-child relational stress in shaping diurnal CT linkage.
Compared to a healthy control group, synchrony in diurnal CT was found between depressed
mothers and children, and the degree of mother-child reciprocity was related to lower adrenocorti-
cal synchrony. When children’s CT production during the day was higher, there was also a tighter
synchrony between maternal and child CT, suggesting that greater physiological stress may render
children more susceptible to the effects of maternal stress physiology. Maternal depression, although
related to attenuated child diurnal CT decline, did not affect adrenocortical synchrony. These find-
ings highlight the role of parent-child reciprocity in shaping diurnal CT linkage (Pratt et al., 2017).
In a group of preschoolers with ASD compared to TD children, we found CT synchrony between
children and both their mothers and their fathers at the three measurement points following the SF
paradigm (Ostfeld-Etzion, Golan, Hirschler-Guttenberg, Zagoory-Sharon, and Feldman, 2015). Fur-
thermore, father-child cortisol linkage is stronger in dyads that show less reciprocity, when fathers
were less sensitive and when children showed less self-regulation. Consistent with the prior findings,
mother-child linkage is stronger in dyads that show less reciprocity and lower maternal sensitivity,
demonstrating that higher CT linkage is observed in less functional dyads (Saxbe et al., 2017).
Finally, in the war-exposed group, early childhood adrenocortical synchrony is present in mater-
nal and child CT levels at both baseline and reactivity to stressors in early childhood (Feldman,
Vengrober, Eidelman-Rothman, and Zagoory-Sharon, 2013). In late childhood (9–11) years, adren-
ocortical synchrony appears in both salivary cortisol and hair cortisol concentrations, and mothers’
reduced CT in the face of chronic trauma initiates a cascade of biobehavioral synchrony, linking to
lower child CT, greater behavioral synchrony, and higher child social engagement, which, in turn,
decreased child externalizing and internalizing symptoms (Halevi et al., 2017).
Overall, adrenocortical synchrony is thought to be a mechanism by which, beginning in utero,
mothers signal to the developing fetus the amount of danger the environment will likely contain.
Studies in rodents indicate that the mother’s species-typical behavior carries a unique effect on con-
solidation of the pup’s HPA reactivity (Gubernick and Alberts, 1983; Rosenberg, Denenberg, and
Zarrow, 1970) and that mothers with lower corticosterone display more maternal behavior and their
infants show lower HPA axis reactivity in adulthood (Francis and Meaney, 1999; Dong Liu et al.,
234
1997). Cross-fostering studies show that maternal behavior has epigenetic effects on pup neural and
behavioral responses to stress and the effects of maternal behavior exceed those of genetic disposi-
tions (Champagne and Meaney, 2001; Kundakovic and Champagne, 2015). These findings provide
mechanistic evidence for the concordance between maternal and child HPA axis functioning and
suggest that variability in maternal caregiving may play a role in shaping the infant’s cortisol produc-
tion and degree of their adrenocortical synchrony (Macrì, Zoratto, and Laviola, 2011).
Parents’ Hormones and the Parental Brain
Hormonal Correlates of Parent Brain Activations

Research in rodents has shown that hormones of pregnancy prepare brain structures which are
sensitized by childbirth and form the “mammalian caregiving network”, including the amygdala,
hypothalamus (particularly the MPOA), and the dopamine-rich subcortical ventral tegmental area
(VTA) (for review; Numan and Stolzenberg, 2009; Numan and Young, 2016). Imaging studies of the
human parental brain, exposing parents to auditory, visual, or multimodal stimuli of their infants, have
revealed that the same network activates, in addition to other cortical networks implicated in empa-
thy, interoception, embodied simulation, mentalizing, and emotion regulation to form the “global
human caregiving network” (for Review, Feldman, 2015b; Swain and Ho, 2017).
Work on the parental brain points to associations between parent’s brain activations and parents’
hormones. Regarding maternal brain-hormone relations, mothers’ plasma OT levels correlate with
two nodes of the subcortical mammalian network; amygdala, mediating maternal vigilance, and
Nucleus accumbens, linked with the subcortical dopamine reward system (Atzil et al., 2011). Salivary
OT in mothers relates to maternal dorsal anterior cingulate cortex (dACC), a component of the
empathy-embodied simulation network (Abraham et al., 2014), and to the hypothalamus and ven-
tral tegmental area of the subcortical mammalian network (Strathearn et al., 2009). Finally, mothers
showing less CT reactivity have higher brain activation to their infant cry in the PAG, insula, ACC,
and OFC, areas implicated in interoception (perception of bodily milieu) and empathy (Laurent,
Stevens, and Ablow, 2011).
Fathers’ OT is associated with activations in the superior temporal sulcus, a key node of the social
brain integrating mirror and mentalizing properties (Abraham et al., 2014). Fathers’ amygdala activ-
ity correlates with fathers’ plasma AVP levels (Atzil et al., 2012). Finally, fathers’ testosterone, known
to decrease in men at the transition to fatherhood (Gettler, McDade, Feranil, et al., 2011), correlates
with lower VTA activation and higher left caudate activation (Kuo, Carp, Light, and Grewen, 2012;
Mascaro, Hackett, and Rilling, 2013).
Long-Term Prediction Of Parental Brain and Hormones

for Children’s Social Development
In several studies, we measured parental neural and hormonal response in infancy in relation to child
outcomes across the first years of life. Among primary-caregiving mothers and fathers, we found
that the coherence of the parent’s embodied simulation network, integrating structures implicating
in mirror and empathy functions, and parental OT predicted children’s OT in the preschool stage as
well as their capacity to use advanced strategies for regulating negative emotions (Abraham, Hendler,
Zagoory-Sharon, and Feldman, 2016).
Another study assessed parent brain response to coparental stimuli—stimuli depicting the partner
as parent, in relation to observed coparental behavior, hormones, and child outcomes. Coparental
stimuli activated the caudate, a critical node in supporting motivational goal-directed social behavior.
Caudate-ventromedial prefrontal cortex (vmPFC) vmPFC connectivity, linking caudate with the
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prefrontal area implicated in intersubjectivity, mentalization, and affect sharing, is associated with
collaborative coparenting and the link between caudate-vmPFC connectivity and reduced child
behavior problem at 6 years was mediated by the parent’s OT. Caudate connectivity with the dACC,
which has been linked with pain perception, envy, and vigilant monitoring of social and aggressive
response, predicts undermining coparenting across time and is linked with AVP (Abraham, Gilam,
et al., 2017). Greater functional connectivity between the two empathy networks in the parental
brain, the embodied simulation and mentalizing networks, predicts lower child CT reactivity and
better emotion regulation at preschool (Abraham, Raz, Zagoory-Sharon, and Feldman, 2018).
Parental Hormones in High-Risk Conditions

A comprehensive review of parental hormones in high-risk populations is beyond the scope of a sin-
gle chapter. The following reviews studies on OT and CT in high-risk populations, focusing mainly
on two systems—OT and the affiliative system and CT and indices of the stress response.
Oxytocin and Affiliative Biomarkers Markers
Postpartum Depression
Several studies have addressed OT functionality in mothers suffering from depression, with few stud-
ies addressing mothers with clinically diagnosed Axis-I depression, not just self-reported depressive
symptoms. Lower OT during pregnancy and the postpartum goes with greater depressive symp-
tomatology in the neonatal period and less maternal behavior (Feldman et al., 2007). Similar findings
were reported by Skrundz, Bolten, Nast, Hellhammer, and Meinlschmidt (2011), who showed that
higher depressive symptoms during pregnancy predict lower plasma OT in the postpartum.
A longitudinal study followed a community cohort of depressed mothers and their families from
birth across the first decade of life. At 6 years, depressed mothers, their husbands, and their children had
lower salivary OT levels and greater prevalence of the more evolutionary-recent protective A allele on
the OXTR. Lower OT was linked with reduced maternal touch and sensitive parenting (Apter-Levy
et al., 2013). Measuring OT in urine in mother and child showed that in both depressed mothers and
their children there was lower baseline OT and lower OT response to mother-child interactions. Such
reduced urinary OT was related to higher maternal intrusiveness and child withdrawal (Pratt et al.,
2015). At 10 years, depressed mothers and children as a group no longer had lower salivary OT, but
child OT mediated links between maternal depression and child externalizing and internalizing symp-
toms as well as child lower empathy as measured by two home-based paradigms (Priel et al., 2018).
Administration of OT to postnatally depressed mothers did not increase the level of sensitive
parenting (Mah, van IJzendoorn, Smith, and Bakermans-Kranenburg, 2013) nor improve depressive
symptoms, but increased their protective behavior to infants in the presence of an intrusive stranger
(Mah, Bakermans-Kranenburg, van IJzendoorn, and Smith, 2015).
With regards to other hormones of the affiliation system, plasma prolactin levels are significantly
lower in depressed mothers who were breastfeeding (Harris et al., 1989). Similarly, depressed mothers
show lower serum prolactin levels (Groer and Morgan, 2007). To date, no study has tested maternal
AVP in the context of depression.
Stress and Trauma

Following a cohort of children exposed to repeated wartime trauma and their mothers from early
childhood to adolescence, we found that by 9–11 years war-exposed mothers had lower OT, but
children as a group did not have lower OT, only those with PTSD. These mothers also had much
236
higher prevalence of psychiatric disorders, particularly anxiety disorders, PTSD, and depression, their
children had higher levels of anxiety symptoms as well as greater prevalence of psychiatric diagnosis.
War-exposed mothers also exhibited lower sensitivity and empathy, and their children displayed less
social engagement, which was related to lower maternal OT (Ulmer-Yaniv et al., 2017). Children
with ASD exhibit lower baseline OT levels, which are momentary normalized during parent-child
contact (Feldman et al., 2014). There is evidence that OT increases during skin-to-skin contact
between parents and premature infants (Cong et al., 2015), findings which are consistent with the
links between maternal proximity and licking-and-grooming with the oxytocin system in animal
models.
Cortisol and Stress Biomarkers

Within the family of stress-related biomarkers, numerous studies assess diurnal or reactive cortisol as
well as other stress biomarkers in relation to high-risk parenting. Most studies on CT and high-risk
parenting address the effects of maternal stress, trauma, depression, or premature birth on the infant’s
CT, but few studies also measure maternal hormones. Importantly, there are studies following chil-
dren from birth to adolescence demonstrating that maternal postpartum depression or perinatal stress
alter various aspects of children’s HPA axis functioning including baseline levels, diurnal patterns, and
variability (e.g., Halligan, Herbert, Goodyer, and Murray, 2007).
Postpartum Depression
During pregnancy and 3 months postpartum, higher depressive symptoms are associated with lower
cortisol awakening response and flatter diurnal patterns (Scheyer and Urizar, 2016). In one study, at
8 weeks postpartum, breastfeeding mothers underwent a social stressor while breastfeeding. Among
depressed mothers, the surge of OT during nursing was reduced and CT levels were higher, suggest-
ing that depression attenuates the anxiolytic effects of breastfeeding on the maternal stress response
(Cox et al., 2015). Mothers with a history of major depression combined with child abuse showed
steeper CT decline and their infants had lower baseline CT, and more maternal comorbid condi-
tions on top of the depression, such as abuse history or PTSD, augmented the disruptions to HPA
functioning (Brand et al., 2010). At 4–6 weeks postpartum, depressed mothers show downregulated
HPA functioning, expressed as lower salivary CT (Groer and Morgan, 2007). In contrast, both clini-
cally depressed and clinically anxious mothers at 9 months have higher CT, augmented CT response
to stress, and slower CT recovery, and these alterations in CT production were associated with their
diminished sensitivity and lower infant social engagement (Feldman et al., 2009).
Following the cohort of depressed mothers and their children from birth to 10 years, we found
that at 6 years, mothers and children did not have altered cortisol levels—both diurnal and reactive,
but had diminished CT variability in daily patterns and in response to stress (Apter-Levi et al., 2016;
Pratt et al., 2017). Similar findings emerged at 10 years of age (Priel et al., 2018), with less flexible CT
patterns (lower AUCi index) at both ages associated with less optimal mothering, including maternal
intrusiveness and diminished sensitivity, and greater child social withdrawal.
Trauma
Mothers with a history of early trauma show less positive affect and flatter cortisol patterns during a
home visit at 6 months ( Juul et al., 2016). In a longitudinal study of children exposed to war-related
trauma, we found in early childhood that, compared to nonexposed controls, children exposed to
trauma since birth had reduced CT variability in response to stress. However, exposed children with
PTSD had low and flat CT patterns, suggesting a “shut down” response, but exposed children who
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Ruth Feldman
were more resilient had elevated nonflexible levels, indicating high arousal of the system. These dif-
ferential patterns were related to differences in maternal depression, anxiety, and PTSD symptoms,
which were higher in mothers of PTSD children, and greater proximity-seeking behavior in the
more resilient exposed child group. Mothers and children also manifest altered patterns of salivary
alpha amylase, and for both CT and sAA there were close links between maternal and child’s hormo-
nal levels (Feldman, Vengrober, et al., 2013). These findings—which differentiate trauma survivors or
trauma-exposed individuals with and without PTSD—are consistent across several samples and vari-
ous traumas, such as survivors of the 9/11 attack, the Holocaust, and abuse (Yehuda and Bierer, 2007).
At 10 years of age, we measured both hair cortisol and salivary CT response from mothers and
children in the same sample. Mothers who lived in a war zone for over a decade had higher hair CT,
indicating greater chronic stress, as well as greater salivary CT production during a home visit. These
altered maternal patterns impacted the child’s pattern via mechanisms of cortisol linkage, charting a
pathway from trauma exposure to higher psychopathology in children (Halevi et al., 2017).
Autism Spectrum Disorders

Mothering children with ASD involves high levels of stress, yet few studies have assessed maternal
CT in the context of ASD, and even fewer compared CT levels with observed parenting. Moth-
ers and fathers of children with ASD have lower morning cortisol levels, indicating effects of the
increased stress on daily stress response (Foody, James, and Leader, 2015). Similarly, 89% of mothers
of ASD children display a blunted diurnal CT response, indicating decreased flexibility of the system
(Dykens and Lambert, 2013), and another study reported lower CT production throughout the day
in mothers of adolescents with ASD (Seltzer et al., 2010).
We assessed CT production in 3- to 6-year-old children during a home visit with mother and a
parallel home visit with father, where they faced the same experimental stress manipulations (emo-
tion regulation tasks and “still face”). We found no differences between the CT response of moth-
ers and fathers to children with ASD as compared to control parents; however, children with ASD
showed blunted CT responses during interaction with mothers, but typical responses during the
visit with father. We interpreted the findings in terms of fathers’ pushing children to act more in
age-appropriate ways and mothers providing social buffering to children’s stress response in a similar
manner to mammalian neonates (Ostfeld-Etzion et al., 2015).
Prematurity
Kangaroo care (KC), or skin-to-skin contact, is an intervention aimed to reduce maternal-newborn
separation and enhance contact among infants born preterm. Skin-to-skin contact decreases mater-
nal CT following premature birth ( Janevski, Vujičić, and Đukić, 2016). Another study showed
reduction of infant CT in a group receiving kangaroo care and linkage between maternal and infant
CT at 4 months only in the KC group (Mörelius et al., 2015). Reduced CT levels across the first
month following birth were also observed in a full-term sample (Bigelow, Power, MacLellan-Peters,
Alex, and McDonald, 2012). In our study of kangaroo care and its long-term impact, by 10 years of
age children who received kangaroo care as neonates had attenuated CT responses to social stress-
ors (the TSST-C) and their mothers similarly had lower CT production (Feldman, Rosenthal, and
Eidelman, 2014).
Hormones of parenting provide biomarkers for stressed or high-risk parenting, hormonal lev-
els are associated with the parent psychological state, history, psychiatric condition, and observed
behavior, and hormones demonstrate the utility of using neuroendocrine measures to expand our
understanding of at-risk parenting.
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Conclusions
Individual variations in hormones play a key role in the development of parenting and are meaning-
fully associated with variations in maternal and paternal behavior. There is much we do not know
which requires future research. First, there are currently no normed curves for hormones across
pregnancy and the first year following childbirth. With the growing incorporation of hormones
into parenting research, there is a critical need for large-scale studies that can define normative
curves across multiple cultures for future research. Second, changing role of fathers and the growing
involvement of fathers in childcare prompts much research to understand the neuroendocrinological
basis of fatherhood. Third, much research is needed to compare hormonal profiles across a variety of
high-risk conditions and to tease apart single from multiple risk, for instance maternal depression in
the context of low-risk environment, from maternal depression occurring in the context of poverty,
premature birth, or child abuse. Finally, much more research and theory-building are needed to test
the hormones of parenting within a global bio-psycho-social theory of parenting that investigates
endocrine systems from a comparative perspective and across levels of analysis, incorporating studies
of the cellular, genetic, and neural levels with behavioral and representational levels, into a theoretical
frame that can define more precisely how hormones of parenting contribute to the successful rearing
of human children.
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7
NEUROBIOLOGY OF HUMAN
PARENTING
Eloise A. Stark, Alan Stein, Katherine S. Young,
Christine E. Parsons, and Morten L. Kringelbach
Introduction
Human infants need prolonged, intensive caregiving to survive and thrive. Human infants’ depend-
ency is unlike many other newborn animals that are mobile shortly after birth, allowing them a much
larger degree of independence from their parents. Early experiences have a lasting effect on our later
cognitive and emotional development (Shonkoff, Boyce, and McEwen, 2009). Parenting our young
represents one of our greatest social and practical challenges, helping to strengthen lifelong learn-
ing and facilitating our advanced cognitive flexibility (Bornstein et al., 2010; Konner, 2010). The
pleasures of social interaction with infants are arguably some of our strongest motivators (Parsons,
Young, Murray, Stein, and Kringelbach, 2010). As such the parent-infant relationship reflects a bio-
logical necessity, ultimately to ensure the survival of the species (Darwin, 1872). Indeed, the systems
motivating parental behavior appear to be largely conserved across mammalian species (Numan and
Insel, 2003).
Here we review the emerging evidence for the way that underlying human brain systems support
the parent-infant relationship. Starting with the earliest elements of the parent-infant relationship,
orienting to infant cues, we demonstrate how the human brain is optimized to perceive cues across
multiple sensory modalities rapidly and efficiently. This “parental instinct” in response to infant cues
appears to be present for nonparents as well as parents, demonstrating how our parental capacities are
well substantiated in the brain prior to parenthood. We also show how the expertise gained through
parenting experience changes the structural and functional properties of the brain in ways that may
optimize future caregiving.
An infant possesses a large range of communicative cues prior to becoming verbal. From the cry
of a distressed infant, their irresistible smell, to the cuteness of a happy smiling face, infants’ emotional
expressions overwhelmingly draw our attention. Responding to infant communicative cues in a
prompt and sensitive manner is a key parenting capacity (Ainsworth, Bell, and Stayton, 1974). A large
body of behavioral and neuroimaging research across species has identified core brain networks that
regulate parenting behavior in mammals (Barrett and Fleming, 2011; Swain, Lorberbaum, Kose,
and Strathearn, 2007). However, studying the human parental brain presents a specific challenge, as
numerous features are unique to human infancy, one example being the prolonged dependency on
parents or caregivers (Konner, 2010). Direct cross-species comparisons are therefore difficult, despite
a wealth of knowledge of maternal behavior and neurobiology in other mammals (Fleming, 2006).
It is known that the quality of caregiving has a profound impact on child development (Bornstein,
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Neurobiology of Human Parenting
Hahn, and Haynes, 2011; Feldman, 2015; Parsons et al., 2010; Parsons, Young, Rochat, Kringelbach,
and Stein, 2012). Furthermore, the sensitivity of caregiver responses to infant cues can have a lasting
effect on a child’s cognitive and socioemotional development (Stein et al., 2013; Stein et al., 2014a).
We have previously described a behavioral framework to characterize the parent-infant relation-
ship in terms of early interactions (Parsons et al., 2010). As the postnatal period advances, interac-
tions between a parent and infant become increasingly intricate and refined. What begins as a simple
orienting to infant cues may culminate in prolonged and complex interactions such as play or con-
versation. Our behavioral framework covers the first 18 months of an infant’s life, and describes six
major components of the parent-infant relationship: (1) orienting system; (2) recognition system; (3)
intuitive parenting; (4) attachment relationships; (5) intersubjectivity; and (6) higher socioemotional
and cognitive functions.
The orienting system is the first interface between caregiver and infant, with early interactions
characterized by an immediate propensity for each partner to seek contact with one another. Ori-
enting to one another serves to instigate close proximity, thereby facilitating subsequent interaction.
Parents demonstrate a basic attraction to infant cues, which helps secure parental attentiveness, for
example, the allure of an infant’s smell, the magnetism of “cute” facial features, and “auditory cute-
ness” such as infant laughter and babbles (Darwin, 1872; Kringelbach, Stark, Alexander, Bornstein,
and Stein, 2016). These multimodal infant cues serve to orient adults to infants rapidly. Infants show
a remarkable predisposition to orient to fellow humans. Newborn infants show a readiness to track
face-like patterns with their gaze, but not matched nonface forms ( Johnson, Dziurawiec, Ellis, and
Morton, 1991) despite having minimal perceptual experience of faces. Newborns also prefer speech
compared to nonspeech sounds matched for pitch and intensity (Vouloumanos and Werker, 2004)
which helps them to orient to adults within their proximity.
A more selective recognition system supersedes the general orienting response. By learning to rec-
ognize each other, parents and their infants are able to actively pursue interpersonal contact, and this
process therefore happens rapidly following birth. Mothers can accurately recognize their own infant
early in the postpartum on the basis of single nonvisual cues, such as smell, cry, or touch (Cismaresco
and Montagner, 1990; Kaitz, Lapidot, Bronner, and Eidelman, 1992; Porter, Cernoch, and McLaugh-
lin, 1983; Russell, Mendelson, and Peeke, 1983). Within the first few days and weeks of life, infants
demonstrate preference for their mother’s face (Bushnell, 2001), voice (DeCasper and Fifer, 1980),
and even breast milk smell (Macfarlane, 1975). It is therefore crucial to view the infant as a dynamic
partner in the parent-infant relationship, with a wide array of communicative cues, and growing
social perceptual capacities. As part of this developmental trajectory, by about 6 weeks old, infants
show a remarkable sensitivity to the qualities of adult communication (Brazelton, Koslowski, and
Main, 1974; Papousek and Papousek, 1975). They actively pursue social interaction with caregivers,
and react in striking ways if such interactions are not forthcoming (Cohn and Tronick, 1983; Field
et al., 1988). Therefore, the reciprocal recognition system functions to ensure prolonged and intricate
parent and infant interaction.
The third element in our behavioral framework is intuitive parenting. The processes underlying
parental orienting and recognition have been conceptualized as instinctive or intuitive, forming a
distinct class of social behavior (Papousek, 2000; Papousek and Papousek, 1987). These behaviors
include altering speech, establishing eye contact, and mirroring infant expressions. They are argued
to occur largely in the absence of conscious intent, and are therefore referred to as “intuitive” (Par-
sons, Young, Stein, and Kringelbach (2017), for review).
We suggest that intuitive behaviors include and follow from orienting and recognition capaci-
ties. For instance, adults’ intuitive behavior also drives interpersonal contact between caregivers and
infants. For example, by attempting to stay within the middle of the infant’s visual field at an optimal
distance of approximately 30 cm (Von Hofsten et al., 2014) and by making direct eye contact with
the infant. When eye contact is achieved, the adult may respond with vocalizations of greeting, often
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in a manner known as “infant-directed speech” or “motherese”, which entails high pitch and exag-
gerated intonation, and is preferred by infants to adult-directed speech (Fernald, 1985). Adults may
also make exaggerated facial expressions (Papousek and Papousek, 1983). Within days following
birth, parents and their infants also unconsciously mirror each other’s emotional expressions (Melt-
zoff and Moore, 1977; Trevarthen, 1977). These intuitive, preverbal interactions form the foundation
of a socioemotional understanding on which complex attachment relationships can be built (Bowlby,
1982; Trevarthen and Aitken, 2001).
In this chapter we focus on the initial aspects of the parent-infant relationship: orienting and
recognition. Studying the first point of parent-infant interaction allows us to explore how intuitive
parenting behaviors emerge and evolve. Understanding these early points of interaction may also
inform us about attachment, intersubjectivity, and higher-order caregiving capacities. For example,
behavioral sensitivity, generally defined as parental availability and appropriate, prompt, responsive-
ness to infant cues, has been shown in two meta-analyses to be a key predictor of attachment out-
comes (Bakermans-Kranenburg, van IJzendoorn, and Juffer, 2003; De Wolff and van IJzendoorn,
1997), although the “transmission gap” should be noted—the gap between what can and cannot be
explained about the determinants of attachment security in infancy (Belsky, 2002; Verhage et al.,
2016).
Recent and expanding evidence has suggested that the human parental brain is a distributed net-
work with precise temporal involvement of subcortical and cortical regions. Of crucial importance
is the orbitofrontal cortex (OFC) in the frontal lobes, which appears to be implicated in different
phases of caregiving behavior (Lorberbaum et al., 2002; Parsons, Young, Mohseni, et al., 2013). Given
the region’s role as a nexus of reward-related processing (Kringelbach, 2005b; Rakic, 2009; Zald and
Rauch, 2006), and the often inherently rewarding nature of the parent-infant relationship, such a key
role in the human parental brain is consistent with current understanding of OFC function.
As well as delineating the brain regions involved in the human parental brain, we are getting
closer to understanding the involvement of different regions over time. A temporal understand-
ing complements our structural understanding, and offers a greater insight into the driving forces
within preferential processing of infant cues, as well as allowing us to separate brain processes into
fast, instinctive processes, and slower, more deliberative efforts. A comprehensive understanding of
the neurobiology of parenting requires us to recognize how parenting may alter the brain, both
structurally and functionally and allows us to explore psychiatric disorders where parenting behavior
can be disrupted, which in turn raises the risk for behavioral, emotional, and cognitive difficulties
in the children. The potential for new neuroscientific methods such as whole-brain computational
modeling, offers great potential to explore the human parental brain in more depth than ever before.
Historical Considerations in Understanding

the Neurobiology of Parenting
Given the relative novelty of methods to explore the neurobiology of the human parental brain, such
as functional magnetic resonance imaging (fMRI) and magnetoencephalography (MEG), the history
of research in this field is relatively sparse. However, several key scientists, such as Charles Darwin,
paved the way for our current understanding, and certain cultural and familial factors play an ever-
increasing role in understanding the neurobiology of human parenting.
Darwin
In The Expression of the Emotions in Man and Animals, Darwin (1872) extolled the virtues of study-
ing animals alongside humans for clues to our rich experiences. For Darwin, emotions had an
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evolutionary history that could be traced through cultures and across species. Darwin proposed that
observing infants is a good starting point for understanding facial expressions, as infants exhibit many
emotions in a “pure” form. In addition, he wrote of the strength of maternal love, and how such
strong emotions can lead to instinctive behavior:
No emotion is stronger than maternal love; but a mother may feel the deepest love for her
helpless infant, and yet not show it by any outward sign; or only by slight caressing move-
ments, with a gentle smile and tender eyes. But let any one intentionally injure her infant,
and see what a change! how she starts up with threatening aspect, how her eyes sparkle and
her face reddens, how her bosom heaves, nostrils dilate, and heart beats; for anger, and not
maternal love, has habitually led to action.
(Darwin, 1872)
Darwin was also fascinated by the visceral cries of hungry, pained, or discomforted infants. He noted
the corresponding facial expression that co-occurs with crying: firmly closed eyes, wrinkled skin
around the eyes, forehead contracted into a frown, widely opened mouth with retracted lips. He
collected and documented images of crying infants as he believed the photographs best showed
the “instantaneous process” of crying. Darwin also wrote about smiling and laughter as examples of
early emotional expressions in infants, such as the instinctive peals of laughter that ensue from being
tickled.
In many ways, Darwin was one of the first scientists to explore infant communicative cues and
how they attracted instinctive adult behaviors, paving the way for what we know today about the
human parental brain.
Lorenz and Kindchenschema

Pioneers of the study of infant cues and behavioral responsivity were Nobel Prize winners and
founding fathers of ethology, Konrad Lorenz and Niko Tinbergen. They focused on visual cues,
proposing that infantile facial and bodily features, such as chubby cheeks, big round eyes, a small nose
and ears, form a “Kindchenschema” or “infant schema” (Lorenz, 1943). They further suggested that
the Kindchenschema is a prime example of an “innate releasing mechanism” that naturally attracts
adults and motivates the provision of care through instinctual behaviors. The duo provided a larger
ethological program, including four goals, to define how we should approach the biological study
of behavior. They involved defining the physiology, the survival value, evolution, and development
of behavior (Burkhardt, 2014; Tinbergen, 1963). These goals are still at the forefront of ethological
research, however there does exist debate about the nature of “instincts” and the concept of “innate
releasers” (Hinde, 1966, 1982; Lehrman, 1953). It may be said that infant visual cues promote caregiv-
ing, but do not determine it. This early theoretical account provided a lasting legacy in terms of how
to conceptualize the impact of infant visual cues on caregiver behavior.
Lorenz focused on visual cues, and other researchers focused on cues in the auditory domain.
Early accounts of infant vocalizations suggested a range of motivations for promoting caregiving
behavior, including terminating an aversive stimulus, empathic responding to reduce another’s dis-
tress, or evolutionarily driven responding ensuring the well-being of offspring (Murray, 1985, for
overview). Later, Murray proposed a model that considered infant acoustic cues as “motivational
entities”, promoting the likelihood of initiating behavioral responses in listeners (Murray, 1979,
1985). The motivational entity model states that the specific acoustic structure of an infant cry alerts
the listener rapidly and universally, whereas other factors such as the context of care or cognitive
appraisal have an impact on the selection of specific caregiving behavior.
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Bowlby and Attachment

Bowlby (1969) proposed attachment theory, which asserts that humans are born with a psycho-
biological system that motivates them to seek proximity to a significant other (he focused on the
mother), with the principal aim of ensuring survival. Attachment theory provides an explanation
of how the parent-infant relationship emerges and influences subsequent development. An infant
needs a secure relationship with one primary caregiver for the child’s successful social and emotional
development, and especially for learning affective regulation. The primary caregiver should provide
a secure base from which the child may face the external world, and to which he or she may return
to be comforted if distressed or reassured if frightened. Alongside the infant’s propensity to form an
attachment, Bowlby also proposed that caregiving is the result of an organized behavioral system.
This caregiving system has the goal of promoting proximity and comfort in the event that the child is
stressed by internal or external cues. Bowlby’s attachment theory remains a strong influence on cur-
rent research into parenting, and evidence for the “caregiving system” may be substantiated within
the neurobiological networks in the human parental brain.
Cultural Differences
Parenting is a complex behavior that varies across cultures (Bornstein, 2012). Cross-cultural differ-
ences have not been well addressed in terms of the neurobiology of parenting, but we could assume
that orienting, recognition and intuitive parenting capacities remain similar across cultures, while
higher socioemotional functions may show greater variation. For example, the alterations that
parents make to their speech when directed at infants (“infant-directed speech” or “motherese”;
Kuhl, 2004) involve high pitch and exaggerated variations. Such infant-directed speech is com-
mon across cultures that differ widely in adult-directed speech (Broesch and Bryant, 2014), which
suggests that early engagement with infants may follow very similar neurobiological mechanisms
across cultures. Contrastingly, play is an area where cultural differences may abound, as it is fostered
through their social relationships (Bornstein, 2007; Sutton-Smith, 1998). It seems that the further
we travel from the initial infant-parent contact, the greater the scope for variations in interaction
driven by culture.
Theoretical Perspectives on the Neurobiology Underlying Parenting
The Pleasure Cycle

Interactions with infants can be intrinsically rewarding, and the parent-infant relationship provides
many pleasurable moments. Indeed, the typical adult response to an infant is a smile, the most con-
spicuous of pleasure responses (Hildebrandt and Fitzgerald, 1979). Adaptive pleasures that ensure
species survival include food, sex, and social interaction (Kringelbach and Berridge, 2009a). Social
interaction, including parent-infant interaction, can be examined within the framework of a com-
plex set of processes, each involving distinguishable neurobiological mechanisms (Kringelbach and
Berridge, 2009b; Leknes and Tracey, 2008). The “Pleasure Cycle” includes at least three psycho-
logical components, named “wanting”, “liking”, and “learning” (Figure 7.1). “Liking” represents the
hedonic impact of a reward, whereas “wanting” promotes the approach towards and consumption
of the reward, sometimes called incentive motivation. “Learning” involves adapting expectations
about rewards and their future consumption. Once reward-related cues are learned, the brain is able
to represent predictive associations and cognitions, underpinned by prediction-error monitoring in
the orbitofrontal cortex (OFC). Each component has conscious and subconscious subcomponents.
In the context of social interaction, the initial “wanting” for a reward (i.e., the inherent pleasure of
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Figure 7.1 The pleasure cycle and pleasure network.
A. The pleasure cycle for social interaction. Fundamental (i.e., rewards associated with behavior necessary for
species survival) and higher order pleasures are associated with a cyclical time course. Typically, rewarding expe-
riences go through a phase of expectation or wanting for a reward. This may or may not lead to a phase of con-
summation or liking of the reward, which may reach a peak of pleasure (e.g., eliciting a laugh from your child).
This period of consummation is followed by a satiety or learning phase, during which the individual learns and
updates predictions for the reward to guide future behavior. Learning can also take place throughout the cycle.
B. The core affect elicited by infant cues such as facial cuteness or laughter is generated by the pleasure network
in the human brain (shown here in one hemisphere only) with the nucleus accumbens and ventral pallidum
and other main pleasure-coding regions. Clockwise views (from bottom left) are from the top, front, side, and
3D perspective. The connections indicate the tentative functional networks mediating hedonic ‘liking’ reactions
and subjective pleasure ratings.
C. Schematic diagram of the human “parental brain”—networks of regions typically recruited in response to
infant cues.
interacting with your child) often results in behavior to obtain the reward (i.e., holding the infant
and initiating conversation), which is subsequently “liked” and learned from.
Fast and Slow Thinking

In Thinking Fast and Slow, Kahneman (2011) suggested that the human mind consists of two compet-
ing systems. His central hypothesis posits a dichotomy between two modes of thought: System 1 is
fast, instinctive and emotional; System 2 is slower, effortful, more deliberative, conscious, and more
logical. To illustrate the two, imagine you see a baby pass you in a pram—you may react instinctively
by smiling. This is an example of the “fast” system. Our brains are hardwired to respond quickly to
certain cues in the environment, and this helps us to survive. Yet there are many situations when
our brains take longer to complete tasks—using Kahneman’s “slow” system. For instance, you may
be holding a crying infant and not know why he or she is crying. You probably really have to think
through the options for why the infant is crying and work out a plan to soothe the infant and address
his or her needs—that is your “slow” system which allows you to process more tricky situations.
Fast responses are important for survival, and slow responses are a vital aspect of caregiving and its
constituent processes.
In accord with the concept of fast and slow thinking, Murray’s motivational entity model (Mur-
ray, 1979, 1985) has since been broadly substantiated within general neuroscientific and conceptual
models of emotional processing, which outline a rapid emotional reaction (“core-affect”), which
can be modulated by contextual factors, attentional states, and cognitive appraisal (Barrett, Mesquita,
Ochsner, and Gross, 2007). This two-stage model also describes the neural hardware supporting
emotional responses, with rapid, imprecise processing recruiting primarily subcortical regions, and
slower, more detailed analyses occurring in sensory and higher-order cortical regions (Barrett and
Bar, 2009; LeDoux, 2000; Rolls, 2000). The orbitofrontal cortex (OFC) is considered to be a central
hub within the dual streams model of emotional processing, providing an interface between the fast
and slow processing routes (Barrett and Bar, 2009).
Caregiving can also be viewed in a stratified way based on fast or slow systems. While instinctive,
rapid processes exist, such as orienting to infant cues and recognition of an infant, there also exists
slower caregiving responses, such as prolonged interactions in play.
Classical and Modern Research in the Neurobiology of Parenting
The Social Brain and the Parental Brain

The human parental brain, by which we mean the neural networks controlling parental behavior
and cognitions, must be viewed within the context of the social brain, as the parent-infant relation-
ship provides the template for all social relationships. Social cognition has been considered to be of
such fundamental importance to human functioning that some have argued it may even represent
the “default mode” of cognition (Schilbach et al., 2006). It is evident that, as an evolutionary neces-
sity, we would require continuous attunement to the social environment to prioritize attention to
biologically relevant stimuli, such as a family member, an infant, or a threatening face in a crowd.
Some major brain structures involved in social cognition include the superior temporal gyrus and
fusiform gyrus, superior colliculus, and primary sensory cortices; premotor cortex, OFC, amygdala,
and ventral striatum; the anterior cingulate cortex, as well as prefrontal regions involved in higher-
level processes, such as social reasoning, theory of mind, empathy, and moral cognition (for reviews
see Adolphs, 2003; Eslinger, 1998; Fiske and Taylor, 2008; Frith and Frith, 2010; Frith, 2007; Lieber-
man, 2007; Moll and de Oliveira-Souza, 2007).
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The “parental brain” can be conceptualized as the network of regions that support caregiving
behavior in humans. Adult responses to infant vocalizations and visual cues, in association with func-
tional neuroimaging, have defined this network. The parental brain incorporates both responses in
nonparents (the so-called parental “instinct”, which transcends the biological relationship between
adult and infant) alongside parental responses that develop as a result of experience. Functional mag-
netic resonance imaging (fMRI) has revealed a network of regions active in response to infant facial
and vocal cues, which overlap extensively with the “social brain”.
These findings reflect activity in the frontal lobe (anterior cingulate, prefrontal, orbitofrontal, and
insular cortices), the temporal lobe (middle temporal gyrus, superior temporal gyrus/sulcus, and
inferior temporal gyrus/sulcus), motor areas (motor cortex, premotor cortex, and supplementary
motor area), and in subcortical areas (basal ganglia, periaqueductal gray, and amygdala). Temporal
regions, such as the middle temporal gyrus and superior temporal gyrus/sulcus, and the amygdala are
involved in the initial processing of affect in vocal cues (Belin, Fecteau, and Bedard, 2004; Ethofer
et al., 2006; Frühholz, Trost, and Grandjean, 2014; Yovel and Belin, 2013), and occipital and different
temporal regions such as the fusiform gyrus, are predominantly recruited for the visual processing
of facial expressions (Fusar-Poli, Placentino, Carletti, Landi, and Abbamonte, 2009). In these regions,
stimuli may be concurrently represented and processed by frontal regions, such as the OFC (Bar
et al., 2006), and latterly processed by frontal, limbic, and motor regions for higher-order processing
and to initiate behavioral responses.
Infant Faces
The unique and instantly recognizable facial configuration of infants is pleasing and rewarding, and
an instinctive reaction of adults on seeing an infant is to smile (Hildebrandt and Fitzgerald, 1978).
Such features are often referred to as “cuteness” and are instantly recognizable (Figure 7.2).
This facial configuration is believed to attract attention and evoke caregiving in adults. The pref-
erence for juvenile facial features appears spontaneously and universally, transcending the biological
relationship between an adult and an infant, and also seems to be conserved across multiple species
(Sato, Koda, Lemasson, Nagumo, and Masataka, 2012). Adults present a general aesthetic apprecia-
tion of the infant face (“liking”), measured by explicit ratings of stimulus attractiveness. Adults also
display a greater incentive salience to view infant stimuli (“wanting”) compared to images of adults
(Parsons, Young, Kumari, Stein, and Kringelbach, 2011). This hedonic stratification is important to
consider when assessing the responsivity to infant cues, as it brings together the different elements of
the parental orienting response (such as affect and behavior).
The attentional magnetism of the infant schema and “cute” infant attributes has become a mainstay
of Japanese popular culture, known as kawaii. In this context, neoteny, the process of giving human or
nonhuman stimuli childlike, juvenile features, has been used to alter toys, cartoon characters, enter-
tainment, and fashion. The popular cartoon character Mickey Mouse has been demonstrated to have
become “cuter” and more childlike during the 20th century in parallel with his rise in popularity
(Gould, 1979). Items with “cute” or “kawaii” attributes have similarly been shown to afford an advan-
tage on tasks involving focused attention for completion (Nittono, Fukushima, Yano, and Moriya,
2012), led by heightened approach motivation and systematic processing. Infantile characteristics in
“baby-faced” men also preferentially activate reward systems in the human brain, and induce positive
affect in viewers (Zebrowitz, Luevano, Bronstad, and Aharon, 2009). The effects of baby-faced adults
upon brain activity and mood point to the capacity of the baby schema to guide attention, motivation,
and behavior at early stages of visual and affective processing (Kringelbach et al., 2008).
Neural evidence of responsivity to infant faces and visual cues has provided an important level of
explanation for the field, as visual cues form an important aspect of parent-infant contact. Greater
257
Figure 7.2 Behavioural effects of cuteness.

A. The proportions of the features of the face can be used to provide objective measurements of cuteness in
infants and adults (Glocker, Langleben, Ruparel, Loughead, Gur, et al., 2009).
B. Adult men and women (who are not yet parents) differ in their liking ratings, but not in the amount of effort
they expend on viewing natural images of infants with varying levels of objective cuteness (Parsons, Young,
Kumari, Stein, and Kringelbach, 2011).
C. Artificially changing the proportions of the faces of humans, dogs, and cats can change their perceived cuteness,
although questions have been raised over the ecological validity of such non-ecological image manipulations.
D. Five-year-old children find the young significantly cuter than the adults of different species (Borgi, Cogliati-
Dezza, Brelsford, Meints, and Cirulli, 2014).
Source: Panels C and D are reproduced from Borgi et al. (2014).
activity within the OFC in response to infant faces compared to adult faces has been recognized
across multiple neuroimaging methods, including: electroencephalography (EEG; Proverbio, Riva,
Zani, and Martin, 2011), magnetoencephalography (MEG; Kringelbach et al., 2008; Parsons, Young,
Mohseni, et al., 2013), and functional magnetic resonance imaging (fMRI; Baeken et al., 2009;
Glocker, Langleben, Ruparel, Loughead, Valdez, et al., 2009; Leibenluft, Gobbini, Harrison, and
Haxby, 2004; Montoya et al., 2012; Nitschke et al., 2004; Ranote et al., 2004; Strathearn, Li, Fonagy,
and Montague, 2008). In addition, fMRI studies of the human parental brain have found differential
activity in response to infant facial expressions in the temporal lobe, frontal lobe, motor cortex, and
subcortical regions (Figure 7.3, and Table 7.1 for studies).
A strict test of the power of the infant facial configuration to elicit specialized processing in adults
comes from circumstances where the infant face is naturally altered. It is also possible to measure
258
Figure 7.3 Infant cues elicit fast responses in the human brain.
A. Infant faces are examples of cute stimuli that have been shown to elicit fast brain responses (~130ms) in the
orbitofrontal cortex (OFC), at the same time as responses in the fusiform face region (Kringelbach et al., 2008).
B. Artificially manipulating the cuteness of infant faces has been shown to correlate with changes in the BOLD
signal in the nucleus accumbens, part of the pleasure system (Glocker, Langleben, Ruparel, Loughead, Valdez,
et al., 2009).
C. Very fast neural responses (~50ms) are found in the human brainstem to both positive and negative infant
vocalisations (babbling, laughter and crying) (Parsons et al., 2013).
D. Similar to the fast brain response to cute visual stimuli, infant crying elicits activity in the OFC (~140ms) at
the same time as activity in primary sensory cortices (Young et al., 2016).
Source: Panel (A) is reproduced from Kringelbach et al. (2008); panel (B) is adapted from Glocker, Langleben, Ruparel,
Loughead, Valdez, et al. (2009); panel (C) is adapted from Parsons et al. (2013); and panel (D) is adapted from Young et al.
(2016).
Table 7.1 Summary of findings from fMRI studies of the “parental brain”, highlighting neural regions reac-
tive to infant vocal and facial expressions (MTG: middle temporal gyrus; STS/G: superior temporal
sulcus/gyrus; FFG: fusiform gyrus; OFC: orbitofrontal cortex; PFC: prefrontal cortex; PAG: periaq-
ueductal gray; VTA: ventral tegmental area; SMA: supplementary motor area)
Infant vocal expressions Infant facial expressions
Temporal lobe
MTG Montoya et al. (2012) Montoya et al. (2012)
Kim, Leckman, Mayes, Newman et al. (2010) Ranote et al. (2004)
Lorberbaum et al. (2002) Barrett et al. (2012)
Riem et al. (2011) Strathearn et al. (2008)
STS/G Montoya et al. (2012) Montoya et al. (2012)
Kim, Leckman, Mayes, Newman, et al. (2010) Ranote et al. (2004)
Riem et al. (2011) Strathearn et al. (2008)
Mascaro, Hackett, Gouzoules, Lori, and Rilling (2014) Noriuchi et al. (2008)
Bos, Hermans, Montoya, Ramsey, and van Honk (2010) Bartels and Zeki (2004)
FFG None Strathearn et al. (2008)
Bartels and Zeki (2004)
Caria et al. (2012)
Nitschke et al. (2004)
Frontal lobe
OFC Mascaro et al. (2014) Montoya et al. (2012)
Laurent and Ablow (2012) Ranote et al. (2004)
Strathearn et al. (2008)
Noriuchi et al. (2008)
Nitschke et al. (2004)
Glocker, Langleben, Ruparel,
Loughead, Gur, et al. (2009)
Leibenluft et al. (2004)
Baeken et al. (2009)
Anterior Lorberbaum et al. (2002) Barrett et al. (2012)
cingulate Laurent and Ablow (2012) Strathearn et al. (2008)
Caria et al. (2012)
PFC Kim, Leckman, Mayes, Newman, et al. (2010) Ranote et al. (2004)
Mascaro et al. (2014) Strathearn et al. (2008)
Laurent and Ablow (2012) Noriuchi et al. (2008)
Insula Mascaro et al. (2014) Barrett et al. (2012)
Kim, Leckman, Mayes, Newman, et al. (2010) Noriuchi et al. (2008)
Laurent and Ablow (2012) Bartels and Zeki (2004)
Caria et al. (2012)
Subcortical
Basal ganglia Lorberbaum et al. (2002) Montoya et al. (2012)
Mascaro et al. (2014) Barrett et al. (2012)
Montoya et al. (2012) Strathearn et al. (2008)
Laurent and Ablow (2012) Noriuchi et al. (2008)
Infant vocal expressions Infant facial expressions
PAG/VTA Laurent and Ablow (2012) Strathearn et al. (2008)

Amygdala Riem et al. (2011) Ranote et al. (2004)
Seifritz et al. (2003) Barrett et al. (2012)
Strathearn et al. (2008)
Strathearn and Kim (2013)
Leibenluft et al. (2004)
Thalamus/ Lorberbaum et al. (2002) Barrett et al. (2012)
hypothalamus Strathearn et al. (2008)
Caria et al. (2012)
Motor
Motor cortex Kim, Leckman, Mayes, Newman et al. (2010) Strathearn et al. (2008)
Premotor/SMA Montoya et al. (2012) Caria et al. (2012)
the potential impact on parent-infant interaction in such circumstances. Perhaps the most convinc-
ing evidence for the significance of infant facial features in eliciting parental care has come from
instances of facial anomaly in the case of cleft lip and palate. Cleft lip is the most common congenital
condition affecting the face and cranial bones, with an incidence of 1 in 700 live births in the United
Kingdom (Goodacre and Swan, 2008). It affects the nasal and mouth regions and can therefore be
conceived of as a specific, local change to the typical infant schema. It is also important to note that
cleft lip does not typically occur with concurrent developmental difficulties—the sole difference in
these children is their facial configuration.
Cleft lip in infancy has been the most studied facial abnormality with reference to adult motiva-
tional processing, and adults will typically rate infants with cleft lip as less “cute” than healthy infants,
and will work less hard to view them (Parsons, Young, Parsons, et al., 2011). Nonparents have also
been shown to react negatively to global changes to the infant face, in cases of fetal alcohol syndrome
and prematurity (Frodi, Lamb, Leavitt, and Donovan, 1978; Waller, Volk, and Quinsey, 2004). The
presence of a cleft lip disrupts typical gaze patterns, with fixations to the mouth region predomi-
nating (Rayson et al., 2016). Mothers of infants with cleft lip look at their infants for less time, and
demonstrate less positive and sensitive interaction compared to mothers of healthy infants (Murray
et al., 2008). These findings together show that changes to the infant facial configuration appear to
compromise adult responsivity.
Infant Crying
Early infant communication is primarily composed of facial and vocal cues. The newborn infant
cries reflexively and communicatively, acting as a “biological siren” to alert the caregiver to the
infant’s presence and needs. MacLean (1990) drew attention to the importance of the universal
mammalian caregiving cue of infant vocalizations as an important indicator of how mothers and
fathers respond to their infant. Infant cries portray a “falling” or “rising-falling” melody, with high
and dynamic pitch (ranging between 250 and 750 Hz; Golub and Corwin, 1985). Single bursts last
for between 1 and 3 seconds. Initially, infant cries are largely reflexive, indicating hunger, tiredness,
pain, or separation from caregivers. Their cries may also take on more variation in acoustics over
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time. For instance, other vocalizations, such as babbling and laughter, emerge postnatally, as infants
gain greater control of the vocal tract (Soltis, 2004). Infant cries also appear to be “graded signals”
in which changes in acoustic features, such as pitch, indicate varying levels of distress (Soltis, 2004),
although limited success has been found in attempts to divide infant cries into acoustically distinct
categories, such as pain or hunger. A higher-pitched cry is frequently perceived as more distressed
and more arousing by adult listeners, both parents and nonparents (Parsons et al., 2014; Porter, Miller,
and Marshall, 1986; Young, Parsons, Stein, and Kringelbach, 2012; Zeskind and Marshall, 1988). The
ability to respond to infant vocalizations, universal in mammalian species, is of fundamental impor-
tance to parental responsivity (MacLean, 1990).
Hearing a distressed infant’s cry has a noticeable effect on behavior. It elicits autonomic arousal in
the listener, as measured by physiological correlates such as heart rate, blood pressure, skin conduct-
ance, and handgrip force (Bakermans-Kranenburg, van IJzendoorn, Riem, Tops, and Alink, 2012;
Boukydis and Burgess, 1982; Zeskind and Collins, 1987). Furthermore, parents who exhibit greater
physiological arousal are more likely to respond rapidly (Del Vecchio, Walter, and O’Leary, 2009),
demonstrating a link between the magnitude of physiological response and parental responsivity.
These physiological responses appear to ready adults to react behaviorally to infant cries.
If such physiological changes are functionally important, in response to hearing the cries of an
infant, then they should translate into altered behavioral responsivity. Indeed, hearing infant cries has
been shown to disturb normal performance on simple cognitive tasks (Morsbach, McCulloch, and
Clark, 1986) and disrupts cognitive control (Dudek, Faress, Bornstein, and Haley, 2016). This evi-
dence of altered behavioral responsivity could reflect the magnetism of infant cries to orient people
away from less biologically salient tasks. Another study found that after listening to infant distress
cries, compared to adult distress vocalizations or bird sounds, participants showed improvements in
fine motor performance (Parsons, Young, Parsons, Stein, and Kringelbach, 2012). Accordingly, infant
cries orient us away from distracting tasks, but they may also afford an advantage in subsequent motor
movements, which may reflect a behavioral readiness to address the distressed infant (Caria et al.,
2012). The ability to respond appropriately to infant cries has also been associated with the ability
to respond appropriately to other infant cues (Frodi and Lamb, 1980), suggesting that responsivity is
not modality-specific, but could reflect a common mechanism.
Infant cries express key affective and physiological information and have been demonstrated to
elicit specialized activity in an array of brain regions. As with infant faces, the OFC appears to be
a key region involved in responding to infant cries (Young et al., 2016). Regions in the temporal
lobe, frontal lobe, motor cortex, and subcortical regions are all also involved in processing infant cries
(Table 7.1).
It remains unclear how brain regions involved in the response to infant cries are sensitive to differ-
ences, such as graded acoustic information. Differential brain activity in response to infant cries must
also be viewed alongside neuroendocrine system activity. For example, peripheral levels of oxytocin,
the “neurohormone of attachment”, have been found to be higher in mothers who demonstrate
secure attachment to their infant, compared to mothers who demonstrate unstable attachment pat-
terns (Strathearn, Fonagy, Amico, and Montague, 2009). Crucially, mothers with higher levels of oxy-
tocin also show correspondingly higher levels of activity in reward-related brain regions (Strathearn
et al., 2009). Associating endocrine factors alongside both behavioral differences and brain activity
promises to elucidate the functional properties of parenting networks by affording further variables
that may moderate the parent-infant relationship.
Laughter
Alternatively termed “auditory cuteness” (Kringelbach et al., 2016), laughter is another infant cue that
communicates the infant’s affective state and may track other cognitive developments by presenting
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a sign of pleasure at cognitive mastery (Addyman and Addyman, 2013). Laughter emerges at around
four months after birth (Ambrose, 1963; Darwin, 1872), and probably proceeds from smiling which
becomes contingently linked to maternal talk and maternal smile by the second and third months
(Lavelli and Fogel, 2002). Smiling and laughter also serve to attract adult attention, eliciting warmth
and care (Bowlby, 1969).
Infant distress cries have taken precedence when exploring the adult’s neural response to infant
vocalizations, yet laughter is powerful and important cue in maintaining caregiver interactions and
facilitating attachment. As an example, the incentive salience of infant laughter appears to be elevated
by oxytocin administration in nulliparous women (Riem et al., 2012), which is argued to enhance
parent-infant bonding. The valence of the infant cue (happy or sad, for example) may also modu-
late the neural responses (Montoya et al., 2012), suggesting that positive and negative affect in the
infant may draw on different neural resources in the adult. The OFC has been implicated in both
the detection and evaluation of reward valence, in addition to monitoring ongoing reward values
(Kringelbach, 2005a). Particularly where a “punishment” is possible, the lateral portion of the OFC
is involved in responding to counteract negative reinforcement.
One fMRI study exploring the neural response to infant laughter and infant crying compared
parents with nonparents. (Seifritz et al., 2003) found a significant group (parents versus nonparents)
and stimulus (laughter versus crying) interaction. Differential neural responses were found in the
right amygdala, the middle cingulate cortex, insula, left ventral prefrontal cortex, and the left tem-
poroparietal junction. Parents showed stronger activity in these regions in response to infant crying,
whereas nonparents showed stronger activity to infant laughter. In addition, for both parents and
nonparents, the infant vocalizations compared to the control stimulus produced differential activ-
ity in the Heschl’s gyri and temporal and polar planes of the auditory cortex, representing an early
stage of cortical auditory processing. These findings suggest that infant vocalizations evoke early dif-
ferential cortical processing and also provide evidence for experience-dependent neural plasticity as
a direct result of parenthood and exposure to infant cues. Despite this preliminary evidence, infant
laughter remains an underused stimulus in studies of adult response to infant cues, and could offer
great insight into the human parental brain by revealing how positive affective cues relate to specific
caregiving behavior.
Touch
Touch represents a further modality in research linking adult responsiveness and infant cues. A large
literature explores the effect of parental touch on infant outcomes. Touch appears to have many
beneficial outcomes, especially with preterm infants (Smith, 2012). There is evidence that the sense
of touch is the first to develop; the fetus even appears able to respond to touch as early as the 21st
week of gestation (Marx and Nagy, 2015). Both mothers and fathers can recognize their newborn
infants by touch alone (Kaitz et al., 1992; Kaitz, Shiri, Danziger, Hershko, and Eidelman, 1994).
Tactile stimulation often seems to be a behavioral response to infant cues from other modalities, for
example, mothers across several cultures preferentially respond to their infants’ vocalizing distress by
picking up and holding their infant (Bornstein et al., in press). Human parents touch their infants
frequently during face-to-face interactions, with infants engaged in touching behaviors roughly 85%
of the time (Moszkowski and Stack, 2007) and parental touch can regulate an infant’s physiological
and behavioral arousal (Hofer, 1994). The type of touch that parents use appears to be highly adaptive
in engaging and stimulating their child. Parents spontaneously give slow moving, gentle touch, which
is optimized to activate a specialized system of nerve fibers called “C-tactile afferents” which are
purported to underpin the rewarding sensation of touch (Pawling, Cannon, McGlone, and Walker,
2017). Despite the wealth of knowledge linking infant touch and the parent-infant relationship, lit-
tle is known about the underlying neurobiology that processes such cues. This is likely to be due to
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practical difficulties of measuring parental responses to their own infant’s touch during neuroimag-
ing sessions, although it should not be ruled impossible.
Olfactory Cues
Olfactory cues also likely drive intuitive parental responsivity to an infant. Whereas auditory and
visual cues are frequently distal infant cues, bringing the caregiver into physical contact with the
infant, olfactory cues may only be effective once the infant and parent are in close proximity. Moth-
ers demonstrate considerable success with recognizing their newborn infant by olfactory signals.
Mothers have been shown to recognize their infants via smell with just 10 minutes prior contact
(Kaitz, Good, Rokem, and Eidelman, 1987). Paternal recognition of their newborn infant by olfac-
tory cues, in contrast, has yielded rather inconclusive results. Bader and Phillips (2002) asked fathers
to sniff the heads of three newborn infants (all bathed identically) and then attempt to identify their
own infant. Contrary to the findings with mothers, the fathers were unable to choose their newborns
correctly by smell at a rate significantly greater than chance. This was found for both fathers with
low exposure (1–10 minutes) and high exposure to the infant (11–810 minutes), and despite greater
exposure to the infant having facilitated visual recognition within the same participant group. These
findings suggest that fathers learned visual facial cues rapidly, similar to mothers, but did not rapidly
learn olfactory cues related to their infant.
There are disappointingly few studies concerning early olfactory recognition by parents and
related caregiving behaviors. Weisfeld, Czilli, Phillips, Gall, and Lichtman (2003) explored human kin
recognition by olfactory-based mechanisms. They found a very low rate of kin-nonkin confusions,
suggesting a possible “family odor cue”, even when controlling for effects of familiarity. Furthermore,
mothers were preferentially able to identify their biological children’s odor, but not that of their
stepchildren, hinting that environmental similarity and continual association are not sufficient for
olfactory recognition. Whether newborns and young infants possess a distinct olfactory cue separable
from any “family odor cues” remains to be explored. It might be expected that young infants possess
distinct olfactory cues, as studies with rodents suggest that the processing of olfactory cues by the
amygdala may mediate avoidance of young pups by nulliparous rat females, leading to fewer mater-
nal behaviors (Numan and Sheehan, 1997) in the same way that aversive responses to foul odors are
mediated by amygdala activity (LeDoux, 1996). Therefore, although olfaction is not the dominant
sensory modality for human parental care as it is for other mammals, olfactory cues may be important
to initiate orienting and recognition, and promote attachment.
The Temporal Dynamics of the Human Parental Brain

Studies exploring the neural response to infant cues, including infant faces, crying, laughter, and even
touch and smell, have clearly defined a core set of regions that encompass the human parental brain.
However, a more detailed exploration of the brain response to infant cues demands that we ascertain
the temporal involvement of these regions.
Early detection of infant vocalizations in the brainstem. During deep brain stimulation for chronic pain,
four adults were exposed to infant vocalizations (a mixture of babbling, cries, and laughter) while
electrodes measured activity in the periaqueductal gray (PAG) of the midbrain (Parsons, Young,
Joensson, et al., 2013). Local field potentials (LFPs) reveal differences to infant sounds compared to
natural control sounds (adult cries and animal distress sounds) as early as 49 ms after the sound onset.
Although the spatial resolution of LFPs is not sufficient to conclusively differentiate PAG activity
from that of neighboring regions, the PAG has been associated with maternal responsiveness in non-
human models (Lonstein and Stern, 1997; Miranda-Paiva et al., 2007; Sukikara, Mora-Ortiz, Baldo,
Felicio, and Canteras, 2010) and has a role in regulating arousal. Its extensive network of anatomical
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connections includes the inferior colliculus (Dujardin and Jürgens, 2005), the amygdala, and frontal
lobe regions including the OFC (Cavada, Company, Tejedor, Cruz-Rizzolo, and Reinoso-Suárez,
2000). These connections may allow for rapid propagation of information regarding infant cues
through a network of cortical and subcortical regions that support the human parental brain. Quick
orienting to the infant may therefore be supported by a state of heightened physiological arousal
initiated by the PAG. A listener’s heart rate, respiration rate, and hand grip strength have all been
shown to be affected by hearing infant cries, and these peripheral psychophysiological measures may
represent the heightened arousal. Early detection of salient infant cues in the brainstem may therefore
constitute the first step in the initiation of rapid, effortful, caregiving behavior, and support more
detailed processing of cue salience and meaning.
Rapid cortical sensitivity: Modulation of the N100. The N100 component, the negative deflection
peak around 100 ms after the onset of a stimulus, is an obvious target for clues about early cortical
processing of infant cues. There is conflicting evidence regarding whether the N100 is sensitive to
the affective content of infant cues. A recent EEG study found that the emotional expression of an
infant face (happy, neutral, and distressed) can modulate the N100 in mothers (Peltola et al., 2014).
However, other studies have reported no early effects due to valence (Noll, Mayes, and Rutherford,
2012; Proverbio, Brignone, Matarazzo, Del Zotto, and Zani, 2006). A further EEG study of mothers
compared processing of one’s own infant’s face to an unfamiliar infant and found greater power in
gamma band activity occurring before 100 ms (Esposito, Valenzi, Islam, Mash, and Bornstein, 2015).
Regarding valence, the contrasting effects may be due to differing task and attentional demands,
whereas directing attention to infant stimuli and their emotional valence may drive earlier process-
ing of such cues (Malak, Crowley, Mayes, and Rutherford, 2015). Selective recognition of one’s own
infant and sensitivity to the emotional content of their cues may begin at a very early stage in visual
processing.
The orbitofrontal cortex: a crucial role in salience detection and beyond. The OFC may be engaged in
several phases of parent-infant interactions, starting with early salience detection of infant cues, and
progressing to ongoing monitoring of the interaction and its social rewards, and subsequent learning
about the infant cues and their corresponding reward value. The OFC occupies the ventral surface
of the frontal part of the brain, receiving projections from the magnocellular medial nucleus of the
mediodorsal thalamus (Fuster, 1997). It receives inputs from the five sensory modalities: visual, audi-
tory, gustatory, olfactory, and somatosensory, plus visceral sensory information (Carmichael and Price,
1995b). It enjoys direct reciprocal connections with other key brain regions, including the amygdala
(Amaral and Price, 1984; Carmichael and Price, 1995a), cingulate cortex (Öngür and Price, 2000;
Van Hoesen, Morecraft, and Vogt, 1993), insula/operculum (Mesulam and Mufson, 1982), hypothal-
amus (Rempel-Clower and Barbas, 1998), hippocampus (Cavada et al., 2000), striatum (Eblen and
Graybiel, 1995), periaqueductal gray (Rempel-Clower and Barbas, 1998), and dorsolateral prefrontal
cortex (Barbas and Pandya, 1989; Carmichael and Price, 1995b). It is therefore uniquely positioned
between subcortical and cortical pathways and comprises a nexus of sensory processing. The OFC
has established roles in affective processing, reward, and social cognition, making it a prime target for
understanding rewarding interpersonal interactions such as in the parent-infant domain. The OFC is
believed to play a key role in coordinating both slow and fast responses to affective stimuli.
Of note, the “affective prediction hypothesis” proposes that the OFC is implicated in “tagging”
emotionally salient stimuli early in time (Barrett and Bar, 2009). This rapid salience detection is
suggested to influence ongoing sensory processing and to prime rapid motor responses. There is
growing evidence that the OFC is involved in the early salience detection of infant cues. Kringel-
bach et al. (2008) found a possible biological basis for the specific relation between an infant’s facial
features and an adult’s caregiving response. They used MEG to record neuronal activity while par-
ticipants viewed images of adult and infant faces. In response to the infant images, there was highly
specific brain activity present within 130 ms in the medial orbitofrontal cortex (mOFC), followed by
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expected activity in the right fusiform “face area” (FFA) as for the adult faces at around 170 ms. For
evidence of this as a marker of “intuitive parenting”, Kringelbach et al. followed the same reasoning
as Papousek (2000) who had argued that parental responses of such fast latency could be considered
to be instinctive or hardwired. This finding has since been replicated (Parsons, Young, Mohseni et al.,
2013), is found in both men and women, and appears to be present in nonparents as well as parents.
As previously mentioned, cleft lip represents a natural deviation from the typical infant facial
configuration, and thus provides a strict test of the power of the infant face to elicit specific activity
in the parental brain. To explore the effects of cleft lip on neural processing, specifically probing the
causal relation between infant facial configuration and OFC activity, functional neuroimaging using
MEG has been used to compare viewing healthy infants with images of infants with cleft lip (Par-
sons, Young, Mohseni, et al., 2013). In this study, the previous finding of Kringelbach and colleagues
(2008) of significant medial OFC activity in response to healthy infant faces at around 130 ms was
replicated. However, when presented with images of infants with a cleft lip, this medial OFC activ-
ity was diminished as in the case of adult faces. These findings suggest that early OFC responses are
sensitive to the baby schema or typical infant facial configuration.
Evidence for salience detection mediated by the OFC in response to infant cues is also forthcom-
ing in other modalities. Young et al. (2016) compared the neural responses of participants listening
to infant and adult cry vocalizations. They observed a peak of early activity localized in the OFC at
125–135 ms for infant cries but not adult cries, in accord with previous findings of early OFC activ-
ity for visual infant cues. They also reported a second peak of differential activity at a later time point
between 190–200 ms in the OFC. Such specific brain activity in the OFC could be characterized as
a potential biological basis for the “innate releasing mechanisms” initially posited by Lorenz, and the
motivational entity model refined by Murray, promoting, but not necessarily determining caregiving
in response to infant faces. The orbitofrontal cortex is a key brain region in the representation of
reward, so could plausibly represent the especially high reward value of infant visual cues. This emo-
tional “tagging” of the stimulus may bias further neuronal processing to garner energy and resources
for providing care for the infant.
Beyond this early salience detection, activity in the OFC has been demonstrated to represent the
reward value of stimuli, which is a function likely to be based on more detailed processing (Kringel-
bach and Rolls, 2003), and crucial for higher-order processing and decision making (Berridge and
Kringelbach, 2008). Orienting to infants appears fast, effortless, and spontaneous, but parenting also
involves slower process of decision making, such as why the infant could be crying and what the
infant needs. When making decisions, the brain must predict and evaluate the reward values of
the stimuli involved, and the reward values of various behaviors involved in interacting with each
stimulus (Kringelbach, 2005b; Rangel, Camerer, and Montague, 2008). The OFC is a key region in
these processes as neuroimaging studies have found that the reward value, expected reward value, and
even subjective pleasantness are represented in the OFC (Gottfried, O’Doherty, and Dolan, 2003;
Kringelbach, O’Doherty, Rolls, and Andrews, 2003; O’Doherty et al., 2000). The OFC appears to
have functional subregions that subserve different aspects of reward-related processing. The medial
OFC (mOFC) has been related to monitoring, learning, and memory for reward, and the lateral
OFC (lOFC) has been proposed to relate primarily to the evaluation of “punishers”, which can lead
to changes in behavior (Kringelbach and Rolls, 2004). A posterior-anterior gradient in the OFC is
also present, with more complex or abstract reinforcers (such as monetary gain or loss) being rep-
resented more anteriorly, and less complex sensory rewards (such as taste) being represented more
posteriorly (O’Doherty, Kringelbach, Rolls, Hornak, and Andrews, 2001; Small, Zatorre, Dagher,
Evans, and Jones-Gotman, 2001).
Responsive and attentive caregiving is likely to draw on a range of functions mediated by the
OFC, as responsivity to infants requires ongoing monitoring, learning, and memory for infant cues.
It also involves the ongoing evaluation of these cues to appropriately adapt behavior. Learning about
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infant cues via orbitofrontal activity may form the basis of experience-dependent plasticity in the
human parental brain. In addition, a region of mid-anterior OFC is believed to track changes in
subjective pleasure (Kringelbach et al., 2003), and may therefore provide a neural correlate of the
subjective pleasure often inherent in ongoing infant interaction. Within dynamic parent-infant con-
tact, prediction-based learning and choice behavior in uncertain environments are likely to occur.
Such processes may allow for optimization of “intuitive” parenting behaviors by learning through
prediction-making and subsequent analysis of error. OFC activity in response to infant cues may
exemplify how the brain coordinates a seemingly “cognitively impenetrable” orienting response to
infants (Fodor, 1983).
Detailed sensory processing (N170 and beyond). Following early neural responses, it is presumed
that infant cues are exposed to more detailed sensory processing. The N170 is a face-sensitive ERP
component that elicits a larger response at occipitotemporal electrodes for human faces compared
to nonface objects. Here, there are mixed results for infant-related processing. Some studies have
reported no effect of valence in infant facial expressions on N170 responses (Malak et al., 2015; Noll
et al., 2012). Contrastingly, another study revealed larger N170 amplitudes to negative compared to
positive infant facial expressions during focused attention in both mothers and nonmothers, as well
as a main effect of valence on latency of N170 responses (Peltola et al., 2014).
For later EEG components, there is greater correspondence between results. These later dif-
ferences are expected to reflect slower processes, such as detailed sensory processing and cognitive
appraisal. Larger N2 responses within the 250–300 ms window were found in response to infant
faces, compared to pre-pubertal child faces, which also had greater responses than to adult faces
(Proverbio et al., 2011). These effects were subsequently localized to the fusiform gyrus, anterior
cingulate cortex, and OFC. Two further studies reported effects of the valence of infant cues on
responses between 200 and 300 ms (Peltola et al., 2014; Proverbio et al., 2006). Both intensity and
valence of emotional expressions affected the amplitude of the P300 response in one study (between
375 and 600 ms; Proverbio et al., 2006), although another study found no difference in a comparable
time window of 300 to 450 ms (Peltola et al., 2014). Last, two studies reported modulation of the
late positive potential (LPP; 500–800 ms) by valence in infant cues (Malak et al., 2015; Rodrigo et al.,
2011). All of these later effects demonstrate differential processing of infant cues at later time points,
possibly reflecting slower appraisal processes.
The effects of infant cues on behavior: Evidence for preparatory motor responses? There is preliminary
evidence that infant cues may also influence a preparatory motor response in the brain, spurring us
into action. The medial portions of the OFC guide autonomic, endocrine, and behavioral responses
to an object (Barbas and De Olmos, 1990; Carmichael and Price, 1995b, 1996; Ghashghaei and
Barbas, 2002; Ongur, Drevets, and Price, 1998; Rempel-Clower and Barbas, 1998) and coupled
with strong reciprocal connections to lateral parietal regions, the OFC is well placed to coordinate
preparatory motor responses (Barrett and Bar, 2009). In concordance with the connectivity between
the OFC and motor regions, premotor cortex and supplementary motor area activity in response
to infant face stimuli, compared to adult face stimuli, has been found in both nonmothers (Caria
et al., 2012) and mothers (Strathearn et al., 2008). Similar motor cortex activity has been found in
response to auditory infant cues (Kim et al., 2011; Montoya et al., 2012). Using auditory MEG, one
study observed differential activity at around 180 ms to infant cries, localized to the motor cortex
(Young et al., 2016).
A study using transcranial magnetic stimulation demonstrated enhanced motor evoked potentials
at 100–200 ms after hearing infant cries, interpreted as reflecting an automatic audiomotor response
(note that this effect may be specific to female listeners, a finding which requires further assessment,
Messina et al., 2015). These findings could be suggestive of a readiness for communicative behavior
with infants as well as for attachment and caregiving. A potential biological mechanism to predispose
us to be responsive to our young appears to transcend the adult’s biological relationship to the infant,
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as motor responses in nonparents demonstrate (as was Kringelbach et al., 2008). It therefore seems
that in both men and women, whether they are parents or not, there is an instinctive neural response
to infant visual cues that serves as a “parental instinct” and promotes responsive caregiving behavior.
Additional data show specific behavioral responses that may be initiated in response to infant cues.
Behavioral data show that “cute” stimuli narrow our attentional focus (Nittono et al., 2012). Sher-
man, Haidt, and Coan (2009) demonstrated that, after viewing images of cute stimuli, individuals
display more careful behavior on a fine motor dexterity task. Viewing cute infant stimuli may change
our behavior in predictable ways that predispose us to caregiving. Paying focused attention to what
we are doing and being careful are both attributes that would suit responsive caregiving. In response
to infant cries, adults have also been shown to score higher on the popular arcade game “Whack-a-
mole” requiring rapid, coordinated and effortful movement (Parsons, Young, Parsons, et al., 2012).
These findings all point towards the behavioral readiness of adults in response to infant cues, which
could be the result of the neural activity we see in motor regions in response to the same cues.
Summary of temporal dynamics within the parental brain. As outlined in Young et al. (2017), this
temporal framework explains the proposed timeline of neural activity in response to infant cues in
cortical and subcortical regions of the human parental brain. To summarize, specialized processing of
infant cues may originate within the periaqueductal gray of the brainstem, a key “survival structure”
which may trigger adaptive physiological responses, and also act as a rapid subcortical route to engage
broader cortical circuitry. From the PAG, the signal rapidly propagates to sensory cortical regions and
the OFC (within 100–150 ms). These early processes are believed to support salience detection and
subsequent privileged processing of infant cues. OFC activity may “tag” emotionally salient stimuli,
influence ongoing sensory processing, and prime cortical motor reactivity and subsequent behavior.
While sensory processing continues, the OFC may become active at later time points to represent
the ongoing reward value of infant stimuli as the interaction unfolds. Through these processes, it is
Figure 7.4 The pattern of neural activity over time.

Proposed timeline of neural activity in response to infant cues across subcortical and cortical regions of the
human parental brain. Evidence of early detection (<100 ms) of infant cues has been observed in the PAG of the
brainstem. Classical sensory processing ERPs (N100/P100) have been shown to be modulated by some features
of infant facial expressions. At 130ms, there is evidence of salience detection, located in the OFC. Subsequently,
further detailed processing of infant cues occurs in a comparable time frame to the initiation of preparatory
motor responses. Beyond 200 ms, other regions of the parental brain may support responsive caregiving behav-
ior. It should be noted that early contributions of subcortical regions, such as the amygdala and ventral tegmental
area, are less well established due to the greater sensitivity of EEG and MEG to cortical sources of activity, rela-
tive to subcortical sources. However, direct recording from subcortical regions, as in the case of the LFP record-
ings of the PAG, indicate that subcortical regions also contribute to early differential processing.
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clear that infant cues mobilize the brain with great force, allowing for privileged access to neural
mechanisms that in a self-supporting and metastable way is able to ignite motivational states across
the whole brain (Kringelbach and Berridge, 2017).
The Effects of Parenthood on the Brain

Parenting is a transformative experience that redefines how one lives. As an infant grows and devel-
ops greater perceptual and communicative skills, a parent must also adapt and learn, automatically
and unconsciously altering interactions to reflect the infant’s needs. Experience-dependent plasticity
refers to continuous brain development that occurs as a result of a person’s life experiences. As with
any skill, parenting represents a likely modifier of underlying neural circuitry, and there is grow-
ing evidence that parenting experience may provide neural processing advantages for infant cues.
Individual differences in the human parental brain have been investigated in four primary domains:
(1) gender differences in sensitivity to infant cues, (2) effects of parenthood on neural responses, (3)
effects of parenthood on brain structure, and (4) the impact of experience with one’s own infant on
parental responses to own and unfamiliar infant cues.
Gender. One prominent question concerns whether men and women show comparable per-
spicacity with regards to infant cues. Bowlby proposed that children are born with a biological
predisposition to form one exclusive attachment relationship to aid survival, unique to their mother
(Bowlby, 1969). However, the emphasis on the mother as the focus of the monotropy has left a legacy
of inequality in the literature surrounding parent-infant interactions, with fathers typically “kept in
the wings” (Solantaus and Salo, 2005). Fathers bring about positive benefits in child cognitive devel-
opment (Malmberg et al., 2016; Sethna et al., 2017), and paternal involvement in caregiving con-
tinues to increase, especially in middle-socioeconomic families (Yeung, Sandberg, David-Kean, and
Hofferth, 2001). Therefore, a comprehensive understanding of parental sensitivity to infant cues and
effects on both parent-infant interaction and cognitive and socioemotional outcomes must include
both men and women.
Behavioral studies have shown that women tend to display greater overt positive appraisals of
infant facial features, with significantly higher attractiveness ratings than men (Parsons, Young,
Kumari, et al., 2011; Sprengelmeyer et al., 2009). These findings corroborate with the notion that
women may be more emotionally reactive to infant cues than men. However, with recourse to
the hedonic stratification of reward into “liking” and “wanting” responses (Berridge and Robinson,
2003), the findings are more revealing. When the incentive salience of infant cues is measured, by
a simple key-press task where participants control the viewing duration of infant images, men and
women show similar implicit “wanting” responses (Parsons, Young, Kumari et al., 2011). This dis-
crepancy between “liking” and “wanting” for men and women could be due to the differences in
explicit (liking) or implicit (wanting) measures. Men are less conscious of the attractiveness of infant
images, or less willing to admit their appraisals, but they show comparable implicit willingness to
work to view the same images. The reasons for the lower explicit attractiveness ratings in men remain
to be explored, but one possibility is the influence of societal expectations concerning gender roles
in parenting (Lamb, 1975).
Neuroimaging studies have also compared the two genders with regards to responsiveness to
infant cues. Some studies have found no gender differences in response to infant cues (Kringelbach
et al., 2008; Parsons, Young, Mohseni et al., 2013), whereas others have found subtle differences.
There is some evidence of greater early P1 responses in women compared to men when viewing
infant faces (Proverbio et al., 2006, 2011). Cingulate cortex responses may also differentiate women
and men in response to infant cries (De Pisapia et al., 2013; Seifritz et al., 2003). However, there is
scarce evidence to support the notion that women are generally more emotionally reactive and more
“biologically primed” to care for infants.
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Effects of parenthood on neural responses. To explore the effects of parenthood on neural responses
to infant cues, many studies have used a cross-sectional design comparing parents with nonparents.
Generally, results show that parenting experience appears to afford advantages in neural processing
for infant-related stimuli. As an example, compared to nonmothers, mothers have been found to
show greater amplitude of N1 peaks in response to infant crying (Purhonen, Paakkonen, Ypparila,
Lehtonen, and Karhu, 2001) and infant facial expressions (Proverbio et al., 2006). Mothers dem-
onstrate shorter N1 latencies in response to infant facial expressions (Peltola et al., 2014). Moth-
ers also show symmetric N170 responses to infant faces, whereas nonmothers display asymmetric
N170 responses (Noll et al., 2012). Comparing both mothers and fathers to nonparents, parents
have displayed greater N170 and P3a responses to infant faces (Weisman, Feldman, and Goldstein,
2012), with the N170 reflecting face processing and the P3a reflecting attentional processing. Results
are not wholly consistent, however, with some studies finding no effects of parental status on P1
responses (Noll et al., 2012). These findings are also unable to demonstrate whether altered neural
responsivity relates to physiological or behavioral differences.
It is difficult to tease apart whether differences in neural processing are the result of experience
of caregiving or due to pregnancy and neuroendocrine effects. One way to explore this balance
is to compare neural responses to infant cues in biological and adoptive mothers, both of whom
share comparable parenting experience, although adoptive mothers have not undergone recent preg-
nancy. Findings thus far have not been conclusive. One study found no significant differences in
ERP responses to infant faces between biological and adoptive mothers (Grasso, Moser, Dozier, and
Simons, 2009), but another study explored responses to smiling, crying, and neutral facial expres-
sions and found differences in theta power recorded at frontal electrode sites (Hernández-González,
Hidalgo-Aguirre, Guevara, Pérez-Hernández, and Amezcua-Gutiérrez, 2016).
Other studies have explored the duration of time spent caregiving (as a proxy for the amount of
caregiving experience) to see whether the amount of time spent in contact with one’s own infant
has an impact on parental brain responses. One study of fathers found that their childcare experience
was associated with changes in functional connectivity between amygdala and superior temporal
sulcus (Abraham et al., 2014). Parsons, Young, Petersen et al. (2017) found that a longer duration of
motherhood was associated with greater infant-specific activity in key parental brain regions, includ-
ing the orbitofrontal cortex and amygdala. This result supports the idea of experience-dependent
plasticity in the parental brain, an effect that would be amenable to longitudinal studies of parents at
different stages of parenthood.
Effects of parenthood on brain structure. There is also a growing interest in how the experience of
becoming a parent may alter the brain structurally. Pregnancy-induced reductions in grey matter
have been reported in women following the birth of their first child (Hoekzema et al., 2017; Kim,
Leckman, Mayes, Feldman, et al., 2010). These changes have been shown to be stable up to two
years postpartum (Hoekzema et al., 2017). Kim et al. (2014) completed a voxel-based morphometry
(VBM) analysis of structural changes in fathers’ brain during the first four months postpartum. They
specifically looked at the period between 2–4 weeks postpartum compared with 12–16 weeks post-
partum and found both increases and decreases in grey matter volume. Increases were found in the
striatum, amygdala, hypothalamus, subgenual cortex, lateral prefrontal cortex, and superior temporal
gyrus, and decreases were found in the OFC, posterior cingulate cortex, insula, and fusiform gyrus.
However, Hoekzema et al. (2017) who explored grey matter changes in the maternal brain, con-
trolled for the effects of becoming a parent by also scanning first-time fathers prior to and following
their partner’s pregnancy as well as a control group of childless men. They found no changes in grey
matter volume between these groups.
Own vs. other infant studies. The human parental brain is clearly specialized to pick up multimodal
infant cues, but it may also garner expertise in processing one’s own progeny. In the pleasure cycle,
learning is a key component of our interaction with rewarding stimuli and helps us to better predict
270
how to obtain and benefit from rewards. Neuroimaging data have complemented behavioral findings
that adults show enhanced responsivity to their own infants. This “own-infant preference” among
parents may be the consequence of increased learning relating to their infant’s communicative cues,
as factors such as temperament are known to affect hedonic ratings of such cues (Parsons, Young,
Bhandari, et al., 2013).
Both infants and their parents rapidly learn to recognize each other by multimodal cues. Mothers
can accurately recognize their own infant in the early postpartum on the basis of single nonvisual
cues, such as smell, cry, or touch (Cismaresco and Montagner, 1990; Kaitz et al., 1992; Porter et al.,
1983; Russell et al., 1983). Within the first few days and weeks of life, infants demonstrate preference
for their mother’s face (Bushnell, 2001), voice (DeCasper and Fifer, 1980), and even breast milk smell
(Macfarlane, 1975).
Mothers display a differential response to images of their own infant’s face compared to unknown
infant faces in key reward-processing brain regions, including the OFC, anterior cingulate, and insu-
lar cortices (Strathearn et al., 2008). This could be simply an effect of familiarity, as would be
expected for any previously acquainted social partner to drive greater neural responses. However,
unique neural activity in mothers to images of their own infant versus familiar infants has also been
reported in regions relating to reward, such as the OFC (Bartels and Zeki, 2004). These spatial dif-
ferences are suggestive of a progressive attunement of the brain networks involved in processing
personally and biologically salient stimuli.
From the perspective of timing, studies using EEG have looked at the neural responses of moth-
ers to understand when, during processing, mothers differentiate between unfamiliar infants and
their own infants. Repetition priming paradigms have shown that repeated exposures of familiar
faces elicit a larger negative brainwave (N250r) at inferior temporal sites compared to repetitions of
unfamiliar faces (Tanaka, Curran, Porterfield, and Collins, 2006), whereas the P300 appears to be a
novelty or attention marker (Polich, 2009). EEG findings have been mixed. One study demonstrated
no impact of own vs. unknown infant on n170 or P300 responses (Weisman et al., 2012), but two
other studies found differential responding to own infant images from 240 to 500 ms (Grasso et al.,
2009) and at 600 ms (with no effect at N100/N170; Bornstein, Arterberry, and Mash, 2013). Mixed
findings may be due to methodological differences including sample size, time windows used in
analysis, and a lack of analysis of spectral content of EEG signals (Young et al., 2017).
Postnatal Depression
Disruption in parental sensitivity to infant cues is commonly found in parental postnatal depres-
sion (PND), which affects substantial numbers of both mothers and fathers, typically 10%–15%
in high-income countries (Howard et al., 2014; Paulson and Bazemore, 2010) and up to 30% of
mothers in low- and middle-income countries (Parsons, Young, Rochat, et al., 2012). PND can
affect parenting in many ways. It has been associated with altered parent-infant interactions (Bigelow
et al., 2010; Manian and Bornstein, 2009; Stein et al., 2014b), including, as mentioned, disruptions
in parental sensitivity to infant cues (Lester et al., 1995). It can particularly affect cognitive processes,
such as attention and motivation, which are vital for developing parenting capabilities. In terms of
responsiveness to infant signals, for example, both mothers with PND (Donovan, Leavitt, and Walsh,
1998; Stein et al., 2010), and adults with depression (Young et al., 2012; Young, Parsons, Stein, and
Kringelbach, 2015) exhibit disrupted sensitivity to negative stimuli, such as distress in infant cries and
faces. PND has also been associated with increased risk for childhood cognitive and socioemotional
problems (Pearson et al., 2015; van IJzendoorn, Schuengel, and Bakermans-Kranenburg, 1999).
In one study of mothers with depressive symptoms compared to healthy mothers, it was found
that during an observation and empathizing task the depressed mothers showed decreased orbital and
medial prefrontal cortex activity, and a greater reactivity of the right amygdala (Lenzi et al., 2016).
271
The authors interpreted this differential brain activity as demonstrative of an increased internally
focused cognitive style, and an expression of emotional dysregulation. However, the decreased orbital
activity may be an index of a disrupted “parental instinct” as posited by Kringelbach et al. (2008),
with lower coordination of affective cortical resources in response to infant stimuli. Research with
healthy mothers has shown that the duration of motherhood has an incremental effect on the moth-
ers’ neural processing of vocal cues in key regions such as the orbitofrontal cortex and amygdala
(Parsons, Young, Petersen, et al., 2017). In depression, therefore, decreased orbital activity may reflect
strains in parental capacity associated with emotional difficulties.
Higher-Order Capacities and Empathy

Infants have an immense capacity to move us emotionally and spur us to action. Evidence suggests
that infants have the capacity to move us beyond simple caregiving to greater moral emotions such
as empathy. In September 2015, international moral outrage and sympathy emerged in response to
the photograph of a cute 3-year-old Syrian refugee who had drowned in the Mediterranean Sea and
was subsequently photographed on a beach in Greece. Following the release of the tragic photograph,
donations to charities surged and there appeared to be a huge rise in the moral concern for the plight
of the Syrian refugees. While the refugees were previously treated largely with indifference and “dehu-
manized” as an out-group, the plight of the infant ignited hearts and minds and Syrian refugees sud-
denly became part of the in-group and were granted entry in large numbers (Kringelbach et al., 2016).
Parental empathy may be particularly relevant for preverbal infants, as we are required to perceive,
experience, and respond to our infants’ emotional states (Kim, Strathearn, and Swain, 2016). Through
the visual and auditory dimensions of cuteness, infants encourage us to interact with them and treat
them as independent psychological agents (Sherman and Haidt, 2011). The process of mentaliza-
tion involves attributing mental states to others, such as feelings, desires, wishes, attitudes, and goals
(Luyten and Fonagy, 2015). Infant cuteness may act more generally to maximize moral concern by
expanding the moral circle, the boundary drawn around those deemed worthy of moral considera-
tion (Singer, 2011). This idea contrasts with the action of disgust, which has the opposite effect of
contracting the moral circle through contagion processes that extend disgust to categories such as
inanimate objects (Rozin and Fallon, 1987). Disgust often operates towards out-group members,
who are dehumanized as a consequence. Therefore, disgust and cuteness appear to regulate the way
people mentalize objects and involve them within our moral concern, albeit in opposite directions.
The expansion of all infants into our moral circle via alluring infant cues such as cuteness might
also explain why infants are immune from the “other-race effect”, a behavioral finding where peo-
ple are more accurate at recognizing faces from their own ethnicity compared to other ethnicities
(Proverbio and De Gabriele, in press). Here, all infants appear to be processed preferentially, regard-
less of ethnicity. In principle, children are universally forbidden targets of harm, which could stem
from their spontaneous inclusion within our moral circle. Cuteness and infant stimuli may therefore
instigate wider social engagement and perhaps empathy and compassion (Sherman and Haidt, 2011).
We may even be able to harness cuteness to expand our moral circle to other minorities and thereby
minimize dehumanization. Indeed, processing of infant cues has consistently been related to insula
activity (Table 7.1), a brain area known to be involved in the empathic response (Bernhardt and
Singer, 2012). As we currently know very little about how empathy plays a role in our perceptions of
infants, and the neural circuitry involved in such responses, this is an exciting area for future pursuits.
Future Directions in Understanding Parental Neurobiology

We suggest a number of avenues for future research, including (1) expansion of neuroimaging to
other modalities of infant cues and to multimodal infant cues, (2) an increase in the ecological
272
validity of experimental studies, (3) longitudinal studies exploring how parental responsivity changes
with caregiving expertise, (4) an appreciation of the changing family structure, (5) applying compu-
tational methods to the study of the human parental brain, (6) an increased understanding of how the
brain responds in cases of disrupted parenting, and (7) consideration of neuroendocrine involvement
in responsivity to infant cues.
As we have described, infants capture our attention through all of the senses, not just via crying
or visual cues. The majority of studies have used unimodal infant cues corresponding to either static
facial cues or short bursts of infant crying. These studies have formed the basis for understanding the
human parental brain, but the infant cues they use only represent a small fraction of an infant’s com-
municative repertoire. Some studies have begun to use dynamic video stimuli (Noriuchi, Kikuchi,
and Senoo, 2008), which potentially offers more multimodal cues. Incorporation of tactile and olfac-
tory cues represents a considerable challenge, yet these important infant cues cannot be ignored.
A further challenge is to understand how parents come to differentiate the wealth of infant affective
expression. For example, it may be that there exists a hierarchy of infant cues, with those of a negative
valence (e.g., a piercing infant cry) prioritized above positive affective cues.
The majority of studies described in this chapter that explore adult responsivity to infant cues
have involved passive viewing or listening paradigms in socially isolated contexts. Measuring social
behaviors in the absence of a real social partner will always present difficulties. One possible meth-
odology to overcome this problem is “hyperscanning” whereby two (or more) individuals can be
simultaneously measured with high-resolution EEG or fMRI (Astolfi et al., 2011). EEG has the
added bonus of allowing the participants a degree of mobility and also allowing for them to sit
face-to-face.
Understanding how the experience of becoming a parent changes the brain and the process-
ing of infant cues is another major future avenue. To this end, longitudinal studies are needed that
track adults of both genders progressing through parenthood: before, during and after pregnancy.
A combination of neuroimaging, behavioral, and neuroendocrine methodologies would likely prove
illuminating. Future studies may be able to use advances in MRI techniques, such as diffusion tensor
imaging, to examine changes in the connectivity of networks that support parenting. Relating ana-
tomical changes to behavioral changes in responding to infant cues may provide us with more clues
to the biological markers of parental sensitivity.
Family life has changed dramatically, and this change is heavily reflected in the family structure.
Increases in divorce, nonmarital childbearing, and cohabitation have resulted in a larger proportion
of both adults and children living in single-parent families and cohabiting unions (Bumpass and Lu,
2000; Eggebeen, Snyder, and Manning, 1996; Goldstein, 1999). An increasing number of caregivers
now become parents in adolescence (Cavazos-Rehg et al., 2010), while at the other end of the lifes-
pan, grandparents are increasingly taking on more parental duties. Same-sex couples may also have
children, with lesbian, gay, bisexual, and transgender relationships now possessing enhanced social
and legal recognition in the United Kingdom via the Civil Partnership Act 2004 and the Marriage
Act 2013 (Chan, 2012). This diversification of family life outside of traditional cultural norms is an
important consideration for those studying the neurobiology of parenting. Some studies have already
begun to examine diverse parent groups, such as homosexual fathers (Abraham et al., 2014).
Novel methods may pave the way for a greater mechanistic understanding of how infant cues
evoke emotional reactions and relate to caregiving capacities. For instance, advances in whole-brain
computational modeling of human neuroimaging data have made it possible to provide probabilistic
causal information on underlying networks and mechanisms (Cabral, Kringelbach, and Deco, 2014;
Deco and Kringelbach, 2014). Rewards can rapidly become motivational agents with privileged
access to consciousness. Infant cues, in particular, appear to be granted privileged routes to a fast igni-
tion of conscious access. The interplay between prefrontal, limbic, and extrastriate areas can be well
conceptualized within the global neuronal workspace model (Figure 7.5) with higher-order limbic
273
Figure 7.5 Network dynamics supporting global access.

A. Subliminally presented stimuli and nonsurvival-related stimuli often fail to provide ignition of activity that is
made available for global access (Dehaene and Changeux, 2011).
B. Infant survival-related stimuli such as crying provide privileged routes to fast ignition of activity leading to
global access providing the necessary slowness for prosocial caregiving and play behaviors (Chanes and Barrett,
2016; Kringelbach and Rapuano, 2016; Mesulam, 1998).
and cortical regions (e.g., OFC) sending prediction signals to, and receiving prediction-error signals
from, multimodal exteroceptive and interoceptive systems.
Given optimal metastability, fast, privileged activity in regions such as the OFC can start a cas-
cade of neural events that can help sustain the metastable state of pleasure with the necessary slow
processes involved in prosocial caregiving and play. Cases in which parent-infant interaction is com-
promised present a challenge to clinicians. However, using neuroimaging methods we are beginning
to gain a greater insight into how disrupted parenting operates within the brain. On the part of the
parent postpartum depression is prevalent, and on the part of the child certain craniofacial conditions
can compromise parent-infant interaction. Gaining a greater understanding of how these difficulties
translate into compromised interaction is key, followed by intervention trials to mitigate negative
consequences.
Both the oxytocinergic and dopaminergic neuroendocrine systems are believed to play key roles in
the parent-infant relationship (Strathearn, 2011). While oxytocin is primarily involved in social pro-
cesses, the dopaminergic system has a wider involvement in reward processing, reinforcement learning,
and decision making (van IJzendoorn and Bakermans-Kranenburg, 2012; Wigton et al., 2015). The
two systems interact across cortical and subcortical brain regions, and may play key roles in parenting.
Some work has begun to probe how these systems interact with brain activity (Bick, Dozier, Bernard,
Grasso, and Simons, 2013), and further linking them to behavior should prove fruitful.
Conclusions
Our knowledge of the neurobiology of parenting has grown hugely, largely due to the technological
capabilities of neuroimaging. The human parental brain represents a distributed network of regions
that become involved on different timescales and in different capacities. This network overlaps sig-
nificantly with the “social brain” although it is specialized to detect and respond to infant-specific
274
cues, such as infant faces, crying, laughter, and even touch and smell. The orbitofrontal cortex rep-
resents a key node in the network of brain regions that compose the parental brain, supporting early
identification of infant cues for salience, and later processing relating to reward and prediction.
The parent-infant relationship is often inherently rewarding and pleasurable. Such a rewarding
relationship may even promote eudaimonia, the sense of well-being and a life well lived. While neu-
roimaging methodology has grown exponentially, so has our understanding of the human parental
brain, and long may it continue.
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PART II
Ecology of Parenting
8
ANCIENT HISTORY
OF PARENTING
Valerie French
Introduction
For institutions now believed crucial in any society, parenthood and family received remarkably little
attention from historians prior to the early 1960s. Convinced that the adult male world of politics,
war, diplomacy, and economics should command center stage in historical narrative and analysis,
historians gave relatively short shrift to social and cultural history. The 1960s produced many changes
in U.S. society as a whole and in historians’ approaches to the scope and focus of their work. One
result of this revolution within the discipline of history is the burgeoning attention given heretofore
marginalized peoples—women, ethnic groups, middle and lower classes—and institutions—family,
childrearing, community groups, and fraternal organizations. Only because of this revolution in his-
torical interest and inquiry is it now possible to reconstruct the often fragmented history of family
life and parenting.
Historians who struck out in new directions discovered, however, that the old sources and meth-
ods of historical investigation would not work. For modern as well as ancient historians, there is a
problem with the evidence. That evidence was produced and transmitted almost exclusively by elite
males, and it therefore represents one particular perspective on their world and homes. Only rarely
do we hear directly the voices of the women, the children, and the other nonelite members of these
male-elite households describing their own experiences from their own points of view.
Moreover, one cannot turn to the masterworks of political history—Thucydides, Tacitus, Machi-
avelli, von Ranke, Churchill—and find much evidence about family life. The new social historians
had to look elsewhere—in letters, diaries, popular literature, manuals of household advice, medical
treatises, law codes, and artifacts from everyday life. Prior to early modern times, we rarely find
sustained and well-developed discussions about parenting and childrearing in our historical sources.
Ancient and medieval historians, thus, must root about in all kinds of evidence, gathering nuggets
of information wherever they can find them; the picture being reconstructed is not a fully finished
painting, but more a jigsaw puzzle with over half its pieces missing and unrecoverable. From bits and
pieces of details, they then try to discern the overall puzzle pattern.
Reconstructing the ancient history of parenting and family life is beset by difficulties with limited
and undoubtedly biased evidence and by theoretical problems. Do we work inductively or deductively?
If deductively, what general theory or set of questions guides our inquiry? If guided by contemporary
theory, will we misinterpret or overlook evidence? If inductively, how shall we give overall shape and
meaning to the individual puzzle pieces we do find? The method of “deconstruction” offers a way
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Valerie French
to mediate and ameliorate these theoretical difficulties. As a method of interpretation and analysis,
deconstruction takes as its starting point the axiom that the meanings of texts and images are embed-
ded in the beliefs held by society at large and tend to serve the interests of their creators. Thus, the
meanings of texts and images are a social construction, made from a society’s beliefs about what the text or
image represents and, often, interpreted to the advantage of the creator. In most cases, regardless of the
conscious intent of the creator of the text or image, the meanings come from the social construction
of beliefs about the content—both of the society in which the text or image was created and in the
society in which the scholar lives and works.
Something that can be constructed can be deconstructed. And scholars in literature, history, art
history, and other disciplines use the method of deconstruction to illumine the underlying and
inherent social meanings of their sources, whether literary or visual. Although often mystified by
theoreticians, deconstruction requires that scholars ask their evidence and themselves what fundamen-
tal assumptions underlie the categories of analysis and their relations to one another, and how these
assumptions give advantage to the user? For example, to end the conflict between Orestes, who has
killed his mother, and the furies, who torture him in retribution, Aeschylus has Apollo declare, “The
mother is no parent of that which is called/her child, but only the nurse of the new-planted seed/
that grows. The parent is he who mounts. A stranger, she preserves a stranger’s seed” (Eumenides,
658–661). This divine proclamation frees Orestes from the furies, to be sure, and decrees the “natural
order” of things. But just what is portrayed here as “natural”? We should not infer that the Atheni-
ans believed the womb was simply an empty vessel for gestation (there is plenty of evidence to the
contrary). Rather, we must deconstruct the statement and see how it serves male-elite interests as
divine sanction of the male-elite domination of the Athenian family and state, confirmation of the
dominant ideology of male superiority, and male appropriation of woman’s unique contribution
to the family—creation of progeny. Aeschylus is here participating in the Athenians’ construction
of “the natural” and then using this historically specific construction of “nature” to perpetuate the
dominant position of elite males.
The method of deconstruction applies as well to historian as to text or image. This chapter was
written from a post-modernist, feminist perspective. That is, the chapter assumes that most truth is
historically contingent, that some set of values undergirds all forms of inquiry and analysis, and that
elite-adult-male is only one of many kinds of valid, authentic human experiences deserving of care-
ful, disciplined historical study. The goal of this chapter is to illumine as accurately as the sources and
this perspective allow the lives and experiences of all ancient people, particularly parents and young
children.
In addition to presenting scholarly research and interpretation of early childhood, childrearing,
parenting, and family life in several ancient Mediterranean civilizations—Egypt, Mesopotamia, Israel,
Greece, and Rome—this chapter discusses important theoretical considerations—prehistory, patriar-
chy, and maternal values—within the context of this historical description and analysis. Of particular
significance is an overview of how historians have created a rich, new scholarship on the history of
the family and related topics.
History of Childhood, Childrearing, Family, and Parenting

Four categories of inquiry significantly overlap one another. About which one the historian writes
depends mainly on the historian’s focus of attention and position in relation to the subject. All draw
from basically the same pool of evidence, but they write from different angles. The historian of
childhood seeks to reconstruct the experience of childhood, trying to ascertain what it was like to
be a child by seeing the adult world from the child’s perspective; the historian of childrearing stands
back from the child’s perspective and looks for common practices, materials, attitudes, and beliefs that
guided adult care of the young; the family historian is concerned with the material and emotional
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relationships within households and may try to stand either inside or outside of the family; the his-
torian of parenting wants to create a picture of children’s growth and development from the point of
view of the parent, to reconstruct the parents’ experiences and perspectives.
A great deal of the work on children and parents in ancient and medieval societies falls mainly
into the category of history of childrearing, and this work has produced a considerable range of
interpretation. In the 19th and 20th centuries, the few pieces of scholarship to treat children and
families were essays in learned encyclopedias and Daily Life in XYZ books, with obligatory chapters
on mothers and the home. Although often providing basic information, these essays offered little
critical interpretation of evidence and generally painted a benign picture of home life, rather like Fun
With Dick and Jane, a first-grade reader popular in the 1940s and 1950s. The English translation of
Ariès’s (1962) ground-breaking Centuries of Childhood challenged the benign picture and prompted
a variety of scholars to expand on his work. Although Ariès’s contention that childhood as a rec-
ognized stage of development between infancy and young adulthood did not exist before the early
modern era has been overturned (French, 1977), the importance of his positioning childrearing and
childhood as central historical questions can hardly be overestimated.
Hard on the heels of Ariès’s hypothesis came the ontogenic, psychogenic theory of Western his-
tory advanced in the 1970s by deMause, founder of the first journal devoted entirely to the history
of childrearing, The History of Childhood Quarterly. deMause (1974) argued that to understand the
history of the evolution of civilization in the West, one had to examine changes in childrearing prac-
tices over time. In the beginning, deMause contended, childrearing was brutish, cruel, and emotion-
ally distancing because parents were unable to identify psychically with their children’s needs. “The
further back in history one goes, the lower the level of child care, and the more likely children are
to be killed, abandoned, beaten, terrorized, and sexually abused” (deMause, 1974, p. 504). Parenting
in antiquity deMause called the “Infanticidal Mode”; in medieval Europe, it was the “Abandoning
Mode.” Western civilization has progressed, deMause argued, because in each successive era parents
have become psychogenically better able to identify with their young and their needs. deMause’s
monocausal theory explains all adult behavior—political, artistic, social, religious and spiritual,
economic—in terms of the dominant childrearing patterns of its historical era. Today, according to
deMause, we see stages of the “Helping Mode,” in which we recognize that children usually know
what is best for themselves, and as parents we follow our children’s lead in childcare.
Leaving aside the question of whether one accepts deMause’s conclusion that civilization has
truly advanced over the past 3,000 years or his belief that children inherently know what they need,
deMause seriously distorted the historical record he sought to illumine. DeMause’s work, along with
that of a few other scholars such as Etienne (1973), is a striking example of what happens when an
historian presents only one portion of the available evidence. DeMause was correct to point out
multiple examples of horrendous treatment and abuse of children; but he was silent about all the
evidence to the contrary—evidence showing adult knowledge of, attention to, and often delight in
children’s special characteristics and needs.
The work of the last decades on the history of childhood, childrearing, family life, and parenting
is neither benign portraiture nor Hieronymus Bosch–like print. Dozens of scholars have extracted
most of the available evidence from a plethora of potential sources (Bradley, 1991, 1999; Colon and
Colon, 1999; Demand, 1994; Dixon, 1988, 1992; French, 1977, 1987, 1991, 1997, 1999; Golden,
1989; Hallett, 1984; Pomeroy, 1997; Rawson, 1991; Riddle, 1992; Strauss, 1993; Wiedemann, 1989).
Their studies have examined such subjects as contraception and birth control, obstetric and neonatal
care, pediatrics, wet-nursing, foster parentage, children’s legal statuses (freeborn, freed, slave), toys and
play, education, efforts at socialization, affective bonds among family members, and images of moth-
erhood. There is no grand overarching scheme or theme that unites these studies save their common
subject matter and general recognition of the diversity of both children’s and parents’ experiences
within these complex societies and across time.
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Few scholars now working in this field have explicitly considered questions about the vantage
point from which they evaluate and interpret their evidence; they rarely attempt distinctions among
the histories of childhood, childrearing, the family, or parenting. Yet the perspective from which one
writes influences the historical product. This chapter tries to understand and interpret the evidence
about families from the vantage points of the parents within the family; it writes a history of parent-
ing of young children from infancy through ages 6 to 8.
Taking the position of parents has its own complications, for there are many different parents.
In many families in the ancient Mediterranean worlds, the vantage points of mothers differed from
those of fathers, and from those of foster or adoptive parents. In addition, in many homes, parents
were not the primary child caregivers; a variety of forms of surrogate parenthood was common.
Moreover, the expectations of parents differed across socioeconomic class and according to sub-
culture. Thus, while working from the basic perspective of parents, the chapter recognizes that par-
enthood was not a unitary construct, even within a single historical period in a single place such as
pharaonic Egypt, classical Athens, or imperial Rome.
Some Background: Prehistory, Patriarchy, and Maternal Values

Historians rarely venture into the undocumented domain of prehistoric societies, those that left no
written records. However, scholarship on the development of the patriarchal state and attempts to
show that at least some complex pre-literate societies were based on a paradigm of maternal val-
ues compel a brief examination of these issues for they bear heavily on our constructs of “father,”
“mother,” and “parent” and our understanding and use of Jungian psychology and all it implies about
parenting.
In The Creation of Patriarchy, Lerner (1986) traced the origins of the patriarchal state to the
prehistoric transformation of ancient Eastern Mediterranean societies from basically nomadic and
gatherer-hunter groups to settled, agricultural communities that had to defend themselves against
external enemies. The hallmarks of a patriarchal state are the axioms that males are naturally superior
to females, that society must have a hierarchical structure, and that production is more valuable than
reproduction. According to Lerner, the evolution of the idea of individual property rights alongside
the development of male-centered political-military-religious organizations paved the way for the
view that all men (not just members of the elite) have rights in all women and that therefore the
state must control all women. Elite male domination of the state and the creation of patriarchy as
the dominant ideology gave primacy to paternal authority within the family—the rule not just of
fathers, but of the oldest father. It is in the state, not in the family, that we must locate the origin of
“naturalized” male domination and androcentrism (male = norm). Lerner’s thesis has contemporary
significance for parenting as well as political ideology. She demonstrated that the patriarchal state and
family are products of human society and not natural conditions of humanity; we can therefore cre-
ate nonpatriarchal states and families.
Lerner and other scholars who have sought to historicize the origins of the patriarchal state
have assumed some kind of nonpatriarchal prehistoric period. Serious consideration of the possible
prominence of a female or maternal social organizing principle began in the mid-nineteenth century
with Bachofen’s work, a selection of which was published in English as Myth, Religion, and Mother
Right (1967). Based mainly on his analysis of an archaic substratum of Greek and Roman myth that
was filled with powerful female divinities and inferences from the famous “Venus” or “Mother God-
dess” fertility figurines found throughout late Paleolithic and Neolithic Europe and in the Eastern
Mediterranean, Bachofen hypothesized that the first civilizations were matriarchal.
While rejecting Bachofen’s hypothesis of matriarchy as simply the opposite of patriarchy but
with women on top, a number of scholars, including Gimbutas (1982), Eisler (1987), Ehrenberg
(1989), and Baring and Cashford (1991), concluded that prior to patriarchy—with its emphasis on
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militarized hierarchical distribution of power and androcentric axioms—there existed an earlier,

widespread prehistoric civilization based on communal or horizontal organizing principles and rec-
ognition of men and women as different but nonetheless equal in value. The ethos of these cultures is
symbolized in the “Venus” figurines, emphasizing as they do female procreative and nurturing capac-
ities by representing women in the final stage of pregnancy and with engorged breasts. These cultures
that prized maternal values of care and nurturing also valued collaborative activity (as opposed to
competition) and peaceful conflict resolution (as opposed to the use of force).
The burgeoning field of “goddess studies” attempted to weave the archeological evidence from
prehistoric Europe and the Eastern Mediterranean together with historical study of the eradication of
the female divine from Western monotheisms and contemporary feminist theory and feminist spir-
ituality to argue for a radical transformation of society today. The hypothesis that prehistoric societies
were “goddess-centered” and therefore based on maternal, life-giving values was vigorously chal-
lenged by Goodison and Morris (1998) and by Eller (2000). Proponents of the goddess-hypothesis
offered the only systematic and comprehensive interpretation of the archeological evidence; critics
have identified problems in the hypothesis but offered no other interpretation. Given the fragmen-
tary nature of the archeological evidence and the inherent uncertainty of interpreting the meaning
of figurines to the prehistoric peoples who made them, the question of the validity of the goddess-
hypothesis can probably never be settled.
What is of particular importance in this scholarship for historians of parenting is the tendency to
assume that there are inherent male principles and female principles and that the form society takes
depends on which principles it accords primacy. Although there are surely other possible organizing
principles, the debate over prehistory not surprisingly has strong parallels in contemporary philo-
sophical and ideological controversies over women’s and men’s “proper” or “natural” positions in
modern society. These views assume male and female principles splash over into everyone’s under-
standing and interpretation of roles played by mothers and fathers as parents throughout Western
history. The tendency to trace the roots of Western father roles back to the beginnings of the patri-
archal state and to look for the origins of mother roles in prehistoric times poses the real danger that
historians of parenting will impose these gendered parent roles on our sources and either distort or
fail to observe parental behavior that runs counter to or is outside of these parental paradigms. These
same epistemological problems no doubt beset scholars in other disciplines who study parenting.
The field of evolutionary psychology, combined with primatology and anthropological perspec-
tives on the development of human societies, offers perspectives on these issues. For example, Hrdy
(1999) argued that “maternal instinct” is a modern social construction, that in the distant past moth-
ers had to balance conflicting demands and agenda and consequently gave different levels of com-
mitment to each infant depending on the particular circumstances facing the woman at the time
the baby was born. This work also compels us to exercise care lest we impose unthinkingly our own
conceptions of parenting on the millions of mothers and fathers who came before us.
The Ancient Eastern Mediterranean

Traditionally, study of Western history has begun with the Greeks. In reality, as the as the ancient
Greeks themselves clearly realized, European Greek civilization was relatively new and rested on
foundations laid by truly ancient societies in Asia—Anatolia (modern Turkey), Syria, Israel, Assyria,
Babylonia, Akkad, Sumer, and Persia—and Africa—Egypt, Nubia, and the land of Punt (today’s Eri-
trea, Ethiopia, and Somalia). The ancient eastern Mediterranean world was a product of the conflu-
ence of important cultural traditions from three continents. The roots of the European West stretch
back firmly into the soils of ancient Mesopotamia and northeastern Africa.
It was in these ancient Eastern Mediterranean cultures that the system of family and social organi-
zation now called patriarchy was established. Its basic assumptions underlie not only ancient Eastern
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Mediterranean but also Greek and Roman civilization. A brief survey of parenting and family life in
Egypt, Mesopotamia, and Israel suggests the variety of ways parents adapted patriarchy to contingent
historical conditions.
Ancient Egypt
Of all the ancient Mediterranean civilizations, these people—from pharaohs to peasants—seem to
have been, from our contemporary value system, the most devoted parents. Throughout the nearly
2000 years that pharaohs ruled the Nile River valley (ca. 3000–1000 bce), the pervasive social expec-
tation was that parents would have large families, enjoy their children, and rear them with love and
care. Egyptians also recognized stages of child development and had separate hieroglyphs designating
infants, toddlers, youth, and adolescents (Colon and Colon, 1999).
The bulk of the evidence comes from literary and artistic representations of the pharaoh and
his family, and of other nobles, giving our picture a decidedly aristocratic cast. However, archeo-
logical evidence from workers’ communities, particularly Deir el-Medina of the New Kingdom
(ca. 1550–1069 bce), parallels that of elite families. Tomb paintings, from the Old through the New
Kingdoms, presumably depicting what the honoree wanted in the afterlife, frequently show father
and mother surrounded by their offspring, eating, playing, hunting, and so forth. By the conventions
of Egyptian art, the children occupy as important iconographic space as do other relatives and trusted
adult servants.
This interpretation of tomb paintings—that parents regarded their children as essential members
of the family—is corroborated by the prominence given their children in the official art of the
heretic pharaoh Akhenaten and his wife Nefertiti (ca. 1378–1362 bce). King and queen have their
children with them at state ceremonies, hold them on their laps, and kiss and embrace them warmly.
There is no mistaking the fondness of these royal parents for their children—or at least as part of
the public message conveyed by these official representations: conjugal and parental affection is to
be emulated.
Popular stories create the same portrait. In a tale from Middle Kingdom Egypt (ca. 2000–1800
bce), Sinuhe, a royal courtier, returns home after a long trip. Pharaoh and wife rejoice at the reun-
ion; they are joined in their welcome by their children who swarm over Sinuhe, telling him about
their jewelry and toys, as their parents look on fondly. Another Middle Kingdom story, the Tale
of the Shipwrecked Sailor, recounts the adventures of a nonaristocratic, small merchant stranded on
the shores of the land of Punt. Befriended by the Prince of Punt, he finally sets off for Egypt, with the
Prince crying after him, “Farewell, farewell . . . to your home! You will see your children” (Simpson,
1972, p. 55). As a piece of popular literature, this tale probably implies that the audience expected the
poor sailor to have missed his children especially.
Social expectations encouraged large families, with 8–12 children considered a satisfactory num-
ber. Childless marriages were regarded as a disaster; such marriages ended in divorce or with the
adoption of children, often those of poorer relatives. Because marriage occurred early for males—in
their late teens—men had ample time to sire large families; females married in their early to mid-
teens, common in ancient societies.
Fathers were proud of and honored for their progeny, and bureaucrats kept records of all births.
During the 11th dynasty of the Middle Kingdom, an army captain boasted that he had fathered “sev-
enty children, the issue of one wife” (Tyldesley, 1994, pp. 67–68); hyperbolic claims were not uncom-
mon in ancient Egypt. A book of advice to young men from the New Kingdom counseled: “Take
a wife while you are young, so that she may give you a son . . . Happy is the man with a big family.
He is honored on account of his children” (Macdonald, 1999, p. 15). This advice shows the habitual
ancient preference for males over females, but in Egypt the preference was mild in comparison to
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other societies; in Egypt, there is no evidence of female infanticide and few indications of any kind
of deliberate neglect or murder of newborns.
Mothers too took pride in their fertility. In an inscription recording the will of the woman
Naunakhte from the workers’ village of Deir el-Medina (dated to the twentieth dynasty of the New
Kingdom), she proclaims: “I am a free woman of Egypt. I have raised eight children, and have pro-
vided them with everything suitable to their station in life” (Macdonald, 1999, p. 10). The rest of
her will reveals that she disinherited four of her children for not tending to her well enough in her
old age, but that she distributed to all eight a portion of the property she had held in trust for them
from their late father (Robins, 1997, p. 81). Naunakhte’s will is but one of many pieces of evidence
showing the unusually high degree of legal and actual independence of Egyptian women. Custom
dictated that males headed their households, yet women suffered few legal disabilities; they could
own and convey property and could sue in courts in their own right.
Naunakhte’s will also shows the expectation of parents that their children would care for them in
their old age. An adage from the late period (ca. 664–323 bce) advises, “Do not prefer one of your
children above the others; after all, you never know which of them will be kind to you” (Tyldesley,
1994, p. 68). Apparently, not all children lived up to this expectation. But most children probably did.
A New Kingdom scribe’s instructions to his son counsels respect and honor for his mother: “Dou-
ble the food which your mother gave you and support her as she supported you. You were a heavy
burden to her but she did not abandon you. When you were born after your months, she was still
tied to you as her breast was in your mouth for three years. As you grew and your excrement was
disgusting, she was not disgusted” (Tyldesley, 1994, p. 69).
Despite the clear evidence of parental love of and investment in their children, we have relatively
little evidence of childrearing practices. Methods of parenting must be inferred from information
derived from medical treatments, school texts, and household archeology. The extensive medical
literature reveals an astonishing variety of tests used to try to determine the potential fertility of a
woman, whether she was pregnant, and the sex of the fetus; there is little about normal childbirth,
except for medicaments and incantations used to induce birth (Nunn, 1996). Birth was handled by
midwives and was apparently outside the scope of male physicians. While pregnant, women were
protected by the pregnant hippopotamus goddess, Tauret; newborns and young children were pro-
tected by a lion-headed dwarf god, Bes.
At birth, the mother conferred the name on the new baby, presumably with the concurrence of
the father. Names were thought to carry great power, and children were often named for deceased
ancestors who were regarded as continuing members of the family long after their deaths; in some
real way, the child bearing an ancestor’s name also carried that person’s being on into a continuation
of life on earth.
Babies and young children spent their time in the household and their care was entrusted primar-
ily to mothers, elder female relatives living with the family, and older siblings. Babies were laid on
cushions when they were not being carried around by their mothers, who used a sling to keep their
small children close to the breast until they were weaned. In wealthy families, there were additional
child attendants, often slaves purchased especially for this purpose. Elite families had elaborate homes
with elegant furnishings. Working class families had much smaller but well-ventilated houses, usually
with a few rooms and a walled-in courtyard, where most of the day-to-day household work took
place. Thus, children were nearly constantly in the presence of watchful, attentive adults who were
interested in and often delighted by their activities; as they became capable, children were expected
to help with daily chores. Men and women spent their leisure time together, often in family activities
such as picnics or walks through the countryside.
Archeological remains reveal a wide variety of toys: dolls with real hair and articulated limbs, rat-
tles, balls, tops, animal pull toys with moving parts, puppets, board games like chess, and slingshots;
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children apparently also made their own toys out of clay which were then sun-dried. Tomb paintings
depict children at play in games that appear to range from leap-frog, piggy-back riding, to tug-of-
war; holding pet cats, monkeys, and birds; and engaging in such activities as acrobatics, swimming,
fishing, and wrestling or rough-housing. The visual evidence shows small children going naked or
with minimal clothing; some wear sandals and jewelry such as necklaces and bracelets. Most boys
had their heads shaved save for a long lock, often in a braid, on the left side; girls’ hair was cut to
neck length, except for the long side lock they shared with their brothers. Altogether the evidence
strongly suggests that parents provided for their young children toys and activities that helped them
develop physically and socially.
Children were breastfed until about age 3 years—a comparatively late age for weaning. How-
ever, given the contamination of Nile River water—which the Egyptians understood as a potential
source of illness—and the prevalence of barley beer as the main beverage, late weaning makes con-
siderable sense for protecting the well-being of the child. However, given the Egyptians’ desire for
many children and the likely suppression of ovulation associated with breastfeeding, late weaning is
perhaps surprising. Working class and peasant women nursed their own children; elite families hired
wet nurses, a profession held in high esteem among Egyptians, in contrast to other ancient societies.
Otherwise, children probably shared in their parents’ diet—bread, vegetables, lentils, beans, fish, some
fruit, and honey; for working class families, bread was the main stay of the diet, with other foods used
sparingly and meat reserved for feast days. Because of the sand that got into the flour, Egyptian bread
was very gritty; mummies show that even in the elite, teeth were chipped and ground down from
chewing the bread, and all Egyptians, even children, suffered from cavities and gum disease.
Parents, then, created for their children a safe and pleasurable early childhood. But by 5 or 6, chil-
dren were expected to begin preparation for their adult occupations. In all classes, girls were taught
all the domestic skills they would need to manage their households. In working class and peasant
families, the vast bulk of the population, sons learned their fathers’ occupations; training began with
boys following men into the fields or apprenticing in crafts. But here too, parents took great care for
their children’s training for adult life.
More privileged children—some girls as well as boys—attended school from ages 4 to 14, where
they learned reading, writing, math, and singing. A number of school texts from the New Kingdom
survive and give us a glimpse of the values and methods of formal education. Children are reminded
to “write with your hand and read with your mouth”; they could expect beatings for slacking off;
boys are admonished not to pay too much attention to girls (Erman, 1995, p. 189ff ).
Sick babies and children caused their parents much anxiety, as did worry about potential illness.
Infant and child mortality rates were high. The Egyptians had extensive medical knowledge and
advice on treatment ranging from medical to magical, but there was not a specialization in pediat-
rics. The few references to children in the medical literature usually advise incantations and charms.
Maternal milk was employed to treat colic, as were poppy pods which may have conveyed some opi-
ates to the baby; milk was also recommended for eye infections, which were nearly epidemic and a
major cause of blindness. Diuretics and laxatives were often used to treat sick children; they may well
have caused more harm than good.
Examination of the mummies of children shows that they suffered and died from a variety of ail-
ments brought on by worms (round, hook, and flat) as well as other parasites, including schistosoma.
Polio is documented as is an eczema-like skin disease. Without modern anasepsis and understanding
of infectious disease, parents were in fact helpless in the face of life-threatening illnesses (Colon and
Colon, 1999). Children at least had the comfort of loving care and prayers, such as this one intended
to drive away evil spirits: “Perish, you who come in from the dark. You who creep in with your nose
reversed and your face turned back . . . Did you come to kiss the child? I will not allow you to kiss
him” (Tyldesley, 1994, p. 79). In ancient Egypt, even the demon wanted to kiss the child.
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Ancient Mesopotamia
The Tigris-Euphrates River Valley was home to a succession of kingdoms—Sumer, Akkad, Babylo-
nia, and Assyria. Scholars have long noted that in general these Mesopotamian cultures had a darker
cast to them than the apparently more easy-going and optimistic Egyptians: Mesopotamian societies
were more often engaged in war, were more litigious, struggled with comparatively unpredictable
seasonal floods, and had mythologies that depicted a grim life and afterlife. This overall more pes-
simistic tone seems to permeate family life as well.
Literary or artistic representations of a happy home life symbolized by parents with their children
are nearly nonexistent in Mesopotamia. Children are nearly totally absent from official and funerary
art; family life is not celebrated, and it is hardly ever even portrayed. Rather, we find proverbs, like
the following from Sumer, suggesting that, from a father’s point of view, having a family was difficult:
“Who has not supported a wife or child has not borne a leash!” (Kramer, 1963). But having children
was nonetheless important; a barren wife could be divorced, and a wife who refused to have children
could be drowned (Eller, 2000).
The Babylonian law code of Hammurabi (ca. 1750 bce) sets forth many provisions about how
parents and the community were to care for children, particularly their economic upkeep; that many
provisions were needed suggests, perhaps, that the ruler believed that parents had a tendency to shirk
these duties. Dowries were returned upon divorce so that the mother could support her children.
Legally the property of their fathers, children were supposedly protected against physical abuse,
against being reclaimed from adoptive parents, and against being disinherited by adoptive parents
who then had their own natural born children. Again, the existence of these protections implies
some problem with parental ill-treatment of children. But fathers had the legal right to expose infants
and sell their children into slavery. And the principle of the lex talionis in Hammurabi’s code gov-
erned children who were in conflict with their parents. A son who denied that he was the child of
either the man or woman who reared him was to have his tongue cut out; a son who said he hated
the man or the woman who reared him and then went back to his father’s house was to have his eyes
plucked out; a son who struck his father was to have his fingers cut off.
A picture of a desirable home life must be inferred, not from literary or artistic representations of
happy parents with their youngsters, but from descriptions of disasters that could be imposed by the
gods or underworld demons, such as these from Sumerian literature (Kramer, 1963).
The mother will not care for her son, the father will not cry out, O my wife,
The concubine will not rejoice in the lap, the children will not be fondled on their knees.
Take away the wife from the man’s lap, take away the child from the nursemaid’s breast.
Sate not with pleasure the wife’s lap, kiss not the well-fed children,
Take away the man’s son from his knee.
These images suggest that Sumerians lacked confidence about maintaining a tranquil home life;
wife could be ripped away from husband, and children from parents, by forces beyond their influ-
ence, let alone control. This interpretation of such passages finds some support in the etymology of
the Sumerian word for “freedom,” amargi, which means literally, “return to the mother” (Kramer,
1963). First used ca. 2500, amargi’s context indicates that freedom was something valuable. If having
freedom was defined as being able to return to one’s mother, it is quite likely that having freedom
meant being able to be safe, to be protected as a small child would be by her or his mother. Assum-
ing this explanation of amargi’s derivation is sound, then we can probably also infer that in Sumerian
culture, mothers as opposed to fathers were regarded as the main source of protection and comfort
for their children.
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The militarism of Mesopotamian cultures shows up especially in Assyria, certainly the most
fearsome of these powers. Aristocratic fathers introduced their sons early, possibly when they were
3 years old, to the military life by teaching them to ride horses and to shoot with bow and arrow.
The fact that in Assyria the morning greeting involved kisses exchanged among parents and children,
however, suggests some expressions of parental affection for the young.
It was generally believed by the Mesopotamians that demons caused illness. One, Labartu, por-
trayed with a pig nursing at her breast and holding a snake in each hand, was particularly dangerous to
newborns (Colon and Colon, 1999). Surviving medical texts, representing both Akkadian and Assyr-
ian medical knowledge can tell us something about eastern Mediterranean societies’ understanding
of the health and diseases of young children. The Mesopotamians seemed to have recognized such
conditions as scurvy, hydrocephalus, polio, nosebleeds, meningitis, colitis, jaundice, gangrene, epilepsy,
ear infections, tuberculosis, abscesses, dysentery, and poisoning from botulism. Recommended treat-
ments included poultices, bathing, and ingestion of a range of herbs and plants, such as frankincense,
myrrh, and thyme. Only elite families would have been able to engage men learned in medical mat-
ters; the poor surely relied on folk remedies and prayer. How efficacious any of these treatments were,
we have no way of knowing (Colon and Colon, 1999).
Overall it seems that the rather pessimistic outlook of adult society in ancient Mesopotamia may
have served to promote more emotional distance between parents and children than seems to have
existed in ancient Egypt. However, no scholar primarily trained in the ancient Mesopotamian lan-
guages has yet investigated parenting, family life, or childrearing in these complex cultures; research
by specialists may cause a revision of these conclusions.
Ancient Israel
Reconstruction of parenting and childrearing among the ancient Israelites is both important and
difficult. Important because the images of and values attached to family life portrayed in the Hebrew
Bible provide the foundation for the major religions of the Western tradition; difficult because the
Hebrew Bible is organized mainly around a narrative recounting the development of a religion and
nation, written from a patriarchal and androcentric perspective that tends to emphasize public as
opposed to domestic life and concerns. Despite these problems, when the Biblical literature is sup-
plemented with archeological evidence, a reasonable although sketchy portrait can be recovered.
The archeology of ancient Israel in the Biblical period (from the patriarchs through the Baby-
lonian exile, ca. 2000–2500 bce) reveals that the vast bulk of the population lived in an agrarian,
subsistence economy based on small family-centered households. These households had at their core
a husband and wife, their children, and other kin—often elderly relatives—and were usually multi-
generational; they probably averaged 10–15 people (Meyers, 1992).
The number of people and their differing needs seem to have created families that were charac-
terized both by love and affection and by tension and conflict. These complex family relationships
provide the basis for much of the imagery and metaphor of the Bible, particularly that between hus-
band and wife and father and daughter. The authors of the Bible frequently use a metaphor contrast-
ing female infidelity and betrayal with male sorrow, anger, and punishment; the female, representing
the Israelite people or Jerusalem, and the male, representing God, experience discord followed by
reconciliation and renewal of the marital or parental bond, signifying restoration of the covenant
(Ackerman, 1992, on Isaiah; O’Connor, 1992, on Jeremiah). The prevalence of these images and
metaphors surely represents a long-enduring use and reuse of a single literary motif but very prob-
ably also reflects a real and continuing concern with harmony and conflict within the dominant
social and economic unit of the Israelites.
In the Bible, God is often described as a parent, usually as a father; most of these portrayals
show “loving ties between fathers and their children” (Gruber, 1999, p. 142). Sometimes God is
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represented as both mother and father, as in the first creation account (Genesis 1:27) where God
creates both man and woman in a divine image, suggesting therefore a deity who embodies both
male and female (Gruber, 1999, p. 125). Less frequently, a writer such as Isaiah, shows God speaking
or acting as a mother: screaming in childbirth (42:13–14); unable to forget her nursing baby (49:15);
comforting her children (66:13).
In a subsistence, agrarian economy, families profited from having many children to help with the
intensive labor required to survive. “Biblical injunctions to ‘be fruitful and multiply’ surely served the
interests of Israelite villagers and of society as a whole” (Meyers, 1992, p. 248). Most families probably
had 3–5 offspring who survived into adulthood (Gruber, 1999, p. 142); given infant and early child-
hood mortality rates, to produce that number of surviving children, women probably had twice as
many pregnancies (Meyers, 1992, p. 248). Despite the dangers of death in childbirth, women seem to
have wanted large families. Indeed, barrenness—explained as God’s closing of the womb—is a theme
and dilemma that recurs in the stories of the matriarchs, Sarah, Rachel, and Leah (Exum, 1985). And
in the eleventh century, Hannah’s prayer of petition for a child and her prayer of thanksgiving for
her first son provided models repeated in both later Jewish and Christian traditions (Gruber, 1999);
she was rewarded with a son, the future king Saul, and then with three more sons and two daughters
(1 Samuel 1–2; Hackett, 1992). Like other ancient societies, the Israelites explained barrenness and
fertility by divine intervention and regularly prayed for divine favor and protection of the mother
and small children.
Despite the clear evidence for a strong desire for many children, there are hints about older tradi-
tions of child sacrifice, such as the story of Isaac and the ruminations of Jeremiah, which condemn
the practice (7:31–32; 19:5; and 32:35). The date or extent of child sacrifice is unknown (O’Connor,
1992).
Desire for children led some Israelites to adopt orphaned children, as Mordecai adopted his
uncle’s daughter, Esther, in the late sixth century (Esther 2:7). Several other passages depict an ōmēn,
a man who provides nursing care to someone else’s child (Numbers 11:12; Isaiah 49:23). General
community solicitude for parentless children is reflected in Biblical law’s demand that orphans and
widows be invited to holiday feasts (Gruber, 1999).
The Bible, supplemented with later Talmudic literature (dating from the Israelites’ return from
exile in Babylonia in the mid-sixth century bce, extending into the Hellenistic and Greco-Roman
periods, and lasting into the fifth century ce), provides some evidence about the beginning of human
life among ancient Israelites. Pregnant women were to avoid alcohol, suggesting an understand-
ing of the possibility of damaging a fetus in utero (Colon and Colon, 1999), and pregnant women
were excused from fasting. Miscarriages were explained by strife and stress in the home, strong and
unpleasant odors, the experience of great pain, or insufficient food. Childbirth was expected to be
painful and explained by Genesis’s account of God’s punishment for Eve’s disobedience. The death
of a woman in childbirth was attributed to her violation of religious law or breaking a vow; an easy
delivery was seen as a reward for righteousness (Ilan, 1996).
Childbirth was superintended by midwives and accomplished by kneeling on bricks or stools.
Midwives could exercise considerable authority to judge from the Exodus account of Shipreh and
Puah, who rejected pharaoh’s order to kill all newborn boys at birth; their refusal saved Moses and
started the Hebrew resistance that culminated in the Hebrews’ escape from Egypt (Exodus 1:15–19).
The twelfth-century story of the wife of Phinehas giving birth during a battle against the Phil-
istines shows midwives comforting the new mother with the news, “Do not be afraid, for you have
given birth to a son” (1 Samuel 4:19–22), suggesting a preference for male over female children, or
perhaps a need for sons to grow up to become soldiers. Preference for male children during the Bib-
lical period, however, may be overstated by the Bible; in a labor-intensive agrarian economy where
many children are desired, females provided not just workers but crucial skilled labor (Meyers, 1992).
But almost certainly a preference for males was clear by the more urban and commercial Talmudic era,
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when the birth of a daughter became a disappointment. Nonetheless, Jews living in the Greco-
Roman world reared all children, and fathers were required to support both sons and daughters
while they were minors (Ilan, 1996).
In a metaphor describing God’s care of newborn Jerusalem, Ezekiel (16:4–12) provides a good
description of the treatment of neonates. The umbilical cord was cut, the infant washed first with
water, then oiled and salted with soda ash, which likely served as a bacteriostatic astringent, and
finally wrapped in cloth. Evidence from Talmudic sources reveals a variety of remedies for babies
who did not breathe readily and for some birth defects, showing “a keen observation of congenital
abnormalities and normal newborn behavior” (Colon and Colon, 1999, p. 29). In Ezekiel’s account,
God provides his infant daughter with the finest embroidered fabrics of linen and silk, sandals made
of badger skin, and jewelry—bracelets, a necklace, earrings, and a tiara; this portion of the story of
divine care reflects, no doubt, the practices of a few wealthy families; such luxuries were simply una-
vailable for the bulk of the population. But these unusually lavish infant accoutrements probably do
reflect a general social expectation for basic material investment in the care of newborns.
Infants were named at birth, often for parents or grandparents. The name was usually given by
both parents, but could be bestowed by the mother alone, as shown when the wife of Phinehas by
herself named her newborn son Ichabod. Circumcision occurred on the eighth day and was carried
out by a man, although Exodus 4:24–26 hints that women may have played a role in this ritual in
very early times.
Mothers, together with older women and siblings, provided the bulk of the care of young chil-
dren. In Biblical times, the demands for women’s skilled labor in the subsistence, agrarian economy—
gardening, transformation of agricultural products into food, weaving and sewing, pot-making and
basketry—probably consumed an average of 10 hours a day. In such circumstances, mothers likely
integrated childcare into their other responsibilities (Meyers, 1989, 1992). Thus, we must picture
children growing up under the watchful eyes of mothers and adults but also having considerable
freedom of movement and action so long as they did not disrupt the extensive and necessary work
of adults. We hear nothing about small children’s toys or games or play. But some time for maternal
play with small children is suggested by Isaiah’s description of a restored Jerusalem as a happy child
“carried on her hip and bounced on her knees” (66:12).
The most frequent references to early childcare in the Bible concern nursing and weaning,
which probably took place around the age of 3 years (Meyers, 1988). The story of Ruth, from the
fourteenth century, shows us an infant left in the care of a grandmother while the mother went
to work in the fields (4:17). But the story of Hannah, from the twelfth century, presents a nursing
mother who is excused from making the Passover pilgrimage so she can stay home with her infant
until he is weaned (1 Samuel 1–2). In Biblical times, mothers very probably nursed their own babies,
although a wet nurse is mentioned in 2 Samuel (4:4). In the later Talmudic period, wet nurses were
used in some elite families, as they were in wealthy Greco-Roman ones. Rabbinic opinion was
divided on the practice of wet nursing, as it was on the appropriate time for weaning; there was
agreement that the minimum age should be 18 to 24 months, but recommendations for the maxi-
mum age ranged from 2 to 5 years. Nursing mothers were encouraged to suckle their children with
such incentives as a reduction in other household duties, an exemption from fasts, and the right to
have intercourse and use contraceptives while nursing (Ilan, 1996).
Another common reference to young children in the Bible depicts their suffering in war, and the
history of the Israelites brought them much strife—both internal struggles and conflict with their
aggressive neighbors. Indeed, the suffering must often have been terrible. Writing of events in the
sixth century, Jeremiah tells how a mother, once honored and blessed for her seven children, is cursed
by losing all of them in war (15:9). And Jeremiah’s prophesy that Jerusalem will endure so horrible
a famine that parents would eat their children (19:9) seems fulfilled in Lamentations where Jeremiah
says the children are desolate (1:16), nursing children and toddlers faint from hunger on the streets
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(2:11; 2:19), children’s tongues stick to roofs of their mouths due to excessive thirst (4:4), and moth-
ers eat their own young (2:20; 4:10).
War and upheaval were regular features of Israelite history, but more peaceful images of daily fam-
ily life appear as well, if less frequently. A mother’s voice in Psalm 131 tells the Lord as she approaches
a place of worship that she has quieted her soul; it is calm, “like the contented child I carry” (131:2).
Involvement of children in family-centered religious activities, including those the central authorities
regarded as acts of apostasy, was probably a regular feature of family life, such as the scene described
by Jeremiah: “the children gather wood, the fathers kindle fire, and the women knead dough, to
make cakes for the queen of heaven” (7:18); the queen of heaven here may refer to the fertility god-
desses so common in the eastern Mediterranean (O’Connor, 1992) or to an earlier Hebraic concep-
tualization of the divinity as both king and consort (Fyrmer-Kensky, 1992).
Other evidence suggests sensible, healthy childrearing practices. Parents were advised to keep
their youngsters out of the sun during the middle of the day. Children were to eat slowly, chew their
food well, and have a good, solid breakfast (Colon and Colon, 1999). Like their parents, children
probably lived on a diet in which the mainstays were bread, olives and olive oil, grapes, and dairy
products; these principal foods were supplemented seasonally with other fruits, legumes, and vegeta-
bles. Meat was probably reserved for feast days. Strict dietary laws and unusual attention to sanitation
in food preparation and handling may well have reduced the incidence of food-borne disease among
Israelite children (Colon and Colon, 1999).
Primary care of infants and toddlers was the responsibility of the mother and older women, but
the Bible clearly reflects an expectation of shared parental authority. Law required respect for and
obedience to both mother and father. Besides the commandment to honor both mother and father
(Exodus 20:12; Deuteronomy 5:16; Leviticus 19:3), Exodus demanded the death penalty for anyone
who cursed or assaulted either parent (21:15–17; Leviticus 20:9). Two injunctions in Deuteronomy
required actions by both mother and father for grown children: turning over a juvenile delinquent
to authorities and displaying the bloodstained marital sheet of any bride wrongly accused by her
husband of not having been a virgin (Deuteronomy 21:18–21 and 22:15).
Traditional wisdom expressed in Proverbs shows the role both parents played in educating the
young: “Hear, my child, your father’s instruction, and reject not your mother’s teaching” (1:8 and
6:20). Parents taught children the tasks they would have to carry out as adults: fathers the heavy work
of the fields, mothers the more specialized labor of tending the gardens, transforming crops into food,
weaving, and sewing. Because both boys and girls spent more time during their formative years with
their mothers and elder females, it seems probable that these maternal figures played the greater role
in socializing the young into the values and mores of Israelite society and religion (Meyers, 1988).
Mothers had a duty to teach their daughters to keen and wail at funerals ( Jeremiah 9:19).
The authority and power of mothers during the Biblical period distinguishes the Hebrews from
other eastern Mediterranean cultures. While the Bible mentions women infrequently (only 9% of
the named people are female; Meyers, 1992) and mainly as the mothers of important and famous
sons, these women often had significant influence in starting and determining the outcome of crucial
events (e.g., the matriarchs, Miriam, Bathsheba, Jezebel, Delilah). One of the most powerful women
in the Bible, Deborah, was a judge who organized and helped to lead an army that freed the Israel-
ites from Canaanite oppression, probably in the twelfth century. The name of her husband is given,
but there is no specific mention of her children. Yet in her victory song, one of the oldest pieces of
literature in the Bible, Deborah identified herself first as a “mother in Israel” ( Judges 4–5). Inferences
from her accomplishments suggest that “a mother in Israel is one who brings liberation from oppres-
sion, provides protection, and ensures the well-being and security of her people” (Exum, 1985, p. 85).
There is every reason to believe that within Israelite households of the Biblical period, moth-
ers exercised authority alongside fathers and served as empowering role models for their daughters.
But the legal and cultural authority granted to mothers in the Biblical period was significantly
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circumscribed by the Talmudic era. In rabbinic literature, mothers have lost their ability to be a
guardian of or agent for their minor children, and with respect to “legal responsibility, women were
usually bracketed with slaves, minors, deaf-mutes, and persons of double or doubtful sex” (Archer,
1994, p. 63).
Many scholars believe that during much of the Biblical period, ordinary Israelites had rudi-
mentary literacy—in sharp contrast to other eastern Mediterranean cultures. Because there is no
evidence of formal schools with paid instructors prior to the Talmudic period, it seems likely that
parents, both mothers and fathers, taught their children the basic elements of reading, writing, and
numeracy, as well as the fundamentals of Hebrew cultic and family law (Meyers, 1988). Law required
fathers to instruct their children in Hebrew history and tradition (Exodus 10:2; Deuteronomy 32:7).
Eastern Mediterranean Civilizations: Some Conclusions

In these societies, it seems clear that the overall outlook and presuppositions of the general culture
permeated to some degree attitudes about children and parenting. Although all of these civilizations
recognized infancy and early childhood as distinctive stages of life and took care to safeguard young
children, the experience of parenting—and of being a child—seems to have been decidedly differ-
ent in each. Because of their particular history during the Biblical period, the Israelites seem to have
blended into their beliefs about parenting elements of the more harsh and pessimistic Mesopotamian
cultures with the more affectionate and optimistic outlook of the Egyptians, combining both with
the Israelites’ unique conceptualization of their relationship with their God.
The Greeks
For the Greeks, there is considerably more historical evidence for investigating parenting, family life,
and childrearing practices. As in the ancient Eastern Mediterranean, patriarchy dominated the way
Greeks thought about these critical issues. Most of the evidence for ancient Greece was produced
in Athens, and an inevitable Athenian elite male bias colors the sources. What historians reconstruct
about Greek parenting pertains primarily to the Athenian upper classes. Inferences about lower
classes, slaves, and other Greek communities are difficult and uncertain. However, some important
differences between Spartan and Athenian parenting can be seen.
Unlike most of their ancient predecessors in the Eastern Mediterranean, the Greeks thought con-
sciously about thinking and knowing, systemizing their knowledge of the human and natural worlds.
Although few Greeks focused their writing on parenting and childrearing, political philosophers saw
parenting and childrearing as crucial ingredients in creating and maintaining the ideal state. Greeks
were observant and knowledgeable about stages of child development and thoughtful about how
parents should respond to the child’s changing needs—not so much for the sake of the child, but for
the welfare of the community.
Polis and Oikos: The Cultural Context of Greek Family Life

The Greek polis or city-state was the fundamental unit of Greek political life. Most were governed
by relatively small groups of families who called themselves aristoi—the best men; because they held
the power, kratos, they became known as aristocrats. For our purposes, it is important to note that
these aristocrats used the household, oikos, as their principal conceptual model for the polis. For the
Greeks, the aristocratic household embraced a good deal more than the family and its basic dwelling;
the oikos included family dependents (e.g., servants and slaves; other people, usually kinfolks, who
were wards; in-laws of lesser socioeconomic status) and all property (e.g., city house as well as coun-
try estates). At the head of each household was the kurios, the senior male direct descendant of the
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former head. Although the kurios in theory had primary power, in fact he was expected to consult
widely with other senior males; collectively the senior males looked out for the social, economic,
and political interests of the family with the kurios serving as their major spokesman. In less affluent
families, the oikos was, of course, much smaller, corresponding more nearly to a nuclear family that
rents its dwelling place. But regardless of wealth and size, every oikos had its kurios.
Within the oikos, especially the more wealthy ones, there arose a fairly strict division of labor
between men and women and a clear demarcation between public and private spheres of action.
The world of the polis was the world of men; the activities within the oikos as a domestic and fam-
ily unit was a space for both men and women. The women supervised and performed domestic
duties—cooking, cleaning, weaving, and childrearing—but were nonetheless accountable for their
work and proper behavior to the kurios. In practice, daily life saw men of affluence moving freely
and frequently back and forth between oikos and polis; affluent women were mainly confined to the
oikos because the family could afford to have servants and slaves go out to get water and conduct the
family’s daily commerce.
Recent work on the domestic architecture of ancient Greece allows us to infer some important
ideas about the interaction of adults and children within the Greek house. The focal point of domes-
tic life was a walled-in but open courtyard with a portico for shade in summer and protection from
rain in winter. Paved in more affluent houses, packed earth in less, the portico and courtyard com-
plex was where most activities took place. Small, often windowless rooms—including a washroom
and kitchen or cooking area—were reached from the portico; only these small rooms were part of
the oikos’s truly private space. Almost always directly adjacent the house’s main doorway, the court-
yard served as a kind of intermediate ground between the private oikos and the public polis in that
guests, both male and female, were received here. From the courtyard, there were usually clear lines
of sight into the rest of the house’s rooms. The only interior room that may have been reserved for
exclusively male use was the andron, a dining area used by the male family members for entertaining
other male guests—and sometimes female entertainers—at dinner; although it is certainly possible
that during the day this room could have been used by other members of the family (Levett, 1999).
From the architecture of Greek houses, we can infer that children constantly interacted with adults
in the household, observed them at work, participated in both daily family religious rituals and other
significant events such as weddings and funerals, met most guests, and had ample space to play and
exercise. In turn, parents and other adults could keep a constant eye on the young.
For less affluent families, the basic house plan was the same, just smaller. Families that ran artisan
businesses—leatherworks, masonry, pottery—generally had their workshops immediately adjacent
to the house; it is not unreasonable to infer that children would have gone back and forth between
house and workshop, perhaps even helping out with the business. Because by law, Athenian fathers
were required either to train their sons in a trade or to provide them with some means of sup-
port, we can safely assume that sons of artisans and laborers began some sort of apprenticeship
with their fathers at an early age. A large proportion of the male citizens earned a meager living as
day-laborers—carpenters, agricultural workers, and ox-drivers. Without domestic servants and with
their husbands otherwise fully engaged, women in these families had to go into the public sphere to
purchase food and household items; indeed, many women from the lower classes also worked outside
their homes for pay as laundresses, midwives, nurses, fishmongers, ribbon sellers, and so forth. Who
looked after children while their parents were away running errands or working is unclear; most
likely it would have been an older relative living in the household, although it is certainly probable
that children from poor families were left unattended for considerable periods of time.
It is into this social, economic, political, and physical context that we put our investigation of
Greek parenting and family life. Only the wealthy elite families had the wherewithal to achieve
the aristocratic ideal of the oikos, but it is likely that less affluent families aspired to the same ideal.
In classical Athens, even among aristocrats, there arose considerable division of opinion about how
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far into the oikos the reach of the polis should extend. And when the state tried to reach into the
home, it was often about matters of childrearing and sexual activities (particularly relations among
men). The more radical democrats drew the line at the front door, claiming a right to privacy in
childrearing and sexual conduct much like that contemporary jurists have found in the U.S. Con-
stitution. The conservative aristocratic families, however, alarmed at the demise of the old, highly
disciplined approaches to childrearing and the erosion of deference to their position, advocated
state-imposed schemes of childrearing that began at birth and continued into early adulthood
(Cohen, 1991; Keuls, 1985).
Reckoning with this aristocratic conservative perspective, especially in Athens, is crucial to inter-
preting parenting, for its major spokesmen—Plato and Aristotle—provide us with the bulk of our
information about Greek views of expectations of parents and child development. It is often difficult
to separate what parents were actually doing from what Plato and Aristotle say parents should be
doing.
The importance these philosophers attached to children and childrearing seems not much differ-
ent from that found elsewhere in Greek society. A household needed children to be complete; a son
was critical for continuing the family line into future generations; at least one daughter was desirable
to care for parents in their old age; and children were needed to tend and honor the family graves.
Leaving an enemy childless, as Medea does her abandoning husband Jason, was the most devastating
revenge possible. These reasons for producing and rearing are fairly devoid of sentimental feelings
about small children themselves, and that is an important difference between ancient and contempo-
rary Western attitudes about children. Nonetheless, Greeks seem to have regarded the experience of
parenting with interest and felt considerable affection for their offspring qua children.
Family Planning
The birth of a child was an important family event, and the child’s formal acceptance into the house-
hold and naming involved considerable ritual and celebration. A collateral issue here is the extent to
which the Greeks practiced infanticide. That infanticide—as we define it, taking away the life of a
newborn baby—was a legal option that was exercised by the kurios is indubitably true. The vexing
historical questions concern how often Greek fathers resorted to infanticide and whether females
were more likely to be exposed than males.
In considering infanticide, we must remember that an infant did not become a legal person until
it was officially accepted into the family; the act of birth itself did not confer legal status on the baby.
The distinction was important to the Greeks. By exposing the child—wrapping it warmly and leav-
ing birth tokens with it—the parents left its fate to the gods; some exposed newborns were rescued
and reared by childless families. Thus, exposure did not legally constitute murder, and newborns
seem to have been conceptualized as being more akin to an unborn fetus than a full person. Given
high neonatal mortality rates, it is not difficult to understand why ancient families waited a few days
to determine whether the baby would survive; formal acceptance and naming occurred about a
week after childbirth. However, Pomeroy (1997) suggested that the delay in performing the ritual
of acceptance may well have diminished the initial strength of parental bonding with the newborn,
creating a more distanced affective tie for both baby and parents.
Many of the crude ancient pots found filled with the bones of newborns—evidence adduced
in favor of significant infanticide—are more likely burials of infants who died from natural causes;
because they were not yet formal members of the family, the disposal of their bodies did not require
the usual funerals and rituals of internment, and they would not have been put in cemeteries. How-
ever often fathers resorted to infanticide, it was not a casual decision, and adults felt real unease about
the act, as demonstrated by ambivalence expressed in literary texts and the hope that exposed infants
would be rescued and reared by another family. There is no consensus among historians about selective
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exposure of females. Despite some work (Riddle, 1992) arguing that adult sex ratios favoring men
do not demand extensive female infanticide as an explanation, most scholars (e.g., Pomeroy, 1997)
maintain that more girls than boys were killed, a reflection of the general social preference for males.
The weight of the evidence and argumentation now suggests that infanticide in antiquity was a
relatively uncommon method of controlling family size. Indeed, it appears that ancient families regu-
lated their size through a variety of methods of contraception and abortion. Most of our evidence
about contraception and abortion comes from sources later than the classical Greek era, but the
information contained therein most likely reflects centuries-old traditional herbal medicine. Riddle
(1992), historian of ancient medicine and particularly herbals, conducted an extensive study in which
he concluded that ancient people used contraceptives and abortifacients for effective planned parent-
hood, regulating with considerable success the number and spacing of their children.
The Social Construction of Children and Childhood

In general, Greeks saw their offspring as inheriting both physical and psychological characteristics of
their parents and ancestors (Pomeroy, 1997). If there were various views about the relative strengths
of male versus female contributions to conception and gestation, there was little doubt that both the
mother’s and father’s families were somehow represented in the children. Adults expected to find
both physical resemblances between children and progenitors and personality and character similari-
ties as well. Parental expectations here may have significantly influenced their behavior towards their
young.
The Athenians had a well-developed social construct of what children were and how they dif-
fered from adults and old people. Above all, Greek parents conceived of their young as plastic, shapea-
ble, unformed, impressionable, ignorant, gullible, imaginative—a physical as well as mental/emotional
tabula rasa. The idea that children were plastic included the child’s body as well as the mind. Thus,
the application of the proper nurture and training was seen as critical in determining what kind of
adults children would grow up to be. Collaterally, adults saw children as sometimes unruly, difficult
to control, in need of discipline.
Judging from the frequency of remarks in the corpus of Greek literature from the classical period,
second to the adult view of the child as plastic comes a recognition of the child as a fearful creature,
as easily frightened, given to tears, hiding in mother’s skirts, begging to be picked up for comfort.
Greeks believed their children would be frightened by strangers, objects of large size, or changes in
the family’s circumstances. Parents and other adults apparently took advantage of (and perhaps even
cultivated) this characteristic by telling stories about ghosts and monsters that would punish children
for bad behavior. In addition to these characteristics, the Greek construct of children saw them as
loving, happy, playful, and affectionate; as naturally imitative; and as innocent, unworldly, and without
sexuality (French, 1991).
Not surprisingly, Greek parents also saw children as helpless, in need of protection, unable to
communicate. One adjective frequently associated with small children in Homer is nepios, meaning
literally “not able to speak.” Babies’ inability to express their needs clearly was apparently a source of
considerable discomfort to Greek parents. In Aeschylus’s play, The Libation Bearers, Cilissa, Orestes’
aged nurse, recalls caring for her young charge: “A baby is like a beast, it does not think/but you
have to nurse it, do you not, the way it wants./For the child still in swaddling clothes cannot tell us/if
he is hungry or thirsty, if he needs to make/water. Children’s young insides are a law to themselves”
(753–757). This recognition of infancy as a distinct phase of life during which a child is in need of
special care shows up in medical writings, particularly with respect to the child’s greater vulnerability
to disease and infection.
Plato and Aristotle on children’s development. Writing in the fourth century bce, Plato and Aristotle
undoubtedly drew a good deal of their knowledge about children and parenting from the larger
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society; neither seems to have devoted himself to empirical studies of the subject. It is important to
recognize how much knowledge about children and childrearing was generally available to these
wealthy male aristocrats; the scope and depth of that knowledge strongly suggest that Greek fathers
were not the absentee, disinterested parents some scholars have posited. Of particular interest is the
developmental approach both take to describing the stages of childhood. Table 8.1 sets out their
principal observations about the different characteristics or needs children have as they grow up and
their prescriptions for the appropriate or proper care for that stage. Plato’s prescriptions come from
Laws, 694D and 789E–795E; Aristotle’s come from Politics 1336a–b; the views they set out in these
passages are echoed and supplemented in other of their works.
The stages that Plato and Aristotle created correlate reasonably well with many contemporary
theories of development, missing only is what today we call toddlerhood. And both Plato and
Aristotle included psychosocial as well as physiological growth. Neither philosopher believed that
his scheme fit all children uniformly. Both commented on the importance of recognizing differ-
ences among individual children and advised tailoring childrearing to suit the particular child. We
can find a good deal of evidence in other sources to confirm that these developmental stages were
widely accepted and that the generally mild, nurturing parenting practices recommended were also
generally agreed upon. Indeed, these other sources allow us to add a good deal of detail to the stages
implicit in Plato’s and Aristotle’s generic discussions.
Childbirth and infancy. Demand’s (1994) Birth, Death, and Motherhood in Classical Greece provides
a comprehensive study of this important topic. Demand accepts arguments that Athenian citizen
women probably underwent six or more pregnancies; the number of pregnancies for women in
other city-states may have been lower, in the range of three to five. Ancient Greeks were well aware
of the dangers of reproduction for mother and child. We will not be far off the mark if we assume
that ancient Greece had maternal and infant mortality rates comparable to those of contemporary
cultures lacking modern medical care, particularly anasepsis. There were three major incentives for a
woman to undergo so many pregnancies. First was the social belief that childbearing was a woman’s
Table 8.1 Characteristics and/or proper care
Age Level Plato Aristotle
Birth to important weaning Swaddled, much rocking and crooning Bodily development most important
(about training 2 years) Carried by nurses “Training of the body [comes]
before training of the mind.”
2–3 years Nurses/mothers ascertain wants Plenty of milk; no wine
according to cries As free movement of the limbs as
Shield from pain, fear, grief, corruption possible
“It is infancy that the whole character is Accustom to endure cold
most effectually determined.”
2–3 to 5–6 years Games played together with other Bodily exercise
children Play in preparation for adult
Mild discipline activities
Allow to cry
Protect from base influences
Keep at home
5–6 to puberty Separate the sexes Boys begin school
Boys go to school; learn gymnastics and Avoid hard diet and severe exercise
music
Girls learn gymnastics also
Puberty to 21 Rigorous training for boys Rigorous training for boys
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primary purpose in life and within the family; a woman with few children had less prestige than a
woman with many children. Second was production of many children so that enough of them would
survive to adulthood to sustain the family. Third was the belief that frequent pregnancy protected
women from the dreaded illness of hysteria, the wandering womb; without the weighty fluids pro-
duced by pregnancy, a dried up womb could break free from its moorings and move about the body,
causing terrible medical and emotional problems.
Demand’s extensive analysis of the Hippocratic writings on obstetrics (some 40 individual cases)
and of related sources reveals the social construction of childbirth in ancient Greece, particularly
Athens. Care of the pregnant wife, supervision of childbirth, and care of the infant were regarded as
the province of women, particularly of midwives. The difficulty of successful obstetrical work was
recognized by the Hippocratic doctors: “It requires much care and knowledge to carry and nurture
the child in the womb and to bring it to birth” (Hippocrates, Diseases of Women I.25). A male physi-
cian was called upon only for advice on handling complications during pregnancy and delivery—and
only when the kurios was able and willing to pay the doctor’s fees.
Assuming survival of infant and mother, the family celebrated the amphidromia (literally “the run-
ning around”) on the fifth or seventh day after birth, decorating the front door with an olive wreath
for a boy and wool for a girl. On this occasion the father carried the newborn around the hearth
signaling his intention to accept it into the family and rear it; the mother and female kin who had
attended the birth purified themselves; friends and neighbors sent gifts and congratulations. Families
who could afford a second celebration for family and friends officially named the new child on the
tenth day.
The day-to-day care of infants was the preserve of the women in the household. An experienced
mother would expect and be expected to supervise this care. However, with the notable exception
of Sparta, Greek girls became wives and mothers in their early teens and needed the guidance and
support of their older female kin, both those of their natal families and those of their in-laws—
particularly the women resident in the household. On the whole, we can infer a pattern of baby care
that involved cooperative efforts among several women. In wealthy families, there would probably be
a wet nurse and perhaps other nurses hired or purchased for their expertise in tending small babies.
And the mid-wife remained on call for postpartum and pediatric problems.
Swaddling was common among the Greeks, again with the exception of Sparta whose example
probably led Aristotle to recommend that babies have free movement of their arms and legs. Les-
sons about swaddling in later writers make it clear that adults believed the bands were necessary to
insure the growth of straight limbs, a reflection of their construct of the child as plastic. Along with
swaddling went several daily massages, whenever the bands were removed to clean the infant. Greek
practice apparently loosed babies from their bands gradually, first leaving the head free, then the arms,
and so on down the body; babies were completely out of swaddling probably by the time they were
6 months old.
Breastfeeding was regarded as the best nourishment, but success here seems not to have been uni-
versal. We can assume that some very young mothers did not produce enough milk and that some
upper-class women preferred to use wet nurses. At any rate, historical sources attest to wet nursing as
a common phenomenon and a means of livelihood for many lower-class women. Many infant feed-
ing bottles from excavations across Greece indicate that human milk was often supplemented with
other foods such as goat’s milk, honey water, diluted wine, and strained broth from soups. After a few
months, babies were probably introduced to bread soaked in one of these liquids and thin gruel or
porridge. Older babies might have mashed vegetables with olive oil and bits of meat that had been
masticated by mother or nurse. Full weaning probably took place by a child’s second birthday at the
latest.
Plato attached special importance to infancy. He argued that it is through its perceptions of pleas-
ure and pain that an infant first begins to gain a conception of good and evil, of right and wrong—a
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kind of proto-behaviorist theory of human motivation. Plato complained that parents were generally
not careful enough to make sure that babies found pleasure in people or activities that should be
emulated. He criticized, for example, leaving infant care (Plato surely is thinking here of aristocratic
children) to lower-class or slave nurses for two reasons. First, he contended, they probably do not do
their work as well as the real (high-born) mother would; second, if they did a good job, the child
would associate pleasure with them—people of low status—not with their aristocratic parents.
Mothers and mother surrogates keep nearly constant watch over the infant during the first year of
life. Crying was generally interpreted as meaning the baby needed something—as suggested by the
nurse Cilissa’s observations in Libation Bearers about the young Orestes; she implies that babies cry to
indicate that they are hungry, thirsty, or need to urinate. We can assume that collectively the infant’s
“mothers” provided a great deal of stimulation through their attention and ministrations. During a
child’s first year of life then, a pattern of attentive, gentle female parenting was well established. At
what point fathers began to play more than observer roles in their children’s life is unclear; but their
close observation of infancy is certainly well-attested in our predominately elite male literary sources.
Toddlers and preschoolers. The dominance of females in parenting is clear not only for infants but
also for male children before they went to school and for females until they married and left home.
But there is good evidence suggesting that fathers began to interact with their youngsters once they
became toddlers. Several passages from Aristophanes’s comedy, The Clouds, portray one middle-class
father, married to an upper-class wife, as intimately involved in his young son’s daily care. The play’s
hapless hero, Strepsiades, reminds his now arrogant and rebellious teenage son how well he took care
of him as a small child.
You haven’t a shred of respect for

The father who brought you up. Why, before you could speak
I knew if you gurgled “Bru-Bru,” you wanted your feeding bottle,
And if you went “Mam-mam-mam” I knew it was bread you wanted—
You hardly had time to say “Kakka” before I grabbed you
And held you out the door.
(Clouds, 1380–1384)
The last of the baby-talk words, “Kakka” (feces) provides one of the very few extant literary refer-
ences to toilet training; apparently, this aspect of childrearing did not provoke the discussion in the
ancient world that it does in our own day. Artifacts and vase paintings, however, allow us to infer
something about potty training. Excavated in the marketplace of ancient Athens, the Agora, was a
nearly fully intact potty chair. Beautifully painted and crafted, this was undoubtedly intended for sale
to a wealthy family. The overall dimensions suggest that the chair was made to hold a child weigh-
ing about 18 to 22 pounds, probably corresponding to an ancient 2-year-old; from this artifact we
can probably safely infer that toilet training took place around this age. Several vase paintings from
Athens depict small children in just this sort of chair. More interesting is the fact that the child could
not get in or out of the potty chair alone; an adult would have to put the child in, and once in, the
child would have to wait until an adult lifted her or him out. Can we infer from this evidence that
parents meant toilet training to signal the onset of restrictions on the child’s behavior? That such
devices were used to make toddlers stay put, at least for a little while? Unfortunately, in the absence
of literary evidence, we cannot confidently infer parental intent.
Strepsiades gives us other glimpses of a father’s close observations of his preschooler’s creative
activities.
When he was just so high he made

His own toy houses, he carved boats,
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He knocked up miniature carts from leather,

Cut frogs from pomegranates.
(Clouds, 877–881)
Even allowing for some exaggeration about his son’s accomplishments, there is no mistaking the
pride father expresses about son. He was a doting parent, as this final passage clearly reveals.
When you
Were a little boy I used to spoil you
And do what you asked—with the very first
Obol I earned as a juryman
I bought you a little pushcart
At the Spring feast.
(Clouds, 860–864)
There is additional evidence of male familiarity with the activities of toddlers and preschool-
ers (French, 1999). Little jugs called choes were given to small children or perhaps their families at
a spring festival when, probably at 3 years of age, children were officially accepted as members of
the father’s clan. Hundreds of these jugs have been discovered in excavations, and nearly all use as a
decorative theme little children—usually boys—at play. The figures of the children are delightfully
plump and happy; the vase painters show children indoors and outside romping, running, throwing
balls, playing with hoops and pets—engaged in precisely the kinds of activities we associate with this
stage of child development. Other literary and visual evidence confirms the range of toys provided
for small children we find on the choes.
In his consistent call for close supervision of children to maintain a stable, conservative state, Plato
(Laws, 793E–794B) provides a revealing description of the social life of preschoolers and the impor-
tance of play and games.
To form the character of the child over three and up to six years old, there will be need of
games . . . Children of this age have games which come by natural instinct; and they gener-
ally invent them of themselves whenever they meet together. As soon as they have reached
the age of three, all the children from three to six must meet together at the village tem-
ples . . . I assert that in every State a complete ignorance about children’s games—how that
they are of decisive importance for legislation, as determining whether the laws enacted
are to be permanent or not. For when the programme of games is prescribed and secures
that the same children always play the same games and delight in the same toys in the
same way and under the same conditions, it allows the real and serious laws also to remain
undisturbed.
. . . Alterations in children’s games are regarded by all lawgivers . . . as being mere matters
of play and not as the causes of serious mischief; hence, instead of forbidding them, they
give in to them and adopt them. They fail to reflect that those children who innovate in
their games grow up to be different from their fathers.
From this remarkable passage we can draw a number of inferences. First, here is clear adult male
recognition of one of the most important characteristics of this age group. Second, Plato sees that
play and games can be powerful tools of socialization of the young to the norms of adult society.
Third, he notes that children differ from one another; some are more likely to lead and innovate.
Perhaps the most interesting inference is Plato’s complaint that adults do not make much effort to
regulate children’s games and seem content to let their preschoolers have control over their own
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play and games. It is important that Plato does not condemn parents for neglecting their youngsters;
rather, he chastises them for letting their children play as the children wish.
A parental tendency to see safeguarding as opposed to carefully controlling preschoolers is con-
firmed for the wealthy by the institution of the paidagogue—literally a person who leads a child
around. Probably shortly after full weaning and toilet training, and around the time the child could
speak reasonably well, affluent parents turned over the hour-to-hour supervision of their boys to a
paidagogue, an older male (usually lower-class or slave) whose job it was to supervise the child, espe-
cially outside the home, and see to it that nothing harmful happened to the boy; we hear of a handful
of girls who had a paidagogue. The paidagogue was also expected to begin to teach the child proper
manners and help him improve his speech. In many cases, the paidagogue continued in service to the
boy and his family throughout his school-years and well into adolescence; literate paidagogues appar-
ently helped the boy with his school work. Within this relationship, the child probably had a good
deal of autonomy so long as the child’s behavior did not court danger or seriously violate standards of
conduct expected of this age group. And father and son had an intermediary who seems on occasion
to have been able to diffuse conflict, especially between an adolescent youth and his father.
Adult interest in and observation of small children is reflected to some degree in the large corpus
of Greek medical literature, beginning with that of Hippocrates (ca. 460–370 bce). Although there
is no treatise specifically focused on pediatrics among the Hippocratic works, references to children
and the occasions on which special treatment was required are frequent. In the beginning of Apho-
risms (1:13 and 16), Hippocrates stresses the differences between children and adults, noting that
children cannot tolerate fasts and need liquid diets during fevers. One essay, On Dentition, discusses
not only development of the teeth but also mouth and throat problems. In one section of Aphorisms
(3:24–26), Hippocrates lists the kinds of diseases most associated with each stage of child develop-
ment: infants are said to be vulnerable to aphthae, vomiting, coughing, insomnia, frights, infections
around the navel and in the ears; teething babies are subject to gum problems, fevers, convulsions,
diarrhea or constipation; toddlers run risks for tonsillitis, spinal disc misalignments, asthma, worms,
warts, leprosy, swollen glands in the neck, and tuberculosis. Elsewhere, Hippocrates mentions other
childhood diseases, such as mumps, convulsions from fever, diphtheria, polio, and meningitis. He also
notes that women put medicaments into children’s porridge to treat worms and other diseases. The
treatise on Epidemics shows that in some cases children were treated with surgery. The Hippocratic
attention to children’s physical health was continued in the biological works of Aristotle, the son of
a physician (Colon and Colon, 1999).
What generalizations about parenting in classical Athens can we make with some degree of confi-
dence from this evidence? First, parents—fathers as well as mothers—seem to have had a solid working
knowledge of the characteristics and needs of their young children; parents and adult society believed
that families should meet these needs. Second, the basic concept of the child as plastic and moldable
guided parental efforts to rear and shape the young into the kinds of adults they believed their socie-
ties needed. Third, parenting seems to have been attentive to children but not intrusive. Once out of
swaddling clothes, small children appear to have been supplied with toys and other objects for play
and allowed to follow their own instincts and desires with enough adult supervision to keep them out
of harm’s way. There is little in the evidence to suggest that Athenian parents became locked in some
early struggle for control; the child’s efforts to gain autonomy were probably usually successful. Fourth,
parenting was often a communal endeavor; society expected that in rearing young children, biological
mother and father would share with and receive support from a range of other adults—including men.
The Spartan Anomaly

This discussion has been based mainly on sources for classical Athens, and it is not unreasonable to
assume—given the cultural hegemony of Athens during those years—that what we see for Athens
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can be extended to other Greek city-states as well—with one important exception, Sparta. Sparta
was recognized by everyone as having a distinctive, if not totally unique, culture and political life. This
difference affected patterns of Spartan parenting significantly.
Classical Sparta lived in a state of permanent military preparedness against threats she perceived
within her own lands—a huge servile population tied to the soil—and on their borders—members
of their own Peloponnesian league as well as Athens and Persia. Until they were in their 40s, Spartan
male citizens lived in military camps and rarely spent time in their homes—the places where their
wives and small children lived.
As a result of Sparta’s thoroughgoing militarism, the rearing of boys up to the age of 7 and girls
until they married was left almost entirely in the hands of Spartan wives. And a Spartan women’s
chief job and basis for judging her worth was to produce more thoroughly Spartan soldiers and wives.
A Spartan mother’s attention was riveted on training her young children to become good Spartans.
Whereas in Athens children were an important, well-observed, and for the most part welcome part
of the household, in early and classical Sparta they were at the center of household attention.
The Spartans apparently practiced eugenics. First, women were not married until their late teens
or early twenties on the grounds that they bore more healthy babies and were less likely to die in
childbirth than young teenagers. Adolescent Spartan girls participated in rigorous physical fitness
programs, and Spartan wives were expected to maintain an extensive program of exercise to main-
tain their health. Second, Spartan men, apparently older ones, could “breed” their young wives with
younger Spartan males whose physical prowess and character they admired; it seems that Spartans
believed that as a man aged, his semen/seed tended to deteriorate also. Third, babies were inspected
soon after birth by a team of Spartan men who, probably on advice of the mid-wife, determined
whether the child was healthy enough to rear. How often sickly or deformed neonates were killed
by being thrown into a nearby gorge is unknown.
Spartan mothers personally supervised every aspect of her children’s growth, assisted by one
or more nurses. According to scattered sources (principally Aristotle and Xenophon), Spartan
babies were not swaddled at all; they were fully breastfed, sometimes by both mother and a wet nurse;
they were never fed even diluted wine; they were bathed frequently and in cold water. By the time
they were toddlers, mothers and nurses were teaching them not to be afraid of the dark, not to whine,
and not to be picky about their food. Lots of vigorous exercise and games under close supervision
filled the children’s days. Mothers were to imbue their boys especially with the strength and courage
needed to withstand the punishing course of military education boys began around age 7. It appears
that the primary mechanism of child discipline was the use of shame; children were encouraged to
tease and taunt their peers who stepped out of line.
For well over two centuries, Spartan childrearing produced men and women who were capable
of continuing the rigorous, highly disciplined Spartan way of life. During the later stages of the
Peloponnesian War in the last decades of the fifth century and into the first decades of the fourth
century when Sparta became the hegemonic power of all Greece, Spartan childrearing practices
changed dramatically. The Spartans suffered such heavy losses of men, and the remaining citizens
became so over-extended, that Spartan women began to take on additional functions within the
state, particularly the management of the huge estates owned by each family and a good deal of the
daily commercial and economic activity of the state. The focus of Spartan home life was no longer
fixed on the children and on inculcating the values of Spartan society; childrearing was increasingly
turned over to the nurses who, because they themselves were not products of the Spartan system,
could not transmit successfully Spartan values to their young charges. By the middle of the fourth
century and for generations thereafter, when Sparta tried to reclaim power, she tried to reform, to
return to the former hard self-discipline of earlier times. She failed, in significant part, because the
habits of parenting had so dramatically changed. Here is one excellent example that demonstrates
how changes in parenting can be connected clearly with changes in adult society (French, 1997).
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The Greeks: Some Conclusions

It is with the Greeks that we find the first systematic observations and thinking, even theorizing,
about parenting and childrearing—spurred by their belief that the stability of the society and the state
depended on producing new generations capable of and committed to maintaining the community.
Greeks created models of child development that closely parallel contemporary theory. Emphasis
within the family on parenting practices advocated by the aristocratic elite was probably more preva-
lent in the upper classes. Because Greek poleis were fairly small, it is likely that elite beliefs about
proper parenting and childrearing practices permeated the entire community; to what extent parents
in the lower classes had the resources and time to emulate aristocratic practices cannot be determined.
The Romans
Historians today, as well as Romans themselves, remarked on the extent to which Rome incorpo-
rated many aspects of Greek culture into their own. Many historians posit a Greco-Roman civiliza-
tion that began soon after Rome added Greece and the eastern Mediterranean world to its empire
in the second century bce. Indeed, there are many commonalities between Greek and Roman
ideas about and practices in parenting and childrearing, and Romans clearly adopted much of what
they found in Plato and Aristotle and other Greeks in their own philosophical thinking. However,
Roman family relationships were probably more complex and variable than those of the Greeks, and
it appears that Romans expressed more interest in and attention to affective bonds between parents
and children than did the Greeks.
Familia, Domus, and Patria Potestas: The Cultural

Context of Roman Family Life
Like the Greeks, the Romans’ model of the state grew out of their conceptualization of the house-
hold; but unlike the Greeks, the Romans developed two distinct constructs. The first and most com-
monly used is the Latin familia which is probably closest to the Greek oikos in that it encompassed
not only the central nuclear grouping of father/mother/children but also co-resident kin, retainers
and other dependents, slaves and all chattel property, as well as all land and buildings. When Romans
meant just the nuclear unit’s people and dwelling, they used the word domus, which is more like
our word “home.” It was the familia that interacted with the state, the public sphere, and it was the
familia to which Roman law applied; it was the domus that connoted the private sphere within which
Romans conducted what we would call “family life.”
Within the home and household, the Roman pater familias, like the Greek kurios, was in charge.
However, the Roman father’s power, the patria potestas, was absolute; a father had the right to kill
anyone—including his grown children; his right to impose less than capital punishment was also
absolute; and he could sell his children as slaves. There are a number of examples of fathers who
exercised their right to execute their grown children. This absolute power retained legal sanction for
nearly a millennium, ending only in the late fourth century ce, after Christianity had significantly
changed the core value structures of the Roman imperial world.
In practice, the exercise of patria potestas was restrained by a number of factors. First, before killing,
exiling, or severely punishing adult children, a father was expected to take counsel with the other
adults, including the women, of the familia; few fathers risked acting in contravention of this family
council. Second, the role of the mater familias traditionally expected her to ameliorate the harshness
of paternal discipline. Third, a Roman law, allegedly instituted by her founder Romulus, limited
a father’s right of life or death over his newborn children; Romans were obligated by law to rear
all sons and at least their firstborn daughters to adulthood. Finally, surely in the early years of the
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Republic, fathers were the main teachers of their sons; Romans expected the bonds forged between
father/teacher and son/pupil to be strong enough to withstand intergenerational conflicts.
The alleged Romulean law compelling fathers to rear newborns suggests some notion of legiti-
mate state intervention in the private sphere of Roman families. We see later evidence of attempts of
the state to address what was apparently a general problem—the failure of Roman families to replen-
ish themselves. The Roman practice of dividing the paternal estate among heirs (daughters as well
as sons) seems to have prompted families to adopt a strategy of trying to have but one or two heirs
survive their father so as not to divide and redivide the patrimony until each heir’s share was so small
as to count for very little. The problems of not having a system of primogeniture (i.e., eldest surviv-
ing male child inherits the entire estate) extended into the middle classes and small farm families.
Given the nearly incessant and large-scale warfare in which the Romans engaged for many centuries
as well as childhood mortality rates, it must have been very difficult to carry out this reproductive
strategy successfully. Romans seem to have consistently erred on the side of insufficient reproduction
rather than risking excessive division of the estate probably because the practice of adopting adult
male heirs was widely accepted and offered powerful families yet one more way to manipulate kin-
ship ties to their political advantage. The ability of the Romans to err on the side of producing too
few children who survived to adulthood suggests not high rates of infanticide—for which there is
little evidence and which was illegal anyway—but reasonably effective family planning methods of
contraception and abortion (Riddle, 1992).
The potentially monstrous powers given to fathers and the centuries-long problem of families
failing to reproduce themselves suggest an important element of the social context for investigation
and interpretation of the Roman social construction of parenting. At the very least, it seems that
there must have been some deep ambivalence about receiving life from the previous generation and
handing it over to the next.
One other aspect of the social context of parenting deserves mention—the relatively high rates
of divorce and remarriage that seem to have characterized the Roman aristocracy and affluent and
aspiring families. Although marriage and divorce surely occurred among the Greeks, the Romans
played this game with considerable gusto; many amicable divorces left the ties of friendship that
once united the families fairly well intact. Thus, the individual Roman familia was most often a set
of blended and reblended families forming a complex web of blood and current marriage/former
marriage relationships. Within the domus, the basic unit of husband/wife/children was stable, but
the individuals playing those roles could change frequently within one generation. When the death
of women in childbirth and the mortality rates in the Roman military are added to the ease and
acceptability of divorce, we have to posit nuclear families for Roman parents and children that were
fluid at the least. When we examine Roman ideas about and attitudes toward young children, it is
important to keep in mind this rather somber background of the patria potestas, reproductive failures,
and frequent reconfigurations of the people within the domus.
The Social Construction of Children and Childhood

There is general agreement among historians of Roman family life that sometime between the
beginning of the second century and the end of the first century bce, the Romans’ social construc-
tion of the young child changed significantly, and changed in such a way as to emphasize affective
bonds between parent and child (Dixon, 1988, 1992). Evidence adduced includes changes in vocabu-
lary used to describe small children, increasing evidence of parental mourning the deaths of small
children, and increasing references to the active presence of small children in literature and art (Evans,
1991). However, the emotional investment Romans of the regnal and early and middle republic
made in their children should not be underestimated. A number of factors prompt this caution. First,
evidence for family life prior to the second century bce is extremely meager and cannot be regarded
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as a random sample of what was once available. Second, the important cult of the Mater Matuta—the
nurturing or nourishing mother—in early Rome suggests considerable familial and state investment in
rearing the young. The cult statues of this goddess show a hefty, mature woman seated on a throne
holding from 1 up to 14 swaddled infants. That one of early Rome’s major cults centered on this
visual representation suggests the crucial importance of the young child in Roman society. Finally,
meager though the evidence is, a number of stories glorify parents who invest time and effort in
rearing their children. For example, Plutarch reports in his life of Cato the Elder that the redoubtable
and stern statesman regularly rushed home from the Senate to supervise the bathing of his infant
son. Nonetheless, it is probably true that, by the time Rome acquired an overseas empire and came
into contact with the Greek East, the Roman social construction of the small child had changed and
families paid more attention to affective bonds between parent and child.
Perhaps the most important single source for the current of Greco-Roman thought that put
emphasis on loving parents and children is Plutarch, the prolific writer of the second century ce.
Throughout his biographies of famous Greeks and Romans, he includes stories of his characters as
youngsters, usually to demonstrate that adult characteristics could already be discerned in childhood.
In a number of his essays, Plutarch reveals the dynamics of his own family, indicating that he and
his wife shared in the parenting of all their children, four sons and a daughter. When his only girl,
Timoxena, died at the age of 2, Plutarch was away from home and composed an elegant letter of
consolation to his wife. In it, Plutarch recounts his own fond memories of his little girl:
There is a special savor in our affection for children of that age; it likes in the purity of the
pleasure they give, the freedom from any crossness or complaint. She herself too had great
natural goodness and gentleness of temper: her response to affection and her generosity
both gave pleasure and enabled us to perceive the human kindness in her nature. She would
ask her nurse to feed not only other babies but the objects and toys that she like playing
with, and would generously invite them, as it were, to her table, offering the good things she
had and sharing her greatest pleasures with those who delighted her . . . our daughter was
the sweetest thing to fondle, to watch, and to hear; and we ought to let the thought of her
also dwell in our minds and lives, for there is much more joy in it than sorrow.
(Pomeroy, 1999, pp. 59–60)
Not surprisingly, given intercultural borrowings, Roman concepts of the child and stages of child-
hood differed little from those of the Greeks. Like the Greeks, Romans conceived of their small chil-
dren as plastic, unformed, moldable, ignorant, unaware and, thus, susceptible to corrupting influences
and potentially fragile. Writing at the end of the first century ce, the rhetorician Quintilian illustrates
well the Roman belief in the moldability of the young and confidence in children’s abilities to learn.
I would, therefore, have a father conceive the highest hopes of his son from the moment
of his birth . . . you will find that most [children] are quick to reason and ready to learn. . . .
Those who are dull and unteachable are as abnormal as prodigious births and monstrosities
and are but few in number. . . . Let us not therefore waste the earliest years; there is all the
less excuse for this, since the elements of literary training are solely a question of memory,
which not only exists even in small children, but is specially retentive at that age. . . . He will
remember such aphorisms even when he is an old man, and the impression made upon his
unformed mind will contribute to the formation of his character.
(Institutio Oratoria, 1.1.x)
Romans also saw their youngsters as playful, cheerful, affectionate, and gregarious. Romans seem
to remark more often on children’s unruliness and their natural imitativeness. Probably due to the
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influence of Stoicism and Christianity, Romans frequently comment on the child’s natural inno-
cence. In the Roman construct of the child, there are three traits not recorded among the Greeks:
competitiveness, curiosity, and natural facility of memory—all three characteristics put to good use
in Roman theories of education (Bonner, 1977).
Just as Plato and Aristotle set out basic theories of child development, so too the Romans Quin-
tilian and Augustine describe similar theories for their own cultures. Quintilian and Augustine give
more attention to issues of discipline than their Greek predecessors. There seems to have been more
ready use of physical punishment among Romans and more contention—and perhaps feelings of
ambivalence—about its successful controlling versus detrimental effects on young children.
Baby and Childcare in the Roman Domus

Pregnancy, childbirth, and childhood for the Romans were just as fraught with the specter of death
as for the Greeks. Work on Roman demography suggests that about a quarter of Roman babies died
before their first birthday and that only half of Roman children lived past the age of 10 (Bradley,
1999). Parents undoubtedly expected a heavy toll to be paid to neonate and childhood disease; but
nonetheless many, like Plutarch, made considerable emotional as well as physical and financial invest-
ments in rearing their young.
Quintilian and Augustine described childrearing practices among the upper classes during Rome’s
imperial era. In much earlier periods of Roman history, before she gained the spoils of an empire that
stretched across the Mediterranean basin and well into Western Europe, the care of young children
would not have been entrusted to nurses and paidagogues; according to the Roman historian Tacitus
(Germania), mothers and fathers and co-resident kin would have done this work. As noted above, in
the centuries before Rome acquired an overseas empire, fathers seems to have taken over the training
and education of their male children as early as the age of 3.
But by the second century bce, both the upper and middle-class Roman domus would be likely to
have several attendants, probably slaves, to nurse and look after babies and preschoolers. The sources
describe a considerable range of parental surrogates: other older relatives, such as grandmothers
and aunts; wet and dry nurses for infants; other people called nurses, some of whom were male, for
toddlers and preschoolers; paidagogues; and tutors to begin teaching the children the alphabet and
simple numbers. It is worthwhile noting that literary references and funeral epitaphs refer to these
caregivers with respect and affection (Bradley, 1991). Many children grew up knowing and being
able to trust a wide circle of adults and in later years remembered this experience positively. Given
the frequent change of the individual people who constituted the Roman domus at any one time,
it is perhaps not surprising that responsibilities for baby and childcare would be entrusted to adults
who were likely to stay attached to a single child and that, as grown-ups, children would remember
these caregivers fondly. The only parental surrogate who seems uniformly despised is the stepmother,
who is nearly always portrayed as wicked and mean. We can only speculate that stepmothers were
expected to favor their own biological children over those in her new family.
One can hardly mention Roman mothers without immediately thinking of that paragon, Cor-
nelia, daughter of Scipio Africanus, hero of the war against Hannibal, and mother of the Gracchi,
her two sons who started the political revolution that eventually led to the demise of the Roman
Republic and the imposition of one-man, imperial rule for the Roman empire. Historical sources
uniformly describe Cornelia as the educated and highly cultivated mother of 13 children who,
when her husband died, refused to remarry in order to devote her time, energy, and attention to the
education of her offspring. Her home was a regular meeting place for discussions among the most
notable and intellectually gifted philosophers and politicians in Rome. Cornelia was unaffected by
the wealth and status of her family. A story has it that once, while walking along a street, Cornelia
was asked by another wealthy matron, “Where are your jewels?” Pushing her two sons in front of her,
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Table 8.2 Characteristics and/or proper care
Age Level Quintilian Augustine
Birth–3 years In charge of nurses and then paidagogue At first can only suck and cry
Highly impressionable; therefore nurses Then begins to smile and laugh
and paidagogue must speak properly and Frustrated at being unable to communicate
set fine moral examples and thus cries even more
Has learned to speak Easily angered; often sullen
Jealous of other babies
Learning to speak; at first just one word at a
time; makes lots of grunting noises; learns
without being taught by adults; is aware of
nonverbal cues, “body language.”
3 to 7 years Still at home but education and training Remains at home
should begin Learns to speak well
Memory is acute and retentive At beck and call of others
Not capable or originality Often disobedient
Needs studies to be amusing
Needs stimulation of praise and sometimes
of competition
Suit tasks to child’s limited abilities
Do not push too hard
7 to Boys sent out to school Boys go to school
adolescence Instruction should be tailored to Often unruly and disobedient
individual student Do not like to study
Needs rest and relaxation Love to play
Play is good for boys Irrational likes and dislikes
Character formation is crucial before boys Curiosity leads to learning and frightful
learn deceit punishment hampers learning
Avoid flogging and abusive punishment
Cornelia retorted, “Here are my jewels.” Cornelia has long been regarded as an incarnation of the
ideal Roman wife/mother: married only once, well-educated, and intimately and absolutely devoted
to the welfare of her children.
In her study of Roman motherhood, Dixon (1988) compelled a revision of our interpretation
of Cornelia, demonstrating that her relationship with her children has been constructed in light of
twentieth-century ideals of motherhood—making her far closer to her young children than actually
was the case. Dixon cogently argued that the conditions within which Romans carried on family life
necessarily led to a more distant physical and emotional relationship between mother and child than
we, from our contemporary perspective, think natural. Dixon concluded that the ideal or stereotypical
Roman mother—including Cornelia—was (1) an important and vocal figure within the household
whose position of influence depended heavily on custom and tradition; (2) not connected more
intimately with her babies and young children than with them as adolescents and adults; (3) closely
associated with the father as a generic parent responsible for the children’s discipline, education, and
development of moral character; and (4) a strong protector of her children’s interests and well-being
but not a special source of affection and tenderness in comparison with other family members—
including fathers. In short, the roles of Roman mother and father do not seem as closely tied to female
and male gender constructs as were the Greeks’—or for that matter, those of our own day. Dixon and
other scholars believe that Roman mothers generally had stronger bonds of affection and influence
with their grown children, particularly their sons, than with their children when they were small.
314
Of particular interest is Dixon’s demonstration of a parallel growth of women’s de facto ability to

bequeath their personal property, usually to their biological children, and social recognition of the
importance of the mother-child bond. This aspect of Dixon’s work allows us to see an historical
change in the Roman’s construct of the “natural” relationship between a mother and her children.
Dixon’s argument together with some literary and iconographical evidence about early Rome sug-
gest that, before imperial times and when nuclear units were relatively stable, Romans thought in
terms maternal care in general for children in general. In the main festival for the Mater Matuta, the
Matralia, women carried in their arms to temples, not their own infants, but those of their broth-
ers and sisters, clearly indicating a collective maternal responsibility for the family’s babies. Perhaps
as Roman women began to think of themselves, together with their personal property, as unique
individuals, they also began to claim a unique relationship with their children, symbolized by their
newly acquired right to dispose of their property by testament as men did. Against the backdrop of
the patria potestas, the growth of a social recognition of the importance of the mother-child bond
also appears to coincide with a relaxation in the beliefs about how stern a father should be with his
children. Plutarch’s essays and biographies indicate that he expected good fathers to be moderate,
even compassionate, in their discipline of their children.
Dixon’s (1988) work on Roman mothers and their bonds with their adult sons was preceded by
Hallett’s (1984) study of father-daughter relationships among elite Roman families. Relying mainly
on Roman legends in which daughters play pivotal and influential roles in motivating their fathers,
and on the evidence in Cicero’s extant correspondence about his doting affection for his daughter,
Tullia, Hallett shows that in many families the strongest dyadic bond of affection was likely to be
one between a father and his daughter. A “daddy’s little girl” was more likely to marry and divorce
compliantly; more likely to work hard to further her father’s political interests among her in-laws;
more likely to bring crucial political and economic intelligence to her father; and less likely to be
alienated from her father by the competition that existed between Roman generations. Conversely, a
“daddy’s little girl” was more likely to receive continued financial support if her in-laws were either
stingy or hard up; more likely to be rescued from an unhappy marriage; and more likely to receive a
larger share of the inheritance. A good deal of evidence suggests that girls and young women were
most likely to receive approval and unconditional acceptance from their fathers than from any other
family member. And that same evidence suggests that men were more likely to receive unconditional
acceptance from their daughters than any other family member.
Parents saw to it that their children had ample opportunity to join their peers outside the domus
for play. The evidence indicates that parents provided a wide range of toys and games for their
youngsters. Roman children shared their parents’ enthusiasm for gambling and used dice, coins, and
nuts in games like heads or tails or odds and evens. Children from wealthy families could expect their
parents to give them marbles, balls, spinning tops, spinning hoops, pull-toys, push-toys, scooters, even
small-scale chariots pulled by a dog, pony, or goat. Archeological excavations have unearthed a wide
variety of dolls from crude figures fashioned from wood or clay to near works of art sculpted from
bone or ivory with articulated limbs. Visual and literary evidence depicts children playing a wide
variety of games from skipping rocks over waves, building sand castles, tag, leap-frog, and blind-man’s
bluff. Adults recalled their childhood play with considerable pleasure and apparently enjoyed watch-
ing their own youngsters engaging in similar activities (Evans, 1991).
Greek medicine’s attention to infants’ and children’s health continued on into the Roman world,
with a good number of treatises that provide significant insight into Roman ideas about pediat-
rics. The Hippocratic distinction between treatment of children and adults was fully developed
by Celsus’s (ca. 30 bce–45 ce) clear dictum, “Children must be treated entirely differently than
adults” (de Medicina 3.7). Pliny the Elder (23–79 ce) preserves much valuable information about
folk medicine in his Historia Naturalis; here we catch a glimpse of the kind of treatments—mostly
herbal and dietary—most Romans probably employed to care for sick children. Some may have been
315
Valerie French
efficacious, such as the use of the plant ephedra for a bloody nose and asthmatic cough, or liberal
use of garlic, oil, and fish sauce. Of particular interest is the extensive treatise on obstetrics and care
of neonates written by the physician Soranus in the second century ce; his references to handbooks
written by midwives suggest that the corpus of medical literature relating to children was far more
extensive in the Roman world than in the Greek. The pediatric sections of late ancient and early
Byzantine medical epitomes and encyclopedias by Oribasius, Aetius, and Paul of Aegina confirm this
view (Colon and Colon, 1999).
The Romans: Some Conclusions

On the basis of the available historical evidence, it appears that the concept of patriarchal power
reached an apogee in the Roman construct of patria potestas, the right of the eldest male to execute
his children, even as adults. Fathers were undoubtedly powerful figures within their families, but their
ability to exercise this power was tempered by informal mechanisms of social control: an expectation
of agreement by a family council and an ameliorating influence of the mother.
The Roman mother also played a key role in parenting not only of babies and preschoolers but
also of adolescents and young adults. She was seen as a protector of her children’s interests. Mother
held a position of influence in the home and was closely associated with the father as a generic parent,
responsible for children’s education and training.
Parenting must also be considered in light of two other important characteristics of Roman
families: failure of the upper classes to reproduce themselves and fluidity in the composition of aris-
tocratic and affluent nuclear families resulting from relatively high rates of divorce and remarriage.
Children of elite families were likely to have a variety of parental surrogates.
Probably from its earliest period, Roman parents regarded their roles as important and devoted
considerable time and attention to rearing their young. Fathers were expected to supervise their sons’
education personally, and there is good evidence that many did. The bonds tying parents to children
may have become increasingly affectionate beginning in the second century bce. The most affection-
ate bond within the family may well have been that between fathers and daughters.
Along with other borrowings, the Romans apparently adopted the Greek social construction of
the young child, stages of childhood development, and interest in pediatric medicine. And like the
Greeks, Romans saw childrearing primarily in terms of influencing the physical, intellectual, and
moral development of the young; education was crucial in this process.
Conclusions
What conclusions about parenting can be drawn from this ancient historical material? Constructing
generalizations about relatively poorly documented societies is fraught with difficulties. The best
we can do is to point out that parenting was mainly a private as opposed to public concern, but
political philosophers and politicians recognized the critical importance of this activity. A consider-
able number of people, including males, in ancient societies had an understanding of how children
grow and develop that parallels contemporary theory. To what extent such people predicated their
parenting behavior on this understanding is unknown. Many parents invested heavily in rearing their
children and loved them dearly. Simultaneously, we can be sure that large numbers of children suf-
fered serious physical and psychological deprivation and abuse. Despite detailed knowledge of what
parents needed to provide for their offspring, we know of no systematic effort to eradicate child
abuse. Perhaps the only safe general conclusion is that the range of approaches to parenting we see
in contemporary society was paralleled in the ancient world.
Despite these caveats, we can see some differences in approaches to parenting among ancient
civilizations. Upper-class Egyptians seem to have delighted in their children, giving them prominent
316
positions in representations of family life. Mothers and slaves probably oversaw the care of babies and
preschoolers, but fathers likely personally taught and prepared their sons to follow in their footsteps
as adults. In contrast to Egyptians, Mesopotamians did not celebrate family life, and children are
nearly absent from official and funerary art. Many Mesopotamians seem to have regarded parent-
ing and family demands as a difficult burden. Legal evidence suggests a perceived need to protect
children from ill-treatment at the hands of their parents. It appears that Mesopotamian parents were
more emotionally distant from their children than were Egyptians. The ancient Israelites made the
family—both its bonds and affection and potential for tension and conflict—a metaphor for writing
about the relationship between themselves and their God.
The Greeks were the first to think systematically about parenting and child development, and
they laid the theoretical foundations for childrearing and education that dominated the Mediter-
ranean world for nearly a millennium (ca. 600 bce to 400 ce). Greek and Roman descriptions of
childhood and prescriptions for parenting reflect an emphasis on the effects of nurture and the need
for parents to invest the time, energy, and resources appropriate for the particular stage of the child’s
development. Alongside evidence that demonstrates keen observation of children and their needs
stands other evidence pointing to neglect and abuse of the young. Among the Greeks, the picture of
parenting and family life derived mainly from evidence for ancient Athens can probably be applied
to most other Greek city-states. But the Spartans developed an anomalous approach to parenting,
leaving childrearing of boys until the age of 7 and girls until they married almost entirely to Spartan
mothers. In classical Sparta, children were the central focus of the household; fathers were nearly
entirely absent due to their fulltime involvement in the Spartan military.
In many respects, the picture of Roman parenting is like the Greek. However, fathers had more
legal power over their children, and the roles of mothers and fathers may have been more similar,
with both responsible for the physical, educational, and moral development of their children. Within
the Roman family, the bonds between fathers and daughters may have been the most affectionate.
Can we compare parenting in the ancient world with that of our own era? The most important
common elements are the centrality of rearing children in the social construction of the family and
the variety of assumptions and practices that underlie childrearing both then and now. Although the
Greeks were the first to articulate a proto-theory of child development, all these ancient cultures
recognized stages of children’s growth and sought to employ methods of childrearing appropriate
for those stages. The most important difference is a tendency in some contemporary Western cul-
tures to romanticize the child and childhood and the appearance of an ideology that seeks to create
child-centered societies; in the ancient world, children were an important and integral part of the
family—but not its sole focus.
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9
MODERN HISTORY
OF PARENTING
Peter N. Stearns
Introduction
The history of modern parenting focuses on changing roles and functions for children—in the
family and in society at large—and how parents both facilitated and reacted to these changes. But it
also reflects larger shifts in society, such as the increasing separation of work from home or changing
gender roles that generated often complicated and difficult parental responses. Modern childhood
and the modern family differ from their premodern counterparts, and parenting is directly involved
in these transformations.
The conditions of modern parenting began to take shape in the 18th century in response to
new ideas about children (primarily, initially, in Western Europe and North America) but above all
to shifting economic patterns soon including the early stages of the industrial revolution. Whether
because of industrialization itself, or efforts to prepare the preconditions for economic change—as
in reform-minded Japan after 1868—many aspects of parenting had to be reconsidered, by families
themselves or in response to new social policies such as school requirements. Analysis of the emer-
gence and evolution of modern parenting thus focuses essentially on the past 250 years.
This said, it is also important to note the several complexities in dealing with the history of mod-
ern parenting. First, and most obvious, is the fact that parenting retained important continuities with
the premodern past. Parenting involves, after all, some biological constants, beginning with the act
of giving birth and extending through the general need for substantial parental attention (directly
or through surrogates) for various aspects of child development in a species in which the young are
unusually dependent for a prolonged period of time. Modern parenting also inherits traditions that,
although often modified, continue to provide guidance. The role of religion, for many parents, is an
obvious example. Debating the precise balance between change and continuity in parenting is in fact
a challenging assignment, and we must guard against overemphasizing change.
Throughout the modern period as well, social class complicates any generalizations about parent-
ing. Not surprisingly, historians have found it easiest to deal with middle- and upper-class patterns,
where the evidence is most abundant. And it is true that in many societies, urban middle classes took
the lead in behaviors that would later be more widely adopted, from new levels of birth control to
an emphasis on the importance of schooling. Even when changes were widely adopted, social dif-
ferences conditioned responses, for example in the ways that parents defined their roles in preparing
school achievement, or the precise levels of birth rate reduction. Throughout the modern period as
well, and still today, significant rural-urban gaps condition historical analysis.
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Modern History of Parenting
Geographic variance constitutes a final complexity, surely the most important one from a global
standpoint, and it has several facets. In the first place, key developments in modern parenting took
shape at different times in different regions. Adaptation to lower birth rates, for example, began to
develop in Western society in the 19th century, but the same process spread widely in Latin America
only much later. The phenomenon has similar although not identical features, but the gap in timing
bedevils comparative generalizations at least until very recent decades.
Second, many regions continue to insist on differences in parenting even as they accept some
common patterns. East Asian parents, and East Asian society more generally, are far more comfort-
able using shame as part of children’s socialization than are their U.S. counterparts. Physical disci-
pline continues to vary quite widely. Historians debate the global relevance of some other aspects
of parent-child relationships, such as the salience of any idea of adolescence outside the Western
context (Fass, 2016).
Finally, and at various points in the past two centuries, parenting in a number of regions was
shaped by particular disasters, most notably in response to war or civil strife. Parenting among refu-
gees, for example, became an all too common subtheme. Unusually acute poverty, or special labor
systems such as indenture, also compelled distinctive responses.
All this adds up to a fundamental challenge in dealing with modern parenting: It is risky, and
potentially misleading, to claim too much attention to overall global patterns. Comparative analysis is
essential, but the comparisons are often difficult—hampered as well by variance in relevant histori-
cal scholarship—and demand considerable nuance. It is easy, to take the most obvious problem, to
overemphasize the relevance of Western patterns of change, to assume that they have wider global
reach than is in fact the case.
At the same time, modern parenting, over the past 150 years, does reflect some dominant themes, as
parents sought to adjust to changes in the economy and to new involvements by the modern state. There
were some common patterns in the ways most parents sought, or were pressed to accept, changes in their
goals and behaviors. These patterns must be balanced against the comparative complexities.
This chapter deals briefly with the emergence of serious historical research on modern parenting,
a relatively recent development that has yielded a number of significant findings and with several
key topics clearly open to further analysis. It then turns to the principal modern patterns that have
ultimately affected most world regions at least to some degree over the past century or two. A final
section offers several more specific case studies, where particular factors have shaped modern parent-
ing or where comparative differentials must be explored within the common modern framework.
The Emergence of the History of Parenting: Debates and Theories

Serious historical work on parenting began to emerge in the 1960s, as a function of the develop-
ment of social history as a major research focus in the discipline. Social historians contended, and still
contend, that coverage of the past must include attention to ordinary people as well as the doings of
elites, and therefore to activities beyond the confines of conventional political, military, or intellec-
tual history. Taking shape initially in France in the 1930s, social history spread increasingly to other
countries, including the United States, in the decades after World War II. Mutual influence played a
role here, as attention to what the French called the Annales school gained ground elsewhere (Murra,
Wachtel, and Revel, 1986), but it was also true that contemporary social issues contributed as well,
spurring historians to look to the past to help ground relevant analysis of current social problems.
Thus in the United States, where concerns about family stability took hold amid a rising divorce
rate, a new group of historians began to focus on family history. Family relationships constituted a
key theme for a cluster of historians who dealt with colonial New England, in work that began to be
published by the mid-1960s. Demos (2000), for example, emphasized the differences between paren-
tal goals and behaviors in one Massachusetts community, and those characteristic of the modern
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Peter N. Stearns
United States. Greven (1988) traced changes in parental behavior by the 18th century and examined
a set of different parenting styles (moderate, strict, and indulgent) that, he argued, took shape by the
later 18th century but would inform subsequent distinctions as well—particularly in the persistence
of a gap between evangelical and more moderate Protestants. This kind of work set in motion a series
of historical inquiries that would take up other aspects of parenting in American history, but might
lead as well to research on other regions—as in Gillis’s (1981) pioneering study of the emergence of
adolescence in Western Europe.
This American current was long overshadowed, however, by the tremendous influence of the first
great study relevant to modern parenting, Ariès’s Centuries of Childhood, published in French in 1960
and translated into English two years later. Ariès, to be sure, focused primarily on social patterns in
general, not on parenting specifically, but his approach provided an important framework for the
study of parent-child relationships, that would be taken up by many successors.
Ariès argued, based among other things on his analysis of artistic evidence, that an explicit con-
cept of childhood as a distinct phase of life emerged only in modern times, from the late 17th
century onward. Prior to that point, Western society tended to see children (at least after infancy)
primarily through the lens of adulthood rather than recognizing particular needs and characteristics.
Parents, correspondingly, viewed children primarily as young adults, linked of course to expectations
of productive work. Modern society, in contrast, began to approach children as a separate category,
requiring special treatment—for example, through greater attention to education. Ariès recognized
some disadvantages to the modern approach, for example in the greater supervision of children
that might result, but evaluation merely embellished his insistence on a clear premodern/modern
dichotomy (Ariès, 1962).
Ariès’s insights quickly guided a number of other historians, both European and American, work-
ing primarily on European developments on the cusp of modern times (Hunt, 1970). Stone (1983),
for example, in a study of the family, sex, and marriage in early modern Britain, emphasized how
different English parenting was during the bulk of his period from what began to emerge in the 18th
century. Stone’s premodern British parents paid little emotional attention to their children, seemed
relatively unconcerned about the high infant death rate, and disciplined severely. Stone’s findings,
including the continued insistence on a substantial premodern/modern gap, were echoed by a num-
ber of other historians in other studies of the European family or in specific work on sites such as
17th-century France.
This kind of analysis was bolstered, if sometimes indirectly, by two more theoretical approaches
that were gaining wide attention in the same decades. Historians of the family did not necessarily
invoke modernization theory, but a modernization framework might well be applicable to the kind of
work that took shape under the Ariès’s inspiration. Overall theory of modernity was developed in the
1950s and 1960s, mainly by American sociologists, and it could include claims directly relevant to a
history of modern parenting. Emphasis on the growing importance of education in modern societies,
for example, or the new kinds of contacts developed between the modern state and ordinary citizens
were certainly compatible with the work emerging in the history of the family (DeMause, 1974).
A number of historians more explicitly picked up on a second general approach, launched by the
German sociologist Elias (1982) under the heading of a “civilizing process”. Elias contended that
from the Renaissance onward, beginning in the upper classes but gradually disseminating downward,
Westerners sought greater control over their bodies and emotions in the interests of respectability
and good manners. Bodily disturbances, such as farting or belching, now must be restrained, and
attention to emotion meant new attention to mastering anger or jealousy. Elias emphasized larger
cultural patterns, but his work could easily be applied to parental behavior. “Civilized” parents, in this
rendering, now had to pay more attention to monitoring their offspring, using discipline including
new forms of shaming to teach them those habits that would ultimately qualify them for a respect-
able adulthood. In this rendering modern parenting emerged a bit more gradually than in the Ariès
322
vision, and it involved more explicit emphasis on parental control, but the notion of a sizeable
ultimate gap between the modern and the premodern parent gained additional support from this
approach. Elias’s work was picked up by a number of historians of the family and was applied to the
19th-century United States in a study by Kasson (1990).
An important complement to the Elias approach was, finally, developed by the Dutch historian
Huizinga who argued, in his Homo Ludens (1955) that modern societies significantly constricted
children’s opportunities for spontaneous, creative play. New forms of societal supervision and the
growing interest of parents themselves in shaping early childhood to prepare for formal education
had the ironic effect of constraining the range of children’s activities that had been common in early
modern times. Parents began to develop more systematic supervision (Huizinga, 1955).
As Huizinga made clear, the general tendency to emphasize a major modern/premodern distinc-
tion in childhood and in parenting could win a negative appraisal with an argument that premodern
children were freer and more natural before the imposition of the apparatus of modernity. The pre-
dominant tone among historians of the family and childhood, however, as with Stone, emphasized
the greater virtues of the modern parent. Less forceful discipline—particularly, physical discipline;
growing attention to the child’s educational needs; even parental love all blossomed in modern times,
in contrast to the bleak circumstances of the premodern family. Stone, arguing for example that one
would expect in the premodern family to uncover no more emotion than can be found in a bird’s
nest, epitomized the kind of positive stamp that a whole generation of interested historians applied
to the modern treatment of children.
The premodern-modern evaluative distinction inevitably produced a marked backlash by the
1980s. A variety of medievalists directly attacked Ariès’s contention that a notion of distinct child-
hood was a modern creation. They showed that in theology, in law, and in other settings that child-
hood was explicitly identified and valued. Pollock (1983) focused on early modern conditions and
on parenting more explicitly, finding abundant evidence of parental affection for children, concern
for their welfare, and grief at their passing well before the 18th century in Europe. On another front,
even more recently, emotions historians like Rosenwein (2016) attacked the Elias approach, finding
clear emotional communities in the Middle Ages that valued emotional restraint. Not coincidentally,
these kinds of complexities in evaluating the relation between premodern and modern families were
echoed, also in the 1980s, by a growing group of historians who attacked the whole idea of mod-
ernization applied to the past, contending that the theory provided a set of oversimplifications that
ignored differences among regions, social groups, and gender factors and should simply be jettisoned.
The result of this thrust and counterthrust, during the past decades, has several facets. Historians
of children and parenting, although returning to the subject after a brief pause, have become far
more cautious about generalizations distinguishing modern and premodern parents—including, for
the most part, any attachment to overarching theories. They look for continuities as well as changes.
They examine particular topics, such as parent-child interactions through consumerism or the sig-
nificance of the emergence of the idea of adolescence in parent-child relationships, rather than offer-
ing overall generalizations about change. They are open to the exploration of the downsides of the
modern parent-child relationship—for example, probing the reasons for the modern emergence of
anorexia nervosa ( Jacobs, 1988)—as well as more positive features of changes in parental discipline.
The result is the steady accumulation of additional empirical work, but also a greater timidity about
large generalizations and a distaste for potentially simplistic and unduly positive renderings of the
modern experience of childhood and parenting.
One other shift, although more tentative, also marks the most recent research in the field. The
whole approach stimulated by Ariès and Elias had been West European in focus. This regional thrust
has been complemented by the continuing work on family themes in American history, but it risked
resolutely ignoring the rest of the world or assuming that global patterns could somehow be shoe-
horned into Western models. Recent decades have seen a welcome surge of interest in aspects of the
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history of parenting in Russia, Latin America, Africa, and particularly China (Hsiung, 2005), even as
Western research has continued. Huge disparities remain in geographical coverage on parenting and
childhood—the Islamic Middle East is a glaring case in point—but a commitment to a more global
agenda has clearly emerged.
The number and range of historical studies on parenting and childhood have increased steadily
over the past two decades—symbolized and encouraged by the formation of both an association and
a Journal for the History of Childhood and Youth (2008). Attention to the material artifacts of childhood,
to the relations between emotions history and parenting, to fatherhood and also motherhood—along
with the expansion of geographical coverage—mark the growing interest in the field. At the same
time, the specificity of much of the work complicates any effort to summarize for a broader audience
concerned with the phenomenon of parenting. What follows picks up on a variety of specific studies
and analytical complexities, but at the same time ventures some wider generalizations applicable to a
historical approach to modern parenting.
Central Issues in the Emergence of Modern Parenting

While modern parenting, as it took shape over the past 250 years, must not be systematically con-
trasted with premodern patterns, it does reflect several sweeping changes in context and behavior.
Many of these first took shape in the Western world, associated most obviously with the unfolding
of industrialization and its social consequences. They would however gain global resonance, from
the late 19th century onward, although duly adapted to regional conditions. Several leading issues
predominate, to which parents had to respond or, in some cases, in which parents themselves took
the initiative to adapt previous expectations and behaviors.
1. The advent of new rates of birth and death, or what is called the demographic transition. What
was the parental role in deciding on limitations in the birth rate, and what impact have these
limitations had on parental behaviors? What was the result of the unprecedented decline in
infant mortality levels, as parents in a growing number of societies began to assume that the
death of one or more children was in fact unlikely?
2. How did parents accommodate the shift from work to schooling as the children’s principal
social obligation? Here was a common trend, in which differences in social class still loom large.
3. How has parenting adjusted, or failed to adjust, to the substantial separation between the family
and work? This change initially prompted considerable differentiations between mothers and
fathers, at least in several societies. It has more recently, again in many cases, been compounded
by new patterns of women’s work. Even in some of the oldest industrial societies, the process of
adaptation continues.
4. How have parents reacted to the advent of new agents in dealing with children, and the poten-
tial for a reduction in parental authority? Governments, most notably, played little role in dealing
with children until the modern era, but they now become a major force—with the obvious
result that parents often spend some time serving as intermediaries between their children and
external authority. In many societies as well, new kinds of expertise developed, providing advice
to governments and families alike about what parents should be doing or what their deficiencies
were. Here was another important challenge to parenting that, in some cases, may have under-
mined traditional levels of confidence.
5. How do parents handle children as consumers, and the increasing array of media and commercial
influences on children? Here is another vital modern topic, gaining growing force over the past
century. It adds another set of new agents—sales outlets and advertisers—with which parents
must deal, but it warrants special attention in adding to the socialization roles of parents as well.
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The confluence of factors shaping modern parenting, or shaped by modern parenting, provides
some coherence to major developments over the past 250 years, unfolding successively in most of
the world regions. The same framework helps organize some of the leading comparative issues, and
offers some important intersections between historical analysis and other disciplinary approaches to
the tensions and rewards of modern parenting.
The Demographic Transition

The vital movement away from the pattern of high although not maximal birth rates, and high infant
death rates, which had been characteristic of agricultural societies, is familiar enough. The implica-
tions for parenting, however, merit special comment, as part of a characteristically modern set of
changes, and some deserve further exploration.
In Western Europe and the United States, some groups began to cut birth rates as early as the
1790s—Philadelphia Quakers, for example. The trend would spread fairly quickly within the urban
middle classes, and also among property-owning American farmers; it spread to the working classes
a bit later, and to peasants and agricultural workers last of all. The process extended for a full century,
generating, by the early 20th century, an average family size that had dropped from 6–8 children to
2–3. It would be briefly interrupted by the baby boom of the mid-20th century (though this was
particularly pronounced in the United States), after which the reduction resumed.
Motivations for the dramatic new demographic structure were clear, if gradually realized: Chil-
dren’s work value went down with the advent of more complex machinery and organizational struc-
tures and early child labor laws. The challenge was compounded by the fact, as British working-class
parents began to realize by the 1820s, that sending a child off to factory work, to be directed by a
stranger, was quite different from traditional, parentally arranged apprenticeships, which promoted
new concerns among many parents (Smelser, 2005). At the same time, the desirability of formal
education went up. In the process, children increasingly shifted from serving as contributors to the
family economy, to becoming cost centers—and the need for new levels of birth control followed
inexorably (Livi-Bacci, 1997; Seccombe, 1993).
During this process in the West, infant death rates initially remained fairly high, although they
may have declined a bit. But a massive transformation occurred between 1880 and 1920, dropping
infant mortality from about 30% of all those born to 5% or less. New public health measures were
most obviously responsible, including government clinics and advice literature like the American
pamphlet Infant Care, launched in 1906. Application of the germ theory, after a decade or so of
adjustment, improved hygiene at birth. Parents began to be able to take some new protective meas-
ures directly. But overall improvements in living standards also played a role, completing the basic
transition early in the 20th century.
Similar patterns elsewhere featured a slightly different sequence and chronology, but shared results.
Japanese reforms, after 1868, included wide imitation of Western public health measures, and pam-
phlets for parents on health and hygiene emerged after 1900. Russian revolutionaries moved sur-
prisingly quickly, given the chaos that surrounded them after 1917, to set up medical services for
pregnant women and young children, and this triggered a significant drop in infant mortality. Similar
efforts followed the Chinese communist victory in 1949, although with a bit more reliance on
traditional regional medicine and “barefoot” doctors. By the later 20th century reductions in infant
mortality became a global trend, excepting only societies locked in warfare or suffering extreme eco-
nomic and agricultural collapse. Whatever their ideology, governments, backed by global health and
humanitarian organizations, realized that they had a vital stake in improving children’s health, if only
to create a sounder labor force and, in some cases, a more abundant military recruitment pool—and
many parents quickly responded (Stearns, 2016a).
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Peter N. Stearns
Birth rate changes came more slowly in some cases, occasionally against government opposi-
tion. Here, parental decisions were even more crucial. Thus Russian urbanites, faced with familiar
early industrial pressures including scarce housing, were cutting their birth rates by the 1930s. Latin
America followed suit between the 1960s and 1990s, with countries like Mexico dropping from 6.5
children per family to 2.5 during this short span. Only parts of Africa and the Middle East have yet
to complete this part of the transition, although rapid reductions in Iran and elsewhere anticipate
the probable trend.
What did this vital transition impose on parenting? First, obviously, parents at some point had to
realize that traditional birth rates had to be modified. Only in a few cases, most notably China after
1978 where the state took a direct role, were larger organizations immediately involved. Parental
decisions might be based simply on budget realities, but they might also reflect new kinds of mobility
aspirations from children themselves. A Mexican mother, explaining in the 1970s how she cut her
birth rate despite opposition from her husband and her priest (who accused her of sexual immoral-
ity), simply stated: “For me, family planning is very important . . . I don’t want my children to grow
up like me, with scarcely any schooling” (Huston, 1979, pp. 82–83; Coale, 1973). The overall global
correlation between improved educational access for females, and birth rate reduction, suggests a
similar role of changing expectations, at least among women.
Were parents, and particularly fathers, also challenged by the new constraints? A few historians
have tried to suggest that some men might find birth control a blow to their sense of masculinity
(particularly if as a result they had no male heirs). Even women who actively participated in birth
control might experience some uncertainty about their maternal role, particularly when their behav-
iors contradicted what their own mothers had experienced and raised religious concerns as well.
The most obvious question about parenting and the demographic transition, however, involves
the potentially more intense interactions between parent, or at least mother, and child once the birth
rate dropped and the need to anticipate some infant death fell off as well. Older children were now
less available for help with infant care. In the West, the later phases of the birth rate drop coincided
also with a decline in household servants and a growing residential split between older parents and
their adult children. All of this could add up to some new parental responsibilities, between birth and
schooling, but now applied to a smaller number of offspring.
Quite possibly the same transition measurably heightened the emotional investment many parents
now made in the individual child, leading to the creation of what one sociologist, examining the
United States at the end of the 19th century, has simply termed the “priceless” child (Zelizer, 1994).
New levels of attention, more elaborate consumer purchases for children, and the introduction of
novel practices (like providing an allowance) all suggested a changing economic and affectionate cal-
culation about parental obligations and opportunities. Recent developments in China offer another
example of increased emotional attachment to the individual child: See the elaborate arrangements
many institutions make to help ease the separation between parent and (only) child on college entry,
complete with tent complexes in which parents can stay until they feel ready to make the break.
The new demographic equation, finally, in combination with some wider developments, such as
the expanding role of medical doctors, almost certainly increased the responsibilities most parents felt
for child health care. What we know about parenting at least in Western society before the 19th cen-
tury suggests some gaps in this regard: Measures (like covering water wells) that might have inhibited
accidents were not usually taken; doctors were not normally called in for a childhood illness (in
contrast to what was done for adults). We know that parents grieved when their children died, but
they may not have felt that there was much to be done to prevent their sadness.
During the 19th century in the West, middle-class culture, as evidenced for example in the new
genre of women’s magazines, increasingly insisted that good parents should be able to prevent the
needless waste of infant lives (Branca, 2014). The same argument infused the government pamphlets
aimed at correcting mistakes in hygiene and feeding. The same goals informed the community
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clinics of communist societies. The same goals, finally, spread to schools themselves. By the early
20th century in the United States, school programs (backed by soap companies) carefully instructed
children on the need to bathe and brush teeth regularly, even when (it was assumed) their immigrant
parents needed some prodding (Vinikas, 1992). Accompanying all this, at least in the cities, was the
emergence of pediatrics as a medical specialty and the advent of more elaborate childrearing advice
literature, in some cases authored by pediatricians directly as with the renowned Dr. Spock after
World War II (Spock, 1946; Zuckerman, 1975). Whether parents learned new responsibilities on
their own, increasingly aware that, now that infant death was a rarity, its actual occurrence would
be more devastating than ever before, or whether outside agencies or influences predominated, an
increasing amount of parental attention was now being devoted to daily hygiene routines, medical
checkups, advice consultation, and possibly greater attendant anxieties.
Extension of parental commitments and expectations around children’s health opened up one
further domain, particularly from the 1920s onward: a growing concern about accident prevention
(Tarr and Tebeau, 1996). The advent of the automobile brought a host of new concerns and, ini-
tially, a rash of accidents that disproportionately affected the young. Increasingly complex household
equipment, requiring electricity outlets and machine operations, and a growing list of household
chemicals posed obvious challenges. Responses were both social and parental. New rules sought to
discipline automobile drivers, and train the young appropriately. Corporations were pressed toward
more responsible efforts at accident prevention around home appliances. But parents—and again,
often particularly mothers—were also urged to exercise new caution, to watch children more care-
fully and train them toward greater safety consciousness. Faced with the complications of a mecha-
nized society, it was easy to see children as more vulnerable than before, with the expansion of
parental responsibility a vital compensation.
In sum, the demographic transition invites attention to the kinds of parental decisions that went
into the process of birth and death rate reduction and to the potential impact on their own expecta-
tions. It could potentially affect the love and attention provided to individual children, and it almost
certainly extended parental responsibilities in hygiene and health. The parent of an unusually small
number of children, expected and expecting to see them all survive, was a new phenomenon in sev-
eral respects. The new pattern also highlighted anomalies: The parents who had more than a small
number of children might now draw new kinds of social scorn. Parents who still endured the death
of a child faced new, often almost unendurable, levels of guilt and sorrow (Stearns, 2007).
Work and the Family: A New Challenge

Famously, with urbanization, the rise of factory industry, and other innovations such as the depart-
ment stores (first introduced, in Paris, in the 1930s), work moved out of the home, where it had
lodged—for most peasants, artisans, and small shopkeepers—literally for centuries. The transition
could be surprisingly abrupt. In 1800 only 5% of New Yorkers worked outside the household, but
by 1840 the figure had risen to an astonishing 70%. By the latter date the word “commuting” was
being added to American English, based on the commutation of train ticket prices for regular cus-
tomers. Work hours themselves would remain extensive for at least several more decades, and now
travel time had to be added in as well. The result, ultimately in all industrial societies, was a vital
new question: Who, now, would be available to take care of household and kids? (Hindman, 2002;
Stearns, 2016a).
Answers have varied, both regionally and over time. In both Western Europe and the United
States, and later Japan, initial response involved substantial withdrawal of mothers from the formal
labor force, in favor of home duty, supplemented perhaps in the middle classes by help from a domes-
tic servant. Gender divisions in parenting unquestionably expanded in consequence. The idea of
relatively distant fathers was hardly a modern one, but it took on new dimensions and became more
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widely applicable when fathers spent the whole day at work, often seeing children at most fleetingly
except on weekends. In Western society, particularly during the 19th century, a new cultural valu-
ation of mothers both responded to the new situation, and exacerbated it. Mothers were held up as
ideal caregivers for children, naturally capable of providing the loving atmosphere that, it was increas-
ingly argued, children required. Of course there were burdens attached: Mothers were supposed to
exercise increasing control over emotions, like anger, that might damage the innocent child if applied
to discipline. And the same maternal ideology increased the disapproval of “bad” mothers, including
lower-class mothers, who could not seem to measure up. But the maternal emphasis was very real,
pushing some mothers, at least, to increasing investment in their children well into the 20th century
(Badinter, 2012; Ryan, 1983). Japan, for its part, emphasizing the importance of training women to
be “wise mothers”, generated its own version of maternalism in parenting. Both in the West and
Japan, educational emphases, around subjects like “home economics”, sought to prepare girls for
their maternal roles, as a gendered part of the development of modern mass education.
The same push toward maternalism—both cultural and economic—raised questions about what
fathers were for, beyond serving as “breadwinners”, as the 19th-century phrase urged, supporting the
family economically. Some families sought to retain a special disciplinary role. “Wait till your father
gets home” became a standard admonition to misbehaving offspring, and it had some basis in fact,
with mothers emphasizing a softer side, fathers expected to anchor more rigorous discipline. Some
U.S. studies suggest that, by the later 1940s, American mothers were taking discipline more directly
into their own hands, but the expectation of paternal sanction persisted in some families (Griswold,
1993; Stearns, 1990).
Beyond this, a variety of movements—perhaps as early as the late 19th century, certainly recurring
from the 1920s onward—sought to involve fathers a bit more directly: by this point, the question
of how to bring fathers back into the parenting picture had real urgency, after decades of maternal
direction. Reductions in the hours of work, among other things, created important new opportuni-
ties. The childrearing literature began to mention fathers more frequently than had been the case in
the mid-19th century. Several movements, for example in scouting or the YMCA, sought to develop
new roles for fathers as guides to children’s activities or, as one organization explicitly urged, simply as
“pals”. By the post-World War II era, at least in the United States (in some European countries, fathers
took longer to shake off more traditional formality with their offspring), attempts to build comrade-
ship around shared entertainment gained ground, and some children, clearly recognizing the greater
importance of their mothers, nevertheless reported that dads could be more “fun” (Strange, 2015).
Not all societies greeted initial industrialization with the same degree of gender division that
predominated in the West and Japan. In Russia, rates of women’s employment remained much
higher, both before and during the Soviet era. The same was true later in China. Women were
nevertheless expected to take particular responsibility in caring for children and the household, but
the maternal cultural emphasis was less pronounced, the practical burdens on the mothers’ time
perhaps even greater. Other regional patterns displayed still greater variety. In sub-Saharan Africa,
for example, men became particularly likely to gravitate toward cities or mines in search of better-
paying jobs, leaving many mothers back in the villages, with childcare falling almost exclusively on
their shoulders.
Gender divisions were not the only response to the movement of work outside the home. New
levels of interaction with grandparents might also result. In the West, during the 19th century, older
parents (particularly, mothers of mothers) became more likely than before to co-reside with an adult
child, among other things because they could help provide childcare (Rosenzweig, 2005). This
approach declined from the 1920s onward, as older parents moved out again, but it continued to
apply to some families, as in the African American community. More formal institutional responses
were slower in coming. But experiments with new types of infant care, through nursery schools of
various sorts and then, after World War II, to other types of day care arrangements responded to the
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new work environment fairly quickly, initially often as part of a charitable outreach organized by
elements of the middle class. Communist regimes were quickly active in setting up day care arrange-
ments in the cities. In several cases, however, some groups and families did not accept this kind of
external substitute for parenting (or grandparenting), or simply could not afford the cost. Thus,
acceptance in the United States (where government was far less likely to play a supporting role)
lagged measurably behind European levels by the later 20th century (Druckerman, 2012).
Finally, response to the challenge of work outside the home constituted a moving target. In
Europe and the United States, the huge move of married women back into the formal labor force
during the 1950s and 1960s set up a host of new challenges—even though the motivation, in seek-
ing more income for an improved family standard of living, had its own links to parenting. At crucial
points, polls suggested that a majority of women actually disapproved of work roles for mothers of
young children, even as actual behavior changed in precisely that direction: There was important
tension around the change. But by the 1970s, almost half the labor force in Western societies was
female, many of them mothers. Responses to this new pattern included some new reliance on
fathers, although gender divisions remained challenging; further birth rate reduction; greater use of
institutional care; and use of other relatives. In many cases as well, a clear further result involved the
harried mother, still expecting to maintain disproportionate responsibility for childcare, sometimes
curtailing career expectations in the process, but rushing nevertheless to keep work and parenting
in some balance. The finding that, in the United States by the 1990s, the most likely driver to run a
red light was the “soccer mom” rushing home to chauffeur children to their various activities, was
more than symbolic (Peskowitz, 2005). For up to two centuries, in many societies parents have been
grappling with the inherently difficult response to the redefinition of the relation between work and
family.
Parents of Students: The Rise of Schooling

Parental responsibilities for providing practical training for their children are hardly a modern dis-
covery, and many cultures had long emphasized obligations to provide moral upbringing. However,
the spread of more universal and arguably more demanding school requirements ultimately required
some adjustments in parenting. Parents had to allow outsiders—teachers—to play a greater role with
their children, often facing some values conflicts. Often, they might need to accept the idea that,
through schooling, their children might move into types of jobs that differed from their own, another
potentially challenging modification of most traditional parental goals. The modern emphasis on
mobility through education involved real shifts in parental expectations, toward at least a vague idea
of one’s children doing “better” than oneself.
The growing emphasis on schooling depended considerably on parental assent; this was, in many
cases, the first adjustment to be made to “modern” conditions, preceding new birth rate decisions.
Even when new laws required attendance in principle, lax enforcement often left an important role
for parents at least for several decades. On the whole, middle classes found it particularly logical to
support schooling as part of good parenting—this affinity may indeed be a defining feature of the
class, even in communist societies, on a global basis. Some working-class parents, however, were ini-
tially suspicious of schools, including immigrants in American cities who worried that family values
and religious interests would be challenged by the new public system. In France, peasants held back
from full embrace of education until the 1860s and 1870s, at which point they increasingly decided
that schools now made sense for boys, because literacy and numeracy would support their com-
mercial farming interests, and for girls because they might prove the basis for careers in areas like
teaching (Weber, 1976). On a global basis, the parental debate has continued into the 21st century.
Thus, an Indian child labor reformer in the 1990s confronts a cobbler working with his son outside a
school, asking why he did not allow the boy to join the class: “Young man, my father was a cobbler
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and my grandfather before him, and no one before you has ever asked me that question. We were
born to work, and so was my son” (Shukla, 2007). Obviously, increasing numbers of parents did come
to agree that schooling made sense, and they were often pressed by government enforcement; but a
parental transition was vital.
As schooling spread, it prompted additional changes in parental behavior, beyond decisions about
whether to support school attendance in the first place. The treatment of young children was affected
by the need to ready them for school, although families responded variously in part according to social
class. As early as the late 18th century, in the West, conscientious middle-class families were buying
newly available children’s books and other educational materials to help pave the way. Special lessons
might anticipate the school experience, requiring parental arrangements; it was the rare middle-class
girl who, by 1850, did not have some instruction on the piano. By the 20th century, parental respon-
sibility for school preparation might expand, for example in helping to arrange some formal sports
experience. Preschools of various sorts spread widely; by the late 20th century, up to 90% of children
in countries like Japan were enrolled. Parental arrangements or supervision did not take over the
whole arena of children’s play, but they made important inroads particularly in the growing cities.
Parents, and often particularly mothers, retained a vital supporting role once schooling began. In
Japan and Korea during the 20th century, increasing maternal investment reflected the importance
of school success in measuring parental adequacy. Mothers, often working part-time if at all to assure
their availability, put great care into preparing school lunches (bento) and in organizing recreations to
compensate for the rigor of school and homework. They worked hard to develop the kind of intense
emotional relationship with their children that would in turn provide strong motivation for school
success. Shame was directly applied to support academic achievement where necessary. Maternal
oversight continued with strong involvement in preparations for crucial examinations and in direct
representation in areas such as selecting a university (Yamamoto and Holloway, 2010).
Regional and social class variations remained important. In East Asia, by the late 20th century,
parents expected no regular contributions from children in household chores, in favor of concentra-
tion on homework and related educational activity. In the United States, chores also declined, partly
because of changes in household requirements (including care for younger children), partly because
parents increasingly found it more efficient to do the work themselves, partly because of increasing
resistance particularly from adolescents. But some ambiguity and resentment persisted. At the same
time, U.S. parents enforced shorter periods of home study than was true in East Asia (Stearns, 2003).
Class differences reflected distinctive orientations as well. By the late 20th century, British working-
class parents were as likely as their middle-class counterparts to express hopes that their children
would become doctors or lawyers. But they put far less time into school-supporting activities or to
preschool preparation, leading to some disjuncture between aspirations and outcomes in many cases
(Goldthorpe, Lockwood, Bechhofer, and Platt, 1969).
The extension of schooling into secondary levels and beyond, a gradual process beginning in the
19th century and then solidified by longer school requirements after World War II, affected parent-
older child relationships. Children’s dependence was in many ways prolonged, and this could lead
to some interesting negotiations and tensions. The idea of adolescence, surfacing first in Western
Europe and the United States in the 19th century, partly reflected concerns about behavior of this
age group once it was largely cut off from formal work (Kett, 1977).
Finally, the spread of schooling everywhere increased the importance of peer group interactions,
in ways that might compete with parental authority. Signs of “youth culture” began to emerge in
the West by the late 19th century, and it would extend on a more global basis after World War II,
fueled among other things by new communications technologies. Later childhood became increas-
ingly associated with musical and other tastes deliberately different from parental preferences, and
sometimes deliberately defiant, as an expression of the gap between peer group loyalty and parental
standards.
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Many parents negotiated the implications of the new relationship between children and school-
ing quite successfully; in no sense was there a widespread crisis. But changes in responsibilities were
significant, often bearing particularly on mothers. Schools, plus attendant peer group interactions,
set up potential tensions with parental authority that had to be handled carefully. In some cases, as in
the United States from the early 20th century onward, one result was a more intense focus on early
childhood as a vital period, relatively free from outside influences, where parents could instill values
that might, it was hoped, last lifelong.
Consumerism
By the late 19th century, it was becoming clear, in the urban, industrial societies that many parents
also had to develop explicit approaches to the rise of consumerism. As in other areas, several types
of changes might result. Consumerism—the opportunity to buy or expect unnecessary objects or
entertainments as part of an enjoyable life—had not been a significant parental issue in premodern
societies, outside the upper classes. It now became a standard experience, inevitably affecting parent-
child relationships.
There is no indication that consumerism raised specific new problems in parenting until the late
19th century, but even before this in the industrializing West some new parental spending developed.
Parents, where incomes permitted, increasingly sought to express their feelings for children through
purchases and gifts. Christmas became a steadily more elaborate opportunity to give presents to the
children, building on more modest prior traditions but also removing some disciplinary features of
the earlier holiday—such as the idea of punishing bad behavior. Children’s birthdays became more
extensive as well—in what had, in most cultures that used the idea of birthday at all, previously been
largely a celebration for adults or directed only toward honoring religious figures.
New interests in family consumerism emerged by the early 20th century. Opportunities to buy
goods particularly made for children expanded, whether the item involved toy soldiers for boys, more
elaborate dolls for girls, or the new category of stuffed animals for babies. Some debate ensued, par-
ticularly around indulgence for infants, but experts and certainly parents increasingly agreed that it
was good to give infants some additional targets for comfort and affection and to begin to habituate
them to the idea of acquisition (Cross, 1997; Jacobson, 2004). By this time also, approaches to envy
were being reconsidered, at least in the United States; what had been long regarded as a sin, with
children urged to control their impulses, now became (up to a point) a virtue, in encouraging higher
consumer aspirations (Matt, 2003). It was at this point as well that middle-class parents began the
practice of giving “allowances” to children, to provide experience in budgeting but also to encourage
some direct ventures into the world of consumerism. Parents (and advice-givers) in many societies
periodically worried about spoiling children and encouraging unhealthy levels of commitment to
materialism, but it was mainly up to parents themselves to decide on appropriate guidelines.
More pressing concerns developed about consumer influences that went beyond parental con-
trol. With wages or allowances, some children could now buy items on their own, and this change
began the process of constructing an explicit consumer market that attracted a host of entrepreneurs.
Cheap, thrilling novels designed for adolescents (called “penny dreadfuls” in Britain) morphed into a
first wave of comic books, sold to children directly unless parents intervened. In response, self-styled
purists like Comstock, in the United States, urged parents to mobilize against sales of this kind of
material to impressionable young minds; violence was the clearest target, but there was concern
about sexual stimulation as well. Thus, a sequence of tensions began between at least some parents
and the purveyors of commercial fare—particularly in the United States, where public regulation
of media was particularly limited. Radio shows for children, then comic books again in the 1950s,
then television fare, and then the internet and video games—each succeeding technology, threaten-
ing to bypass parents—drew its critics, including many righteous parents themselves (Gurstein, 1998;
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Starker, 1989). More widely, parental efforts to channel children into more approved fare—such as
Disney materials, which began to emerge from the 1920s onward, offered a more positive alterna-
tive to the greatest commercial dangers. Most parents in consumer societies renounced any effort to
censor children’s entertainment in undue detail, although there were always a few holdouts for no
comic books or no television at all. Many, however, took on a new responsibility for sorting through
the available offerings, in the interests of appropriate caution and guidance.
This tension was accompanied by a growing commitment to helping children learn the ways of
consumer behavior—taking them shopping, giving them some range of opportunity to select items
like clothing. Occasions of this sort could produce shared pleasure, but they could also be sources of
quarrels over appropriate styles, particularly when parental and peer group standards clashed. In some
cases—perhaps most clearly among working-class and immigrant families, children by adolescence
saw consumerism as a means of separating their identity from that of their parents, and this distinc-
tion could complicate the relationship. The parent as consumer, instructor, or sometime ineffective
monitor was an interesting new role ( Jacobson, 2004). It could blend with other new interests, such
as the need to provide an outlet for children who were being pushed toward greater school success,
as in Japan.
Finally, in the world of consumer goods and entertainments, in which parents became increasingly
accustomed to showing their affection for children in part through purchases, many parents gained
some sense of responsibility for keeping children actively amused, in what some scholars have called
a new “happiness imperative”. The idea of the bored child is an essentially modern construct. In
the late 19th century, boredom might suggest a character flaw in children themselves; childrearing
manuals and etiquette books urged children to correct the condition by finding ways to become
more usefully engaged. But as consumerism gained ground, by the mid-20th century, boredom more
commonly turned to an accusation against parents or other adults, an indication that they were fail-
ing in their duties to provide stimulation. Arrangements for play opportunities; additional purchases;
acceptance of television viewing or regular trips to movie theaters even amid some hesitations were
some of the methods parents might emphasize in this new role. More elaborately, family trips to
amusement parks, as these emerged in the United States, Europe, and then East Asia, or the growing
custom of the family vacation, helped fulfill this additional parental role with some regularity. The
“entertained” child is another modern phenomenon that gave parents new opportunities for some
shared pleasure, but some new obligations as well (Stearns, 2012).
Parents and Other Authorities

One final feature of modern parenting, gaining increasing currency from the late 19th century
onward, involved parents as intermediaries with other agencies or authorities that either sought
to guide parental behavior or to intervene directly with children themselves. Both schooling and
consumerism, as we have seen, generated new attention as outside forces that impinged on parental
territory. The need to pay some attention to what children were buying or watching drew parents
into some new issues. Parental (often maternal) investment in supporting school success might bring
active engagement with teachers and other school authorities. This could take the form of participat-
ing in various kinds of parent associations, where volunteering extended the parental commitment to
schooling; or it might involve representations that sought to protect children against real or imagined
school abuse. At several points in the 20th century, American parents banded together successfully to
limit the amount of homework assigned (Schlossman and Gill, 2003). By the early 21st century Chi-
nese school authorities reported increasing interactions with parents and grandparents, now fiercely
focused on their small cohort of children and correspondingly eager to make sure they were treated
fairly. In the United States, an intensification of parental concerns created the phenomenon known
as “helicopter parenting” by the early 21st century, where either a mother or a father might actively
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intervene on their children’s behalf to seek a better grade or a better slot on a sports team—the tar-
gets could be varied (Stearns, 2003).
Parent as intermediary, however, also took on additional features. The parental role in dealing
with government agencies, voluntarily or through compulsion, undoubtedly needs more histori-
cal attention than it has received. The parental role in receiving and filtering new kinds of expert
advice about childrearing is more familiar, but by the same token important to note as an essential
innovation.
With the State

From the late 19th century onward, a number of societies—beginning with the West and Japan—
devoted more attention to the policing of youth, particularly in the growing cities. Acts that used
to be tolerated, in the category of minor vandalism, were now criminalized. And at the same time
juvenile delinquency was treated separately from adult crime, with more hopes of rehabilitation. All
this set up a situation in which parents—historians are now speculating, especially fathers—had to
serve as intermediaries with authorities on behalf of their errant offspring. This was a new category
of responsibility, extending parental roles in trying to teach children to obey social rules (Lassonde,
2016). Parents might in fact learn to appeal for police intervention in hopes of curbing a wayward
child—the relationship was not always antagonistic.
As early as the late 19th century, working-class and immigrant parents in major Western cities
were also learning to deal with new agencies such as settlement houses and orphanages. Along with
schools, settlement houses might help some immigrant children adjust to new values, but in the pro-
cess raising some issues of parental control—particularly when, as was commonly the case, children
proved far more adept at learning the national language and other habits. In most industrial societies,
working-class parents became adept at using orphanages, not usually to abandon children outright
but to bridge periods in which parents simply could not maintain adequate support—planning to
restore their offspring to the family when conditions improved.
The expansion of welfare state functions, in this case mainly in the 20th century, added to parental
contacts with the state. Governments began regarding some parents as clearly incompetent, inter-
vening not only in cases of delinquency or truancy, but also where economic or health conditions
seemed to threaten the well-being of the child. By the late 20th century, in places like the United
States or Australia, intervention might even be provoked by family obesity or tobacco use. In some
instances this approach involved taking children away from parental care, but in others the state
moved in primarily to redress parental behavior (Sandin, 2016). Single and working-class parents
were disproportionately involved in these new relationships. Both communist and capitalist govern-
ments increasingly defined some standard criteria for the treatment of children, and when these were
not met parental behavior had to be modified. In some cases, extensions of ideas about children’s
rights could give children themselves some voice in this equation as well. Here was an important
complication of parental authority that warrants further analysis as part of modern parental history.
Historians have paid some attention to the overall issue of child abuse, which was an arena where
some parents might increasingly be required to deal with the intervention of state authorities (by
2016, up to three million reports of some form of abuse were being handled annually in the United
States). We know that social attention to child abuse has fluctuated, with periods of crisis yielding
to a surprising degree of inattention. Some scholars believe that rates of parental abuse have risen
in the modern centuries, because of a decline in community monitoring, more broken families,
and more confusions over what children are for. At the same time, however, standards applied to
parental behavior have become more rigorous, making some actions—including extensive physical
discipline—seem abusive that once were perhaps more widely tolerated. Overall evaluation is diffi-
cult, but it bears directly both on a historical understanding of modern parenting and on the specific
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issue of a new need for some parents to serve, however reluctantly, in interactions with government
officials (Pleck, 1987).
With the Experts

Parents had long been the target for advice. In many premodern societies, religious authorities were
the primary source, urging parental responsibilities for moral upbringing and religious training and
sometimes thundering against wayward families. In some regions the decline of religious fervor
by the 19th century supported a less denominational approach to childrearing, with more general
advice emanating from self-appointed moral authorities, often women.
The early 20th century, however, saw a new kind of advice literature, increasingly emanating from
scientifically trained experts or their popularizers. Pediatricians wrote widely about children’s health.
In some cases, government-sponsored pamphlets gave an official stamp to this new literature. The
pamphlet Infant Care in the United States, first published in 1906 and issued in regular new editions,
became the most widely circulated government pamphlet through the 20th century. By the 1920s,
this new advice literature branched out, both in topic and in venue. Not only books, but also maga-
zine and newspaper articles now carried the latest wisdom about caring for children’s emotional as
well as physical well-being. Whole journals, like the American Parents’ Magazine, launched in 1926,
expanded the genre still further.
Expansion also involved the range of topics covered. Health and hygiene remained central, but
psychological issues gained growing attention. In the 1920s for example, the dangers of sibling rivalry
were highlighted in a good bit of the new childrearing literature—a topic that had been largely
ignored in the 19th century. The new wisdom, derived in part from some social work experiments,
held that young children might be consumed with jealousy at the arrival of a new sibling. The situ-
ation was dangerous to the new baby, for toddler jealousy could provoke violence, but also to the
socialization of the toddler himself or herself, for successful adults must learn to handle emotions
of this sort appropriately. Parental intervention might be vital, and a host of suggestions followed
concerning corrective measures (Stearns, 1989). Children’s fears and grief were other emotional
categories winning new attention.
The United States was a disproportionately important source of this new kind of childrearing
literature, but there were echoes, or direct sales and translations, not only in Europe but in Japan,
Latin America, and elsewhere. Dr. Spock’s parenting manual, for example, The Common Sense Book of
Baby and Child Care (first published in 1946) was translated into 39 languages, which means in turn
that some parents outside the United States now needed to make some decisions about whether to
learn from this kind of international influence.
Quite apart from specifics, the new literature rested on two common assumptions. First, experts
knew better than parents, at least in many cases; parents could not be sure that their own intuition,
or the patterns they had inherited from their own parents, were satisfactory. Second, expert advice
often changed fairly frequently—for example, from the strictness of the behaviorists in the 1930s
to a slightly more permissive approach with Dr. Spock and others after World War II. These shifts
could further confuse well-meaning parents and potentially increase their dependence on the latest
expertise.
Did the new literature suggest a growing lack of confidence among parents themselves, and/or
did it promote a decline in self-assurance? Lasch (1979), among other historians, has seen the rise
of dependence on external experts as a sign of a real crisis of parental authority. Certainly further
urbanization and the increasing separation of nuclear families from older relatives created a situation
in which new kinds of advice might seem essential. Government support for the new expertise, and
often a sense that parental guidance itself must change to prepare children for changing economic
and social roles, added to the pressure. Many parents, of course, blended their use of advice with their
334
own assumptions and values. On the whole, urban middle classes provided the most avid consumers
of expertise; social differences were important in this category as well. But the new requirement, for
conscientious parents to take external advice into account and to sort out its applicability added an
interesting further twist to modern parenting.
The Modern Parent

Without becoming entirely new animals, modern parents differed from their premodern counter-
parts in several ways, though the changes in some cases emerged only gradually and were marked
not only by personality differences but also by distinctions by social class. Modern parents had to
make decisions about birth rates that would gradually differentiate them from the patterns their
own parents had pursued. They had to figure out how a numerically reduced role would affect their
larger commitments as parents and their overall self-perception. They developed a greater stake in
the physical and (probably) emotional health of the individual child, taking some new measures in
consequence. Modern parents had to figure out how to care for children when one or both parents
were away at work. This problem was eased but not eliminated by the birth rate reduction. It might
lead to new gender divisions (and quarrels), to new dependence on other relatives, and to new
acceptance of state agencies.
Modern parents clearly came to accept the reality of formal schooling at least to some degree
but many played a deliberately active role in converting to a sense of responsibility for schooling
and to new activities in preparation and support. The ubiquity of consumerism prompted additional
adjustments in training children, in tensions over taste, but also in using consumer goods to motivate
and to express affection in novel, sometimes problematic ways. Finally, modern parents had to make
decisions about dealing with external authorities—consumer agents, the state, and self-styled experts
all impinged, requiring some explicit reactions or passivities.
These general patterns came to have global applicability, although with marked differences in tim-
ing. They affected communist as well as capitalist societies—indeed communist leaders were quick
to begin to install their version of modernity as a framework for parenting soon after the revolution-
ary takeovers. To be sure, the most elaborate evidence about parental adjustments comes from the
Western experience, but the transitions have not been Western alone, and they have guided relevant
research on the history of parenting in a number of other regions ( Journal of Social History, 2005).
At the same time, attention to the largest developments that have shaped modern parenting, even
with some allowance for differences in timing and regional specifics, cannot possibly capture all the
essential elements in the history of modern parenting or do justice to the widening scope of research.
The following section turns to several case studies that highlight additional regional and comparative
experiences. The review is hardly exhaustive, but it suggests additional components to the historical
record that amplify, or push against, the larger patterns. The goal is to add some sense of the range of
historical cases and differentiations that must be added in any overall approach to the unfolding of
modern parenting. Five brief sketches add diverse variations on modern parenting, suggesting several
distinctive issues and analytical targets.
Regional Case Studies: Modern Parenting in Crisis, Continuity

Amid Change, Comparative Dimensions
Regions and Groups in Crisis: The 19th Century

By far the most important category to explore, outside the general parameters of modern parenting,
involves those many situations over the past 250 years where poverty or disruption shaped dominant
parental concerns and capabilities.
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During the 19th century, for many groups new or extended forms of exploitation overwhelmed
any real contact with issues like modern forms of birth control, much less consumerism, forcing
distinctive kinds of parental adjustments. In many parts of the Americas, for example, a significant
minority of people had to manage parenting under the conditions of slavery well into the modern
period. To be sure, the disruptions of the transatlantic slave trade, which had torn so many African
children from their parents, eased after 1808. But internal slave trading actually increased in conse-
quence, exacerbating the problem and the fear of family separations. Because of sales that split a slave
family or other circumstances (including rape by a member of the White slave-owning family), up to
a half of all slave families in the United States faced the absence of the father. In response, many slave
families developed a special reliance on the authority of the mother, often supported by her female
relatives (Gutman, 1977).
Parenting among American slaves was also strongly conditioned by poor material conditions.
Slave owners paid remarkably little attention to slave diets, even for potential mothers, with the result
that infant death rates were high and undernourishment of young children even more common.
Children had to be weaned early—at 3 or 4 months, so the mother could get back to work—and the
ensuing diet was typically inadequate. Parents also had to try to mediate on their children’s behalf, but
in fact heavy work obligations and frequent punishments—including whippings—were unavoidable.
Parents often managed to help carve out a bit of space for children’s play, and a surprisingly large
minority—up to 10%—also managed to teach their children to read and write, despite the com-
mon opposition from the slave owner. Careful slave parents also taught their children how to display
protective deference to Whites. Boys were taught to “bend their body forward with head down, and
rest the body on the left foot, and scrape the right foot backward on the ground, while uttering
the words ‘how do Massie and Missie’ ”. At the same time, songs and stories, like the Br’er Rabbit
sequence, were intended to provide a cultural outlet for children, while also teaching intense loyalty
to the slave community as a whole. This was, clearly, a distinctive parental experience, and elements
of the adaptations might persist even after slavery itself was abolished (Mintz, 2004).
In the “settler societies” of the 19th century—the United States, Canada, and Australia most
obviously, although elements of the pattern also developed in Latin America—indigenous parents
continued to face some special burdens as well. Increasingly driven off the land and into special
enclaves, many indigenous people sought to maintain traditional cultures in an unprecedentedly dif-
ficult environment. Even worse was the recurrent tendency of the dominant majority to seek ways
to impose new habits in ways that interfered directly with parental control. Schools on indigenous
reservations often sought to limit parental influence. Even more dramatic was the removal of chil-
dren outright, to be sent elsewhere to be reared and “civilized”. Removals were particularly severe in
Australia by 1900, particularly affecting girls. The children involved were sent to distant parts of the
country, and their names were changed; contact with and even discussion of the home family were
strictly forbidden, as was use of the native language. This type of seizure, and the even wider fear of
seizure as an element in indigenous parenting, persisted well into the 20th century.
Indentured labor, widely introduced in European-controlled colonies in the late 19th century as
a substitute for the now-abolished slave systems, shaped parenting in important ways as well. Many
indentured workers, primarily from various parts of Asia, did not understand what they were signing
up for, or their lack of control over their own lives during the period of indenture. As with slavery
itself, material conditions were harsh, and child labor was expected from the age of 5. Britain abol-
ished this particular system only in 1917.
A host of situations, in other words, strongly conditioned parenting in the 19th century, requiring
modifications of traditional patterns with little or no contact with the modern parenting trends that
were beginning to take shape particularly in the West. Other situations could impinge as well. In
most European colonies, in the age of imperialism, most family situations were relatively untouched,
as Europeans had neither the interest or the means to interfere with established parental patterns.
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In a few cases, however, parents were pressed to send their children into domestic service, in White
households, or into factory or mine work, which would effectively reduce parental authority at least
over older children.
Huge questions can also be raised about working-class parents even in the industrializing centers
themselves, where, overall, modern parenting patterns were emerging. Many workers, often immi-
grants at least to the city, conditioned by poverty and demanding work outside the home, sought to
cling to key parental traditions in difficult circumstances. In fact, growing poverty in the countryside,
in the first decades of industrialization, often forced some painful adjustments, as in Japan where
many rural parents essentially sold teenage daughters into the silk mills simply because of economic
necessity. Even after conditions began to ease, many working-class parents continued to expect their
children to work, contributing wages to the family economy, often from a fairly early age; and in part
because of these expectations, they were often relatively slow to simplify their parental burdens by
cutting the birth rate. They also faced the increasing efforts by the state to require schooling, which
raised new concerns both about costs and about potential conflicts with parental values. Gradually,
of course, adjustments were made. Workers themselves, in places like Britain, contributed directly to
the new laws restricting child labor, as they realized how different children’s work was in the factories
and how much direction by strangers interfered with parental authority. Still, a habit of relying on
income contributions at least from older children persisted in many cases, as did some discomfort
with new demands like schooling. Exploring the ongoing impact on parenting of working-class and
immigrant encounters in the 19th and early 20th centuries must be added to the overall assessment
of modern adjustments (Cunningham, 2006; Heywood, 1988; Humphries, 1995).
The 20th and 21st Centuries

A variety of hardship situations continued in the more recent phases of the modern period, even
though not only formal slavery but also special systems like indenture were widely abolished. Harsh
working conditions as industrialization spread challenged parents in many ways. In late 20th-century
China, for example, the rapid expansion of the urban labor force found many parents leaving their
children back in the home village, to be cared for by grandparents, with only periodic visits on occa-
sions such as the Chinese New Year. In many regions, sheer poverty forced other parental decisions.
Sale of children for sex slavery or other forms of exploitation was one outcome, as were efforts to
market some body organs for use in transplants. With an estimated one-sixth of the global popula-
tion still in dire poverty in the early 21st century, parental survival strategies could be brutal.
Particular attention now focused on parental adjustments due to migration and conflict. From
World War I onward a growing number of people were displaced either by wars directly, or by
border adjustments after war (such as the relocation of over one million Greeks from Turkey after
1923, or the massive number of displaced persons fleeing after World War II), or by more recent
civil conflicts in parts of Africa, Central America, and the Middle East. Important migrations, includ-
ing accumulations of refugees, have also resulted from flights from poverty and famine, particularly
although not exclusively in Africa (Scholvin, 2016).
The experiences involved are not always distinctively modern: Conflicts and famines affected
parents and children earlier in history as well. But the scale of displacement has expanded greatly.
Boundaries between civilians and the military were progressively blurred, from the 1930s onward,
which increased the direct impact of war on parenting. We increasingly realize the disproportionate
involvement of children in many of these disruptions, and with this the special burdens on parents
as well.
It has been estimated that 150 million children have been crippled or maimed as a result of
regional wars since the 1970s (along with an equivalent number killed outright; Goodenough and
Immel, 2008). In some civil conflicts, as in Cambodia in the 1980s, where children were often
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deliberately killed in front of their parents, or in some of the African struggles, children have been
explicitly and disproportionately targeted. Millions of children have also spent long periods in the
many refugee camps established, usually in constrained material conditions, to handle the various
kinds of refugee populations fleeing conflict or hardship. Many conflicts have also seen masses of
children separated from their parents, at least for a time: Separation involved over 100,000 children in
Rwanda in 1994, although aid workers were able ultimately to arrange some reunions. Even where
parents and children, or at least a parent and children, have remained together, resulting ill-health or
psychological stress profoundly complicates the parental task. In one refugee camp in the nation of
Georgia, 83% of the children were judged to be suffering from some degree of psychosomatic stress
(Goodenough and Immel, 2008).
Parenting in these situations involves a range of special reactions. Grief must be one, perhaps
particularly intense for parents who initially had indulged a modern expectation that children would
not normally suffer or die. Deliberate decisions about separation can be another: many parents in
stress, for example in Central America or in 2017 Syria, have deliberately sent even relatively young
children away, to attempt entry into a more prosperous region such as Western Europe or the United
States, with no hope of subsequent reunion. Exploitation can be another response of parents in stress.
Sales of children for prostitution or other degrading economic activities become common in many
refugee camps. In the current (2017) refugee crisis in the Middle East, sales of young girls as wives to
older men who already have at least one spouse have drawn attention as well.
Parents in the contemporary period have become increasingly divided not just by familiar divi-
sions of power and wealth, but by conditions of (relative) stability or instability. The spread of indus-
trialization has cut into the most stubborn regional economic divisions, leaving a growing majority
of fathers and mothers actively participating in the key trends of modern parenting. But a large and
fluctuating minority are largely cut off from these trends, either by extreme poverty or by one of
the many recurrent disasters that continue to afflict key regions of the world. Grasping this variety,
and the kinds of desperate decisions still facing some groups of parents, is an essential component of
modern global history overall.
The Challenge of Illegitimacy

Single parenting undoubtedly increased in many modern societies, sometimes as a result of family
dissolution, even more often as a result of a rise in the rate of births out of wedlock. The most gen-
eral cause was the reduction of effective community controls over sexual behavior, supplemented in
some cases by changes in community norms that made single parenthood more culturally acceptable.
Greater commercial activity in the countryside could generate a rise in illegitimacy, but the trend was
more obvious as greater urbanization took hold.
Several factors argue against positing the rise of single parenthood as yet another standard feature
of modern parenting. Most important, the trend has not taken hold in key parts of Asia, where even
today only 5% of births occur outside marriage. Modern social conditions, in other words, may be
compatible with continued emphasis on a more conventional family context. Changes in single
parenthood also reflect other regional factors. Some increase in illegitimacy occurred in Western
Europe in the late 18th century, thanks to growing commercialization and geographic mobility;
but the rates then stabilize in many cases by the late 19th century, and resumed a much more rapid
increase only recently—complemented as well by an increase in divorce rates. Single parenthood
clearly requires assessment in any study of modern parenting, but with appropriate attention to
regional and chronological variables.
High rates of illegitimacy have been particularly marked in the Latin American experience,
beginning in the colonial era with the disruption of indigenous social patterns and the increase in
nonmarital sexual activity by the European colonists. Well before modern times, illegitimacy rates
338
above 20% were reported in many Latin American cities. The pattern continued into the 19th
century and on to the present day. Percentages rose, in some cases, to 30% to 50% of the total. The
pattern was class-specific: Urban middle and upper classes maintained a more conventional family
context, although individual men from these groups might also be involved in other sexual connec-
tions. One result was a pronounced and explicit gap in social standards, with “respectable” elements,
usually linked to the government, expressing shock and dismay at lower-class behavior, in ways that
could impinge on parenting directly (Bartell and O’Donnall, 2001; Hecht, 1998, 2002).
The most important result, for modern parenting, goes beyond the social divide to the inherently
difficult task of figuring out how so many single parents, mainly mothers, managed and defined their
responsibilities. High rates of illegitimacy did not necessarily reflect disarray, although they were
correlated with considerable poverty. Many mothers retained close ties with their offspring—even
children who tried to make a living on the street, often in combination with a network of female
relatives and neighbors. Large numbers of children in Latin America “circulated”—that is, they were
sent to other families that needed labor; exploitation could result, but also incorporation in an affec-
tion family context—results varied. As in the Western working classes, parenting was also sometimes
supplemented by periodic recourse to orphanages—where some children might spend time, not
because their parents were absent, but because parents needed some time to get back on their feet
prior to reclaiming their offspring. Interpreting the various patterns possible amid conditions that
shocked so many middle-class observers is a key element in an overall analysis of Latin American
parenting (Hecht, 2002; Scheper-Hughes, 1992).
The same task applies to the more recent surge in single parenting in the West. By 2009, for
example, 41% of all children born in the United States had unmarried mothers. About 40% of births
in Western Europe fell in the same category (having risen from only 20% in 2000, again reflecting a
very recent intensification), with some national variations (percentages were greatest in the Nordic
countries, lowest in some but not all countries with a Catholic or Orthodox religious tradition) but
a common pattern overall. Australia also experienced a sharp rise in single parenting. As in Latin
America, however (where by the early 21st century rates had in some cases risen to 74% of the total),
social class and associated ethnic differences strongly affected patterns within particular countries
(Macfarlane, 1980; Ventura and Bacharach, 2000).
The rise of single parenthood in the West carried the same warning signs that had long applied
to Latin America. It sometimes reflected involuntary involvement, the result of sometimes fleeting
relationships that had not intended parental responsibilities. The correlation with relative poverty
was troubling, for the tensions associated with single parenting, including difficulty in combin-
ing work with parenting, often perpetuated economic hardships going forward. By contrast, some
recent trends reflect not dislocation, but changes in cultural norms. In some instances, illegitimacy
followed from a decline in the popularity of marriage, not a rejection of parenting, as both parents
in fact participated despite the absence of marital ties. Many single mothers, often using networks of
relatives including their own mothers, worked hard and often successfully to provide attention and
stability for their offspring. Factoring this modern phenomenon into an overall analysis of modern
parenting requires some flexibility, particularly in the recognition that no single pattern of parental
engagement is involved.
Fathers
In most regions of the world, modern parenting has brought increased attention to the roles of
mothers, certainly as a matter of fact, in some cases (as in Victorian imagery or the Japanese “wise
mother” focus) through distinctive cultural celebrations as well. Mothers’ involvement is directly
highlighted in most instances of single parenting. It also follows from the greater commitment of
fathers to work outside the home or, even when mothers also work, from the tendency that mothers
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rather than fathers remain the more flexible parent, disproportionately called on, for example, when
a child falls ill.
Fathers’ authority in the family might be diminished by more than the absence at work. Modern
economies, emphasizing employed labor more than property ownership, reduce (although without
eliminating) the importance of inheritance, diminishing one of the characteristic holds fathers had
over children in traditional rural societies. Fathers’ breadwinner status might in fact be vital to the
family economy and to children’s future prospects, but it did not bring the same level of control
that threats of disinheritance might generate in earlier settings. Where ideologies also were brought
to bear that claimed essentially natural maternal virtues—providing love, avoiding anger—that men
by definition were held to lack, the challenge of interpreting modern fatherhood simply deepens
(Griswold, 1993; Lassonde, 2016; Strange, 2015).
There is no question that modern conditions, including the decline of effective community con-
trols, increase opportunities for men to be bad fathers—unnecessarily absent or absent altogether,
sometimes seeking to avoid even minimal legal responsibilities to provide child support. Whether
actual child abuse on the part of fathers (or sometimes of course mothers as well) increases is harder
to determine historically, as we have seen, but some historians contend that the decline of commu-
nity supervision applies here as well (Pleck, 1987).
But this is not the whole story of modern fatherhood. Several historians have been working to
show how fathers maintained substantial emotional commitment and involvement in parenting, even
during the most disruptive decades of industrialization. Figuring out how fathers try to return to
greater parental engagement, for example when hours of work decline, is another important goal—
in working-class as well as middle-class settings. Many fathers during the 20th century deliberately
worked on reducing their emotional distance from their children, seeking shared recreations and
greater informality and friendliness is seeking a closer relationship—not only with sons, but with
daughters as well (Strange, 2015). How the result plays out against maternal expectations adds a
further complexity. Even aside from single parenthood, many mothers continue to find the paternal
input inadequate (Hochschild, 1989).
Here, too, regional variables may apply, although they deserve further attention. The maternal
roles continues to seem more obviously predominant in many East Asian societies, than in parts of
the West—for example, in taking the lead role in basic decisions such as selecting colleges for their
offspring (Chua, 2011).
Individualism and Alternatives

One of the most intriguing comparative tasks, in dealing with the history of modern parenting,
involves sorting out two arguably discrete strands of the Western experience, and juxtaposing them
with developments in other societies, particularly in East Asia but also in parts of the Islamic world.
Here is the issue: In Western Europe and the United States, the emergence of many of the key
trends of modern parenting, from the late 18th century onward particularly as a result of changing
economic structures, coincided with a fundamental cultural redefinition of childhood. Both Enlight-
enment values, emphasizing the goodness and educability of the child, and Romantic concepts
that stressed the distinctive nature of each child, reinforced each other in this domain. The result—
gradually, to be sure, against earlier Christian concepts including the idea of original sin—generated
increasing attention to the child as an individual, and to the responsibility of parents and other
authorities to protect and nourish this individual value. The new emphases were fully compatible
with the more basic modern trends—for example, the new emphasis on education or the decline in
the birth rate—but they were not inevitably attached.
Thus, it was possible to devise an equally modern set of parental goals—complete with promo-
tion of educational achievement and lower birth rates—without accepting the full range of Western
340
individualism. This is what has taken shape in Asia over the past century, forming a significant and
fascinating comparative contrast with Western norms. The differentiation is subtle—in Asia, too,
greater individualism emerges to a point, as in the pressure toward school achievement—but the
distinction remains quite real.
Individualistic goals in Western parenting begin to show up quite early, and amplified during the
past two centuries. The initial impulse, toward greater individualism was not immediately applied to
parenting by the ardent authors and intellectuals crafting the new standards, but the applicability to
children showed up quickly, in some of the formal discussions about educational reform that marked
the later stages of the Enlightenment. More to the point, similar impulses emerged in some striking
changes in parental practices.
Thus during the 18th century growing numbers of parents in Western Europe began to abandon
the long-standing practice of swaddling children (interestingly, the shift did not affect southern and
eastern parts of the continent, where cultural standards were more stable). Swaddling was increasingly
condemned as inimical to the proper development of the individual child, preventing rather than
encouraging physical development and childish expression. More parental (mainly maternal) atten-
tion to the young child was required as swaddling diminished, but the move toward greater freedom
seemed worth the effort. By the late 18th century, in yet another important corner of parenting,
naming practices changed. Instead of naming children for older relatives or biblical figures, parents
increasingly sought more novel options, suitable for a more differentiated and individual child. One
former practice—reuse of a name previously given to an older sibling who had died—dropped away
altogether. Parents were thinking of their children in a more individuated fashion (Shorter, 1975).
Further shifts in parenting, designed to reflect and encourage children as individuals, emerged
widely in the 19th century, sometimes linked to other developments such as more affectionate ties
or the early stages of modern consumerism. Children who died increasingly received a spot in a
cemetery, often with a modest headstone (occasionally even a more flamboyant marker, depending
on family means)—rather than a more casual burial which had been standard practice before. Atten-
tion to birthdays joined attention to the child as individual with expressions of affection through
gifts. Childrearing advice urged greater attention to the child’s individual personality and needs—in
contrast for example to more undifferentiated expectations of obedience—and to the importance of
promoting a child’s self-esteem cropped up as early as 1842 (Beecher, 1856, p. 224). By the end of the
century huge changes in sleeping arrangements worked toward the same individuating goals. Intro-
duction of the crib was part of an increasing tendency to have even young children sleep separate
from their parents, often in a different room. Older children were increasingly given rooms of their
own, as opposed to sharing with siblings—a move obviously facilitated as well by the declining birth
rate. These trends, once launched, would intensify during the 20th century and gain greater accept-
ance across social class lines. Amid some variation by personality and circumstances, Western parents
increasingly expected to think of their children as individuals and to adapt their own parenting habits
to further individual development (Reid, 2016; Yalom, 2008).
This Western pattern was not widely emulated in key parts of Asia, even as other features of mod-
ern parenting took shape and even as global interactions promoted some awareness of the Western
model. We do not, to be sure, as yet have the kind of rich histories of Asian parenting during the
19th and early 20th centuries that provide a clear counterpart to Western developments, but part of
the story, and certainly the more recent results, are clear enough.
Japanese society, during the Reform period, wrestled explicitly with the issues of Western indi-
vidualism and its relation to childrearing. Initial excitement about the Western model, based both
on study trips by reform groups and by Westerners widely employed in the new mass education
system during the 1970s, yielded quickly to insistence on the importance of group and family
loyalties over the inculcation of individualism (Stearns, 1998). The Confucian tradition in China,
as it had developed into the late imperial periods, was not immune to some acknowledgment of
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Peter N. Stearns
children as individuals. Recognition might focus on the nuisance of a willful child, but there might
be a hint of admiration as well. Nevertheless, into the 20th century, both advice and recollection
continued to highlight the importance of family loyalty over promotion of individual qualities.
Unusual reading interests thus could be actively discouraged, recalled later as a blow to creativity.
Children’s ability to share group goals and undertake collaborative tasks was widely praised, taking
unusual measures, for example, to assist an ailing family economy or simply working constructively
with adults in the fields. As the most careful student of the history of childhood in modern China
noted, the goal of helping a child “realize” himself or herself was never identified. Control of
individual needs, good manners and self-restraint, and obedience to parents and elders gained far
greater emphasis (Hsiung, 2005).
Of course, 20th-century developments might modify these approaches. But while Chinese com-
munists attacked Confucianism, they did not substitute a more individualist approach to parenting.
Nor has Japanese commitment to group harmony, over individualism, been supplanted. East Asian
cultures, to be sure, have incorporated tremendous encouragement for individual academic achieve-
ment into family goals, but without really elevating paternal encouragement to individualism more
generally.
Debates occur in other Asian cultures as well. An anthropologist describes the tension, in late
20th-century Lebanon, between global signals about cultivating the child as an individual and the
continued commitment to the primacy of family loyalty ( Joseph, 2005). Other tensions are relevant.
The custom of celebrating the birthdays of individual children does gain ground, in Westernized
circles in the United Arab Emirates or Egypt, but others are less persuaded, both because of a desire
to keep a child’s individual identity within proper founds and from a religious concern that only the
birthdays of holy figures should win social support ( Joseph, 2005).
The gap between individualism in Western parenting, and the primacy of family and group
identities in many parts of Asia, shows clearly in differentiations over the use of shame. A key part of
the modern history of parenting in the West involves a steady drumbeat of opposition to the use of
shame in dealing with children. From a social standpoint—reflecting new Enlightenment values—
shame seemed degrading, an affront to individual dignity; from the parental standpoint, according to
the kinds of advice emerging in the 19th century, shame was counterproductive, a discouragement
to individual initiative and self-esteem. The attack on shame did not eliminate the emotion, but it
did gradually persuade most parents that they should reconsider traditional practices, using milder
discipline or, even better, positive incentives to encourage good behavior (Stearns, 2017).
The same evolution simply did not occur in East Asia, even as the other features of modern
parenting began quickly to take shape. Many traditional ideas about shaming were preserved out-
right, adjusted mainly to the promotion of success in school. Mothers continued to threaten young
children with withdrawal of affection—“I do not love you right now”—not only as a means of
immediate discipline, but also as a way to instill lifelong sensitivities. Contemporary polls reveal the
continued regional disparity. Asked about the appropriateness of severe shaming in response to a
child’s bad behavior, 49% of Taiwanese parents indicated full approval, while literally no parent in
Chicago expressed anything but horror at this approach (Fung, 1999; Wong and Tsai, 2007).
The importance of prior cultural norms, and their adaptability to more modern parental settings,
constitutes a major invitation for further work that will, among other things, do more to bring Indian
and African examples into the mix. The distinctions around individualism as a parental goal already
suggest the importance of this kind of analysis in modifying simplistic, or unduly West-centric,
approaches to modern parenting. A few other targets are obvious: The continued preferences of
Indian and Chinese parents for boys, with resulting differentiations in abortion and abandonment,
contrast with gender expectations that began to even out in the West at least by the late 19th century.
But we need other, and less invidious, comparative targets as well as part of a desirable next stage in
historical analysis.
342
A U.S. Anomaly?
National comparisons may also have some utility in setting up case studies within the broader
framework of modern parenting. In the past, scholars have attempted to probe German or Russian
parenting practices to see if they provided a disproportionate basis for promoting authoritarian per-
sonalities, with attention to such features as paternal role or persistence of swaddling. More recent
changes, most obviously in German parenting since the 1950s, have largely sidelined this approach
(Farnen and Meloen, 2000).
Special features of U.S. parenting have long drawn some analysis. In the early 19th century,
European observers noted the unusually strong emotional attachment between American parents
and their children, and particularly the lack of deference that American parents tolerated or even
encouraged. They, and scholars since, have speculated that the deep roots of democratic culture
affected parental styles, giving children more voice. It is also possible that an unusual need for chil-
dren (because of an overall labor shortage) and the unusual options for children to leave home given
the open frontier might have encouraged parental solicitude.
More recently, and probably unrelatedly, another intriguing distinction has opened up, reflecting
a new kind of comparative differentiation amid an ongoing process of historical change. To be sure,
a number of observers have noted continued and significant parallels on both sides of the Atlantic,
reflecting for example the shared results of the surge of married women/mothers into the labor force
and a widely debated “end of innocence” for children exposed to increasing doses of media fare (Pol-
lock, 2010). But within this shared framework, an apparent and, arguably, troubling gap emerges as well.
One study highlighted a startling difference between many U.S. American parents and most
national clusters of European parents in the present day: American parents are more stressed than
their European counterparts and express lower levels of happiness or satisfaction than their Euro-
pean counterparts. In Europe, in most instances, parents are at least as happy as childless couples, and
often a bit happier; whereas in the United States the reverse holds true (Glass, Simon, and Anderson,
2016). The historian can suggest, further, the evidence for the declining levels of American paren-
tal satisfaction began to show up several decades ago, when comparable types of polling revealed a
drop in happiness claims (Stearns, 2003). Another study further notes the loss of self-confidence and
higher rates of depression among current American parents, compared to their baby boom parental
counterparts (Gopnik, 2016).
Two major factors, doubtless operating in combination, explain both the comparative differen-
tial and the contemporary-historical shift. First, U.S. parents are famously less supported than their
counterparts in other industrial societies. They are far less likely to receive paid leave, their medical
expenses are less well covered, the availability of affordable childcare is notoriously lower. American
parenting, in other words, is objectively harder than it is in many other modern settings, even as
Americans participate in many common trends such as the commitment of many mothers to work
outside the home.
But second—although this aspect is inherently more subjective—U.S. parents seem to have devel-
oped a set of fears and anxieties that compound their difficulties and color their attitudes. Beginning
in the 1970s and 1980s, many American parents began to become more concerned about their
children’s well-being, even safety. Key symptoms were a great exaggeration, according to opinion
polls, in the perceived risk of abduction by strangers; a new and factually groundless concern about
poisoned or otherwise altered candies in neighborhood settings during the customary Halloween
practice of trick-or-treating; and a distorted belief about rates of racial violence in schools. These
fears reflected larger, and equally inaccurate, perceptions about overall crime trends and other dan-
gers (Glassner, 1999; Stearns, 2006).
These anxieties were not merely theoretical, to be voiced to the nearest pollster. Americans began
to tighten up their parental practices. Supervision of children increased. Parents who in the 1960s
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Peter N. Stearns
freely let their children walk to school or take an urban subway now drove them directly or at least
waited with them at a suburban bus stop. Parents who were more tolerant of independent a ctivities—
like playing unsupervised in a neighborhood park—came to be called “free range” parents and were
not infrequently subject to arrest.
These palpable concerns, even about the safety of one’s own neighborhood, were then ampli-
fied by a continued commitment to maximize the preparation for adult success—symbolized most
obviously in gaining entry to an acceptable college. Parents began to do more of their children’s
homework, often even taking over the writing of the college application: It was easier, and more pre-
dictably successful, to do some of this directly than assuming that the child should take care of herself
or himself. Another interesting symptom of the combination of ambition and lack of confidence:
American parents came to lead the world in the medication of children, particularly for the malady
now labeled as ADHD—to which American children, in the eyes of their parents and teachers, were
somehow unusually prone. And perhaps on the more positive side: By the 1990s American parents
were spending more time with their children than their counterparts had in the 1950s, despite the
new levels of work and commuting commitments.
All of this added up to a challenging package, reflecting a mixture of good intentions, doubt about
children’s capacities, and lack of confidence in the social environment outside the immediate family.
Mintz (2004) described modern American parenting as alternating between a desire to protect and a
desire to prepare for a successful adult future. It might be suggested that, over the past decades, many
American parents have tried to adopt both priorities simultaneously—and have made their own task,
and their own satisfaction levels, far more problematic in the process.
The comparative challenge here deserves more attention, for while the fact that many American
parents have diverged from contemporary European patterns is clear enough, the reasons deserve
more analysis. The policy context remains relevant, as does the distinctive configuration of American
suburbs which worked well enough when the birth rate was high but became far more constraining
when children are widely scattered. Larger changes in associational activities, making Americans feel
more isolated, also factor in (Putnam, 2000). There remains, however, a contemporary historical chal-
lenge in explaining why American parents have come to feel less comfortable, more reluctant even
to rely on available childcare facilities to ease their burdens (Druckerman, 2014). Here is yet another
case, and a significant one, in which dramatic regional variations form part of the larger analysis of
modern parenting.
Conclusions
Historical analysis of parenting has always included the need to accommodate variety and to com-
pare often surprisingly different approaches to a common phenomenon. This injunction applies
from hunting and gathering societies onward. Distinctive environments, particular cultural norms,
special experiences with war or other disruptions—a host of factors shape actual parental goals and
functions.
The variety theme remains vital in modern parenting, and the examples above merely scratch
the surface. We need more work on the impact of regional cultures, but also more attention to the
experience of social class and, as the section on modern fathering suggests, to the impact of gender.
Parenting styles are closely associated with family life and with larger group identities, which can
make them resistant to many external influences and slow to change.
At the same time, as the major themes of modern parenting clearly demonstrate, change does
occur, and sometimes—as with quiet revolutions in birth rates—surprisingly rapidly. Granting the
importance of modern variety, and particularly the impact of the various devastations of the past two
centuries, a few final general points are warranted about the most global features in the history of
modern parenting.
344
For whatever the region, modern parenting has involved a number of shifts in function. Some
changes seem to have been handled fairly smoothly: many mothers, at least after some exposure to
formal education, seem to accept, or even take the lead in, the reduction of the birth rate. Many par-
ents figure out how to use consumerism to promote their bonds with their children, without being
overwhelmed in the process. Other modern developments have been more troublesome, however,
including in some cases adjusting to the new claims of the state to intervene in certain childcare
issues or negotiating decisions about caring for children amid work outside the home. And all this
is quite apart from the many special circumstances, generated by war or poverty or exploitation, that
force far more difficult parental adjustments.
Overall—and with particular focus on the overarching modern trends—two questions remain,
and both are challenging. First: Has modern parenting become easier or more difficult over time,
especially in comparison to premodern patterns? The most obvious answer would center on the
many ways the parental task has been simplified. Community support for parenting may have
declined with urbanization, but a host of experts and government resources are now available, most
notably toward the protection of children’s health. Lower birth rates and other changes reduce the
availability of older siblings in childcare—a major change; but there are obviously far fewer children
to worry about. Anxiety about assuring the survival of a smaller number of offspring may rise, but
the fact that most parents will not endure the death of a child must be taken as a huge relief.
Parents themselves, and the wider social environment, can intensify a sense of obligation, making
parenting seem more difficult. Parents in many societies are urged to pay more attention to their
offspring’s emotional development. In many cases, the new commitment to children’s happiness
imposes another set of tasks. The need to deal with outside agencies—schools, commercial solicita-
tions, government offices—can take time and also impose some concern about measuring up to
external standards. Worrying about whether children have the consumer goods they and their peers
expect, and about whether they are doing well enough in school, constitute important features of
modern parenting, and play a major role in parents’ own assessments of their adequacy. The conun-
drum is clear: Basic parental success may objectively become easier to achieve, but does not take into
account a set of new, partially self-imposed standards that, at the least, complicate the process.
Answering the basic question about parental ease or difficulty may also vary regionally, and per-
haps over time. Current U.S. parents seem to think that their task has become harder than it was in
the “good old days”, but this is not necessarily the case globally. A great deal depends on the avail-
ability of, and willingness to utilize, outside assistance, such as day care facilities. Responses are also
conditioned by the many other things modern parents must do beyond the basic parental tasks, most
notably in combining parenting with work commitments outside the home.
These considerations lead to a second kind of general assessment: about the new range of choice
in modern parenting. Increasingly—again, amid some important regional distinctions—modern par-
enting becomes an option, not a social requirement, that must be fit within other adult responsibili-
ties and aspirations. Thanks to birth control devices, more and more parents can choose when they
are ready to take on these responsibilities. A surprising minority of people continue to claim that
pregnancies were unintended (49% in the United States, but 38% in the developed nations on aver-
age), but the trend toward deliberate planning gained ground steadily. Globally the rate of unintended
pregnancies dropped by 20% between 1995 and 2016 (Finer and Zolna, 2016). Equally important
was the fact that choosing to have no children at all became an increasingly popular course. Here is
a final aspect of modern parenting that benefits from a historical vantage point.
In the United States, Europe, and Japan, many adults by the late 20th century decided that child-
lessness was preferable to parenting: work and adult consumerism, combined often with a real or
imagined sense of the difficulties of dealing with children, shaped this decidedly modern decision.
Many others were concluding that while parenting was a desirable expression, it must be confined
to a single child—another, if less radical, reflection of modern priorities. Even more widely, parents
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Peter N. Stearns
in virtually every urban society increasingly had to realize that the parental role was no longer all-
consuming, thanks to smaller numbers, the role of caregiving institutions such as schools, and grow-
ing adult life expectancy. Deciding how to combine some parenting with other adult interests—a
key element in modern feminism—and how the other interests would condition parenting in turn,
constituted a final area where traditional approaches to the role needed some modification. Here,
as in several other facets of modern parenting, many families are still working out their preferred
responses, as the challenge of adapting to modern frameworks continues.
For choice matters. In the United States, parents who feel that a new child was unintended
are likely to face other difficulties, in providing economic support and arranging other aspects of
appropriate care—for intentionality clearly reflects, but also contributes to, other differences in socio-
economic conditions, particularly given rapidly growing rates of single parenting (Cheng, Schwarz,
Douglas, and Horon, 2009). At the other extreme, people who decide not to have children at all,
for whatever set of reasons, could become a social problem in their own right. The dramatic birth
rate decline in places like Japan, reflecting concerns about the costs of parenting (including the
affordability of urban housing) and the wider range of adult interests, creates an unprecedented, and
arguably dangerous, lack of young people and a decline in demographic vigor. This development has
prompted new government measures, but so far without appreciable impact. The need to support
modern parenting, and to figure out what measures of support are most useful and why over time,
parental needs have expanded, may be a significant challenge going forward.
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10
EPIDEMIOLOGY OF PARENTING
Rebecca M. Pearson, Miguel Cordero, and Priya Rajyaguru
Introduction
Epidemiology studies the occurrence, distribution, and determinants of health-related events, states,
and processes in specified populations as well as the application of this knowledge in order to con-
trol health problems (Miquel Porta, 2016). Put another way, epidemiology is the science that first
describes and then studies causes behind the distribution of population health so that we may inter-
vene to prevent disease and promote health (Keyes and Galea, 2014).
Populations Not Individuals

Epidemiology describes what occurs in populations and not individuals. In Sick Individuals and Sick
Populations, Geoffrey Rose (2001) pointed out that the distribution of sickness across populations can
have unique causes that sometimes do not explain individual differences within these populations.
This distinction between causes of individual cases and causes of population rates is crucial to under-
standing how epidemiologists think, their methods, and interpretation of findings.
Think of a question, such as does parental discipline in early childhood affect later school achieve-
ment? To answer this question, an epidemiologist may test a hypothesis by looking at the strength
of the association between the exposure (parental control measured by an attitudinal scale) and the
outcome in the child (child achievement in a standard math test) in a large sample of individuals.
Even when an association is found within such populations, it is key to application to real life that
there will still be many individuals whose parents show high levels of control but whose children do
poorly at math.
Descriptive Epidemiology of Parenting

It is important to first describe health-related events of interest to highlight potential disease burden,
particularly at-risk populations and the potential magnitude of variation in different health states.
Descriptive epidemiology relies on having clear and consistently used definitions of the health states
of behaviors of interest. As described later, there are difficulties in defining and measuring parenting.
However, in this section we describe patterns of parenting using data from the existing literature on
aspects of parenting that have been robustly linked with several outcomes in the child and are con-
sistently measured; stimulating home environments and parental practices such as cognitive stimula-
tion, as well as harsh discipline (Byford, Kuh, and Richards, 2012).
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Rebecca M. Pearson et al.
Discipline
Discipline is an important aspect of child rearing all over the world. Traditionally discipline aims
to address undesirable behavior by preventing its reoccurrence and letting the child know that the
behavior is wrong. One way in which parents and caregivers practice discipline is using corporal
punishment. Corporal punishment is defined as hitting or spanking a child “with the intention of
causing the child to experience pain, but not injury, for correcting or controlling the child’s behav-
ior” (Straus and Donnelly, 2005).
A substantial amount of research links corporal punishment to negative social, psychological and
behavioral consequences for the child (see Thompson Gershoff, 2002; Gershoff and Grogan-Kaylor,
2016 for an overview). However, see the next sections regarding the challenges of inferring causal
links.
Corporal punishment can be considered a human rights violation and goes against the Conven-
tion on the Rights of the Child (CRC; UN General Assembly, 1989). Because of the CRC, corporal
punishment used both inside and outside the home environment has been legally banned across 53
countries worldwide with a further 56 countries expressing a commitment towards prohibition.
There are, however, several countries where the use of corporal punishment in the home environ-
ment is still legal, for example in the United States and the United Kingdom.
In view of these legal differences it is interesting to describe the prevalence of corporal punish-
ment use by parents and primary caregivers globally. The following are reported estimates based on
the available data. Using data from MICS Lansford and colleagues (Lansford, Sharma, et al., 2014;
Lansford, Deater-Deckard, Bornstein, Putnick, and Bradley, 2014) reported important variations
across 24 developing countries in parental practices over 30,470 families with 2- to 4-year-old chil-
dren. Their data shows on average 29% of parents report believing that using corporal punishment
was necessary to rear a child properly and 63% of children had been exposed to corporal punishment
in the last month. But there were large variations across countries with regards to reported corporal
punishment used over the past month, ranging from 28% in Bosnia and Herzegovina and 84% in
Jamaica (Lansford and Deater-Deckard, 2012). However, in some high-income countries corporal
punishment also remains widespread. Utilizing a nationally representative U.S. survey, 76% of men
and 66% of women interviewed in 2016 agreed that sometimes a child needs a “good, hard spanking”
(Child Trends, 2018). This was consistent with other studies exploring Northern American parent
attitudes towards corporal punishment, in which 71% approved of it as a disciplinary practice (Straus,
Douglas, and Medeiros, 2016). Moreover, in a U.S. longitudinal cohort study following children born
between 1998 and 2000, almost 50% of American parents reported spanking their 2- to 5-year-old
children in the last month (MacKenzie, Nicklas, Brooks-Gunn, and Waldfogel, 2011; 2015). In the
UK using a large population-based sample of children born between 1991 and 1992, 24% of mothers
reported smacking their child at least sometimes at the age of 1.5 years (unpublished data). Mov-
ing forwards in time by approximately 9–10 years using data from a large, nationally representative
sample of British children born between 2000 and 2002, 8.6% of mothers reported smacking their
child if they were naughty at least once per week at the age of 3 years (Millennium Cohort Study,
unpublished data in submission).
Stimulation and Play

Stimulation and play alongside other educational activities are an important aspect of child emo-
tional, psychological, social, and cognitive development (Bornstein, 2016; Bornstein et al., 2012).
Maternal education in particular has been linked to later life success including socioeconomic sta-
tus and income in the offspring (Bornstein, Putnick, Bradley, Lansford, and Deater-Deckard, 2015;
Grantham-McGregor et al., 2007). It has also been suggested that the presence of books in the family
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Epidemiology of Parenting
home increases the child’s time spent in education, on average by three more years of schooling than
for children from homes with no books. This finding holds irrespective of a caregiver’s level of edu-
cation, occupation, or class and applies to both rich and poor countries (Evans, Kelley, Sikora, and
Treiman, 2010). In developing countries, using UNICEF’s (2017) global databases the availability of
children’s books in the home varies widely, from 92% in Belarus to 11% in Honduras to less than 1%
in Mali (2005–2016). There also appears to be substantial variation according to family wealth and
financial status in almost every country with available UNICEF data (see https://data.unicef.org/
topic/early-childhood-development/home-environment/#_ftn3).
Looking elsewhere, utilizing a large Brazilian cohort of children born in 2004, Barros, Matijase-
vich, Santos, and Halpern (2010) created a stimulation score based on the number of activities each
2-year-old child was engaged with in the preceding week. These included activities such as watch-
ing TV, visiting someone else’s house, owning a story book, being told a story, and being taken to
the playground or park. They found that the highest percentage of children (approximately 60%)
had been exposed to three or four of these activities in the preceding week. In the UK using the
ALSPAC cohort of children, 83% of mothers reported taking their child to the playground at least
once per week at 2 years of age and 60% reported teaching their child at 1.5 years of age (ALSPAC
paper, pending publication). In comparison using the Millennium cohort of children born some
9–10 years after the ALSPAC children, 30.7% of mothers reported playing with their child more
than once per day and 97% reported teaching their child songs and how to count at age 3 (Millen-
nium Cohort Study, pending publication).
Why Causality Matters?

Rather than just describing patterns (as above), epidemiologists are particularly concerned with
inferring causality. Causality is crucial, because only if an identified link between a risk factor and
an outcome (i.e., parenting and behavioral problems) is causal, will changing the risk factor within
a population prevent the outcome (i.e., disease). Focusing on noncausal risk factors will be inef-
ficient at best. However, the causal link does not have to be deterministic (i.e., only one cause for
one outcome and if you have that cause the outcome is inevitable). Such a proposition is unrealistic,
especially in the context of complex behavioral outcomes which will have multiple causes. Thus,
epidemiological evidence seeks to identify factors that contribute to increased risk in a causal way and
separate those from factors that are linked to the outcome in a completely noncausal way. To high-
light why causality is important, consider the counterfactual model.
Counterfactual Model
Put simply, what would have happened if things had been different? What would have happened if
everything in the situation had been the same, but the only difference was that the potential causal
factor had not occurred? This is the counterfactual model.
Consider an adult, orphaned as a child and recipient of institutional care, who later develops
depression. Let us assume that being an orphan causes depression through the negative experiences
of not having a stable primary caregiver. In a counterfactual situation, if the same child had never
been orphaned, the risk of depression would be reduced: Institutional care is a causal risk factor. The
child also slept in a shared bedroom. In a counterfactual situation, if this same child was given their
own room in the orphanage, but nothing else changed, the risk of depression would not be reduced.
Sharing a bedroom is not the causal factor. In a population sample with adequate numbers of orphans
and nonorphans, however, if we just looked at sharing bedrooms and depression risk we would likely
observe a link. The link is inevitable because it is the norm in institutional dormitories to share bed-
rooms but not in most family homes. Shared bedrooms are thus a proxy for institutional care within
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populations. In other words, those who share bedrooms are more likely to be orphaned, and thus a
link between shared bedroom exposure and depression is created within a population. If we assume
this observed association with shared bedrooms was causal, to prevent depression, we would remove
shared bedrooms. Removing would not reduce the risk of depression because sharing a bedroom is
not the true causal risk factor and only associated with depression due to sharing a bedroom being a
marker of the true causal risk factor (institutional care). The distinction between true causal and non-
causal factors is very difficult in observational work especially with social and behavioral outcomes.
Causality is essential to making a difference. If parenting is not causally associated with outcomes,
then changing it will make no difference to child outcomes and we should look elsewhere for modi-
fiable factors which may help promote child development.
What Can Epidemiology Contribute to Causal Parenting Research?

In the field of parenting research, there is an extraordinary large corpus of theoretical and empirically
based knowledge. This Handbook is one such example of how diverse perspectives converge to study
this corner of mammalian behavior. It remains difficult, however, to make causal links between small
variations in what parents do and the resulting adult traits of their children. There are several reasons
E D
Exposure/treatment Disease/outcome
Potenal outcome for 1 individual in real and counterfactual scenario
D(E=1) D(E=0)
Figure 10.1 Counterfactual model of causality.
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Potenal outcome for populaon with the same reality and counterfactuals
D(E=1) D(E=0)
Figure 10.1 Continued
for this. First, parents and children share the same environments and common genetic background.
Second, parenting may be involved in the causal pathway but not necessarily be the primary cause of
the outcome. Third, parenting is likely to also be influenced by child outcomes and so determining
whether child outcomes influence the parent rather than the other way around is difficult. Perhaps
because of such complexity, parenting has been somewhat neglected within the epidemiology field.
For example, a keyword search for “parenting” in the International Journal of Epidemiology yields just
three papers, all published in the last decade.
Despite such sparse evidence, parenting has been repeatedly implicated as a risk factor for obesity,
a variety of mental health disorders, school achievement, and social and adaptive outcomes.
We propose that a systematic approach using an epidemiological framework can help to strengthen
causal inference and arrive at new insights that can help to improve overall population health. We
define and apply terminology and methods that are regularly used by epidemiologists to conceptual-
ize and discuss challenges in current research on parenting. Using examples from the literature we
illustrate how epidemiology can contribute to making new progress in parenting research.
Parenting Variables Acting in Different Roles on the Causal Pathway

If you are interested in modifiable exposures, such as parental behavioral factors, diet, or other exter-
nal environmental exposures such as chemicals, the obvious thing to do is to measure the exposure
and then look at how variation in the measure relates to health outcomes. However, the theoretical
or hypothesized connection between the two variables can take many forms, and we define termi-
nology used to describe the different connections below.
1. Parenting as a risk factor for the outcome, where those “exposed” to a certain parental variable
are more likely to have the outcome; for example, stimulating parenting (exposure) may have
better executive functions (outcome) in the child.
2. Parenting can be the outcome and caused by the other variable; for example, depression (expo-
sure) in mothers may lead to less stimulating parenting (outcome).
3. Parenting can be a mediating variable, in this case it is part of the causal chain from the exposure
to the mediating variable to the outcome. In this case, parenting is both an outcome and an
exposure. For example, depression (exposure) may lead to less stimulating parenting (mediating
variable) which in turn leads to lower executive functioning (outcome).
4. Parenting can be a confounding variable. Confounding happens when all or part of the apparent
association between the exposure and the outcome is accounted for by other variables that are
also linked to the outcome and exposure, but are not consequences of the exposure (i.e., part of
the causal pathway; Miquel Porta, 2016).
5. Parenting can be a moderating variable. Moderating variables alter the strength or direction
of the association between an exposure and an outcome. For example, greater levels of father
involvement may reduce the association between maternal depression and poor child outcomes,
because the father compensates for the mothers reduced emotional availability.
Life Course Epidemiology Models

In addition to the definitions above, we need to consider how parenting coincides with other risk
factors over time. We cannot view parenting in isolation, and its influences occur over an extended
period of development. This situation adds yet further complexity to causal inference.
Life course epidemiology models have attempted to define and term how different timings of
exposures can have long-term effects on later health by two principal paths: (1) exposure in a sensi-
tive period and (2) accumulation of risk with repeated chronic exposures.
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EXPOSURE OUTCOME
Child
1. Parenng execuve
funcon
2. Maternal
Depression Parenng
Child
3. Maternal
execuve
Depression
funcon
Parenng
Maternal Child
Depression execuve
funcon
4.
Parenng
Parents who show low

levels of structured
parenng
Child
5. Maternal execuve
Depression funcon
Parents who show high

levels of structured
parenng
Child
Maternal execuve
Depression funcon
Figure 10.2 Potential outcomes for individuals and populations in a real and counterfactual scenario.
Sensitive Periods
propose that an exposure will have greater impact if it occurs during a specific time window rather
than the same exposure during a different time window. It is often proposed that parenting may have a
greater impact earlier in life due to rapid brain development (Gupta et al., 2017; Lomanowska, Boivin,
Hertzman, and Fleming, 2017). There is some evidence, for example, that orphans living in institutional
care who are then fostered into a sensitive family environment at a younger age leads to better outcomes
than those fostered later (Ghera et al., 2009; Vanderwert, Marshall, Nelson, Zeanah, and Fox, 2010).
However, in more subtle examples within healthy populations, there is not always evidence for timing
effects, rather repeated and chronic exposure is most often associated (Evans et al., 2012). Early exposure
is also hard to separate from greater duration of exposure; for example, if you are exposed to adverse
parenting from an earlier age it is likely to continue throughout childhood. This would represent a
longer total number of years exposed to adverse parenting than if you are only exposed from a later age.
Cumulative risk assumes that the long-term outcome occurs through the composition of environ-
mental, socioeconomic, and behavioral exposures during life course. Accumulation of risk can occur
as follows:
1. With independent and uncorrelated exposures

2. With correlated exposures:
a. “Risk clustering”
b. “Chains of risk” with additive or trigger effects.
First, two risk factors may contribute to the child outcome, but they are unrelated to each other.
For example, being a girl is a risk factor for depression and so is having a mother with depression.
Thus, if you are a girl with a depressed mother you have a greater absolute risk of depression than a
boy with a depressed mother. However, being a girl is not correlated with having a depressed mother,
so the two risk factors act independently.
Second, variables may be linked by common causes. For example, low maternal education is asso-
ciated with parenting that lacks cognitive stimulation but also with poor diet. Thus, poor diet and
lack of stimulation are correlated, and both contribute to risk of lower child cognition. The cluster
of risk factors relates to those which relate to maternal education.
Third, exposures may chain in a sequence, with each exposure increasing the likelihood of sub-
sequent exposure, as well as subsequently influencing the risk of an additive effect on the observed
outcome (Braveman and Barclay, 2009; Maughan and Collishaw, 2015). For example, observing vio-
lence in the home is associated with violence in the child at school which contributes to the child’s
development of depression.
Threats to Detect Real Causal Effects in Parenting Research

Separating causal and noncausal associations is very hard from studying populations. There are sev-
eral threats to causal inference that have been well thought out in epidemiology. In this section,
we describe each one, and we use examples from the parenting literature to understand sources of
threats to causality and potential approaches minimize the role of such threats which can minimize
the problems of invalid casual claims.
Precision Versus Accuracy

Estimation of the association between X and Y can be affected by error and bias introduced by the
study design that then leads to inaccurate estimations. Although often confused, the concept of bias
356
differs from precision. Precision concerns the fact that we only have an estimate of any association
in any whole population of interest (for example, all parents and children in the world), based on
observations taken from one smaller sample. There will always be some variation between statistics
taken from different samples, even if each sample is representative of the true population. The more
precise the statistics, the smaller the variability between the sample statistics and the more we can
narrow down the likely values of the parameter within the total population. The precision of a single
sample statistic can be considered by calculation of a confidence interval and is directly related to the
sample size (larger samples are always more precise).
Accuracy, however, is concerned with whether the observed estimates from the study population
systematically differ from the true associations in the true population. Such inaccuracy in the study
would be through biases (see below) introduced by the study.
Selection Bias
We described above in the section on precision how the selection of a single sample from a target
population will be prone to sampling variation (random sampling error). Sampling variation alone
will not affect the accuracy of our estimate of prevalence or presence of associations. It just means
that we can only estimate a range of values (i.e., the “true association” will likely lie somewhere
between the confidence intervals).
However, if there is systematic error in the selection of participants we may end up with an inac-
curate estimate. For example, parents who volunteer to advertisements for studies often have a personal
interest in parenting. The prevalence of disease or exposures in this volunteer group may be very
different from that in the larger underlying population, hence this difference may result in either an
overestimate or underestimate of the true prevalence (Howe, Tilling, Galobardes, and Lawlor, 2013).
Nondifferential Selection
Provided any errors in the selection of participants occur equally between all groups being compared
(e.g., exposed and unexposed), the estimate of the association between exposure and outcome will
be unbiased. For example, if younger mothers are less likely to be recruited into a parenting trial, but
this occurs equally across both arms of the trial, this is nondifferential selection and would not bias
the association between groups.
Differential Selection
If, however, any systematic bias in the selection of participants occurs differentially across groups,
then selection bias may be present and result in either an underestimate or overestimate of the
association between exposure and outcome. For example, if younger mothers are less likely to be
recruited to one arm of a nonrandomized comparison of two treatments for parenting (e.g., because
they are more afraid of stigma from attending parenting classes), this differential selection could bias
the association between parenting interventions and outcome.
Evidence of Selection Bias in Parenting Research

Selection bias can occur from the initial recruitment of participants or when participants drop out
of studies or choose not to complete all measures. Selection bias is especially relevant for parenting
research which relies heavily on parents agreeing to participate with their child. Selection bias may
lead to systematic differences in parents who do and do not take part.
The Avon Longitudinal Study of Parents and Children is such an example (Boyd et al., 2013;
Fraser et al., 2013; Golding, Pembrey, Jones, and Team, 2001). Among those parents who attended
357
visits for general assessments, there are differences among those who completed parent-child interac-
tions. In the final study sample of this analysis, women of higher education and social class who were
more likely to be married and to have breastfed their infant were included (Pearson et al., 2011).
Measurement Bias
There will also be errors in the measurement of the exposure and/or outcome in any epidemiologi-
cal study. For example, an individual’s blood pressure will vary from day to day or even throughout
the day, hence different measurements taken on the same individual will vary at random around their
average blood pressure. Alternatively, the device measuring blood pressure may be imprecise equating
to random variation in readings. Indeed, there will always be some degree of random error (which is
not bias) in the measurement of exposures and outcomes. If, however, the device is inaccurate such
that it always underestimates or overestimates blood pressure, or the health care professional using
the device rounds measurements up or down, then there will be some degree of systematic error.
Measurement Bias in Parenting Research

Parenting is difficult to measure as though like blood pressure in its variation, it is also an abstract
concept not always directly observable. Hence one relies on the manifestation of such behavior or
the individual’s subjective report of their experience; lending itself to both nonsystematic and sys-
tematic errors or biases as described below.
Nonsystematic measurement error causes one main problem that associations between measures of
parenting and later outcomes will be attenuated (likely resulting in previous studies reporting under-
estimates of true associations).
There are methods which can help account for such measurement error. Approaches which just
add up all item scores on a survey, for example, do not allow for the fact that the observed scores
are only a manifestation of the underlying hypothetical construct of interest and thus contain meas-
urement error. In contrast, factor analytic techniques, such as those employed in structural equa-
tion modeling (SEM), assume that observed information reflects an unobservable construct, and
measurement error is one of the sources of variance (Kline, 2010).
If the meaningful (i.e., that which relates to the construct) and error sources of variance are not
accounted for (as in scores which just add up all items from a survey) any “true” differences between
constructs may be masked. For example, if all observed parenting item scores are summed, scores
represent (1) true differences in the individual’s parenting construct and (2) measurement error.
Construct relevant variance may be considered as that which is shared between items theoretically
assigned to the same parenting constructs because if they all measure the same construct what they
all share should represent that construct. The nonshared variance is likely to be the error of each
individual item. By deriving construct-specific latent traits based on shared variance across items
only, SEM factors allow different items to contribute differentially to the construct of interest, and
measurement error is accounted for in the model. Minimizing the role of nonsystematic measure-
ment error in associations should reduce the issue of underestimation of associations. Indeed, there
is empirical evidence that if the association between the same two variables is first modeled by using
both variables as sum scores and then using both variables as latent traits, the association between the
two latent traits was larger than with the sum scores (Pearson et al., 2015).
Reporting Bias
For many dimensions of parenting that involve rare events, such as harsh discipline and internal
feelings of love or irritation, parental report is the most appropriate measure. Observations do not
358
capture such events. Importantly, measurement error only leads to bias if it is systematic (it is sys-
tematically related to both exposure and outcome). This is, however, highly likely if the outcome is
also maternally reported (due to shared method variance). For example, more educated mothers may
overestimate their frequency of book reading with their child (as they know they should) and their
child’s ability in reading (as they value such skills). The overestimation of book reading in educa-
tion mothers would create an association between reported book reading and child ability. Studies
with multiple informants reporting child emotional symptoms have demonstrated that associations
between maternal and child depression when relying on maternal report of both are substantially
overestimated (Ringoot et al., 2015) as compared to associations between maternal depression and
child self-report or teacher or father reports of child depression. The same is likely true for parental
reports of parenting and child outcomes. One potential approach, if systematic measurement error is
thought be relevant, is to adjust for variables which represent the sources of such biases. Such vari-
ables may include social desirability, education, and neuroticism which are likely to explain some of
the reporting biases.
Another source of measurement error lies in trying to capture parent report of parenting or
parental style across multiple children. Parenting is likely to differ for each child, and thus studies
should clearly state which child they are referring to and conduct different measures for different
children, particularly relevant for within-family and twin designs (see later). Parenting differs by
child gender and birth order (Pearson et al., 2011)—which are two clear examples why parenting
by the same mother is likely to differ across siblings. Different times and contexts and development
stage will also require and evoke different parental and child behaviors. For example, the amount
of autonomy versus direction a parent provides changes due to context (safe versus harsh environ-
ments), age, and child ability.
Recall Bias
An alternative to parent report is to measure parenting experienced by the child as reported by the
child. However, unless measuring current experiences of parenting this approach will rely on retro-
spective recall which can be subject to recall bias. In Children of More Caring, Less Controlling Parents
Live Happier Lives,—the authors reported: “We found that people whose parents showed warmth
and responsiveness had higher life satisfaction and better mental wellbeing throughout early, middle
and late adulthood.” This study examined parent-child relationship quality and positive mental well-
being using Medical Research Council National Survey of Health and Development data (Stafford,
Kuh, Gale, Mishra, and Richards, 2016). Well-being was measured at ages 13–15 (teacher-rated hap-
piness), 36 years (life satisfaction), 43 years (satisfaction with home and family life), and 60–64 years
(Diener Satisfaction with Life scale and Warwick Edinburgh Mental Well-being scale). The Parental
Bonding Instrument captured perceived care and control from the father and mother to age 16,
recalled by study members at age 43. The authors report a greater well-being observed in the off-
spring of parent combining higher warmth parental care and lower psychological control. However,
it equally may be the case that individuals with greater life satisfaction look back at their experiences
more positively.
A particularly salient example of potential recall bias is in the context of Adverse Childhood
Experiences. The Adverse Childhood Experiences (ACE) study was a large epidemiological study
examining effects on health of adverse life experiences in childhood and health outcomes in adult
population (Felitti et al., 1998). Since first reports, dozens of papers have shown large increases of
risk in almost every health condition from cardiovascular disease to smoking, suicide and adolescent
pregnancy, nearly every mental health disorder, school achievement, unemployment, and all causes of
death in adults (Afifi et al., 2008; Anda et al., 2008; Hemmingsson, Johansson, and Reynisdottir, 2014;
Liu, Croft, and Chapman, 2013; Pietrek, Elbert, Weierstall, Muller, and Rockstroh, 2013; Schilling,
359
Aseltine, and Gore, 2007). Comparing individuals who do not report having these experiences, those
reporting multiple ACEs had higher risk in almost every outcome. Another paper studying whether
associations between ACEs and health outcomes are the same for prospective and retrospective ACEs
measures showed moderate correlations between prospective and retrospective measures. Compared
with prospective ACEs, retrospective ACEs showed stronger associations with middle life health
outcomes that were subjectively assessed (e.g., self-reported physical health and psychopathology)
and weaker associations with outcomes that were objectively assessed (e.g., waist circumference,
cholesterol HLD, working memory performance).
Performance Bias
It is often assumed that observation of parenting provides a less biased measure than relying on
report. Although research can take steps to ensure the observer and ratings are unbiased (training
and high inter-rater reliability, being blind to any information on the parent or study hypoth-
esis), researchers can only observe what the parent shows them. The presence of knowing you
are being recorded/observed and the physical presence of an observer will alter behaviors. If this
tendency to behave differently is related to the outcome/exposure of interest, it will bias the
results.
Observation may promote socially desirable or appropriate behaviors and suppress socially unde-
sirable or inappropriate behaviors (e.g., adults may display higher rates of positive interactions with
children; Baum, Forehand, and Zegiob, 1979; Zegiob, Arnold, and Forehand, 1975). This result may
be differential according to different maternal characteristics; that is, some mothers may behave more
positively, while others may become self-conscious and thus behave less positively (Weber and Cook,
1972). Observed parenting may thus be associated with positive child outcomes, not because parent-
ing influences child outcomes, but because parents who can perform well on observed tasks also have
children who can perform well.
There are emerging methods to help overcome performance biases in parenting research. Lee
et al. (2017) compared footage recording the same play interactions from a traditional researcher
present setting with two different cameras: third-person point of view (3rd PC) and using cameras
worn on headbands (first-person cameras [1st PCs]) to record first-person points of view of mother
and infant simultaneously. Importantly in a second stage of the study, the dyads are left alone with the
1st PCs for many days to record natural mother-infant behavior at home.
Data from 1st PCs during sessions recorded alone at home captured more “negative” maternal
behaviors per minute than observations using 1st PCs while a researcher was present (mean differ-
ence = 0.90 (95% CI 0.5 to 1.2, p < 0.001) representing 1.5 SDs). Based on the assumption that
mothers are more likely to display more behaviors that are socially considered “negative” parent-
ing when demand characteristics are reduced, the finding of increased negative parenting provides
preliminary evidence for reduced demand characteristics. A further study demonstrated that a more
extreme “negative” maternal behavior of corporal punishment is recorded more frequently by using
passive audio recording in the home than reported frequency from mothers (Holden, Williamson,
and Holland, 2014). Demand characteristics may be especially high in educated mothers, due to
knowledge of research and “best parenting.” This potential bias poses a challenge to measurement of
parents’ behavior, in which consistent associations are made between maternal education levels and
observed parenting
There are many ways in which 1st PCs could lead to reduced demand characteristics. It may
because no researcher is physically present or could result from longer duration of recording. Briefer
observations can be unstable, and observations lasting longer than an hour are likely to include sam-
ples of behaviors from highly varied activities or contexts, thus being more varied themselves (Miller,
Shim, and Holden, 1998).
360
Reverse Causality
A link between two variables does not tell us the direction of any association. Bi-directionality or
reverse causality should always be considered in parenting research. In previous work, parent behav-
iors were considered antecedents, and child behavior as an outcome, and when both were correlated
often interpreted as an effect of one person’s behavior (the parent) on another individual (the child;
Maccoby, 1992). However, it can easily be the other way around. Cross-sectional studies which
measure both variables on the same occasion (for example parenting and child behavior) make this
causal distinction impossible.
Longitudinal studies can improve temporal understanding; for example, parenting may be meas-
ured at age 3 and then looking at the extent to which it is associated with behavioral outcomes in
the child several years later. However, the extent to which this approach eliminates reverse causality
is not always clear cut. For example, just because parenting was measured first in the study doesn’t
mean that in the real-world parenting influences behavior first. It may be that behavioral problems at
age 3 already influence parenting at age 3 and then the behavior problems persist several years later.
The only way longitudinal designs can overcome reverse causality is if either at the first time
point the proposed outcome does not already exist or influence the exposure. An example of this is
a study reporting an association between maternal cognitive style during pregnancy and offspring
cognitive style at age 18 (Pearson, Fernyhough, et al., 2013). The authors proposed that the most
likely mechanism of association is that the child mimics their mothers’ way of thinking. An alterna-
tive is that the mother mimics the child’s. However, in this design, as the child was not yet born, this
is implausible. If the measure was taken later point (even if mothers were measured before the child)
it would not be possible to rule out that the child’s cognitive style had not at some point influenced
that of the mothers.
Alternatively, longitudinal designs may measure the proposed outcome at baseline and then adjust
for this in the analysis, thereby accounting for any pre-existing association. For example, a study
reported a strong association between affiliating with goth subculture at age 15 and later depression
at age 18. It seems highly likely that pre-existing depression influenced teens to affiliate with such
subculture rather than being a goth caused depression (Bowes et al., 2015). Thus, the study took
advantage of a longitudinal design and looked at the association between goth affiliation at age 15
and later depression, adjusting for baseline depression. Perhaps surprisingly there was little evidence
for attenuation of the association, suggesting that reverse causality did not account for the observed
association.
Bi-directional Associations
In reality however, many associations, especially between parent and child, are likely to operate in
both directions. Bi-directional effects have been studied in the context of intervention or longi-
tudinal studies using repeated measures (Barnett, Gustafsson, Deng, Mills-Koonce, and Cox, 2012;
Besemer, Loeber, Hinshaw, and Pardini, 2016; Del Vecchio and Rhoades, 2010; Hummel, Kiel,
and Zvirblyte, 2016; Moilanen, Rasmussen, and Padilla-Walker, 2015; Te Brinke, Dekovic, Stoltz,
and Cillessen, 2017). Such repeated measures allow the temporal links from parenting at time 1 to
behavior at time 2 and vice versa while accounting for the time 1 correlation between parenting and
behavior. Any association between parenting at time 1 and behavior at time 2, over and above the
variance explained by the baseline association, can be inferred to be independent of reverse causality.
Findings of such bi-directional links are mixed but there is emerging evidence for robust but rela-
tively small bi-directional effects. This is likely to be partially due to the statistical power needed to
detect associations over and above variance explained by stability in parenting and behavior (Barnett,
Gustafsson, Deng, Mills-Koonce, and Cox, 2012; Besemer, Loeber, Hinshaw, and Pardini, 2016; Del
361
Vecchio and Rhoades, 2010; Hummel, Kiel, and Zvirblyte, 2016; Moilanen, Rasmussen, and Padilla-
Walker, 2015; Te Brinke, Dekovic, Stoltz, and Cillessen, 2017).
Confounding
Latin: “Confundere” is to mix together. Unlike bias which is introduced by study design, confound-
ing is a consequence of the natural co-occurrence of variables in the real world. There are many
potential risk factors for parenting that could also be independently associated with child outcomes,
thus creating spurious noncausal associations between parenting exposure and child outcomes. For
example, a previous study reported an association between more responsive observed parenting at
age 12 months and later child IQ at age 4 (Pearson et al., 2011). An obvious confounding variable
here is maternal education. Maternal education influences maternal parenting due to knowledge and
problem-solving skills. Maternal education also directly influences child IQ both through common
genetics but also through the mothers’ skills/knowledge transfer. Maternal education would not be
considered a mediating variable as it is unlikely to be a consequence of maternal parenting, given
that the mother’s education largely occurs in pre-parenthood life. In this study adjusting for mater-
nal education diminishes the association. The attenuation of the association provides evidence for
confounding by education.
However, even if associations remain after controlling for confounding variables, residual con-
founding remains a likely possibility (Fewell, Davey Smith, and Sterne, 2007). It is unlikely that the
study considered and measured every possible confounding variable, thus resulting in unmeasured
confounding. In addition, given that just like exposure and outcome variables are measured with
error so are confounding variables and therefore the variables used to adjust for do not fully capture
the confounding construct (Fewell et al., 2007).
One example of likely unmeasured cofounding in the example of parenting to child outcomes is
genetic confounding where shared genetic risk for poor parenting and negative child outcomes cre-
ates associations between parenting variables and child outcomes. Given that genetic basis for com-
plex phenotypes is still unknown and thus impossible to capture in existing data, such confounding
remains unmeasured in most studies.
Mediation Versus Confounding

The distinction between mediation and confounding is very difficult. The key difference is the
direction of the arrow of association between the third variable (possible confounding or mediat-
ing variable) and the exposure variable. If the variable is correlated with or causes the exposure (see
Figure 10.2 number 4), it is a confounding variable. However, if the variable is a consequence of the
exposure it is on the causal pathway and thus a mediating variable (see Figure 10.2 number 3). This
distinction is subtle but important. Confounding introduces observed associations that are noncausal.
Mediating variables highlight part of the causal associations. Failing to correctly account for con-
founding may lead to falsely concluding an association is causal when it is not. However, including
a mediating variable as a confounding variable in analysis may lead to falsely concluding an associa-
tion is not causal when it is. The distinction between mediation and confounding must be driven by
theory and clinical expertise or design (i.e., in longitudinal studies we may be sure that the variable
preceded the exposure). See Table 10.1 for considerations of interpretation of analysis adjusting for
variables which are confounding or mediating associations.
As well as the possibility of including mediating variables, inclusion of potentially covariates into
models can also introduce further forms of bias. Less known is a type of bias created through analy-
sis choice and the inclusion of adjustment variables between an exposure and an outcome. These
phenomena have been described as Berkson’s bias, Simpson’s paradox, backdoor (basing path) and
362
Table 10.1 Interpretation of parenting and child outcome associations with variables conceptualized as confounding and mediation
Scenario Analytic Model Diagram Results Interpretation assuming no other Translation

important biases and adequate
statistical power
Parenting as exposure Univariate E 

D Evidence of association No causal association Changing parenting would not
for child outcome Multivariate with No longer evidence of help to change the outcome.
including variables confounders association Consider other variables.
conceptualized as
confounders
Univariate E 
D Evidence of association Association is unlikely to be Changing parenting still may
Multivariate with Association remains due to measured source of help modify the outcome,
confounders after adjustment confounding. Causality still is subject the further evidence
a possible explanation to the
association.
Parenting as exposure Univariate E 
D Evidence of association Exposure may explain changes Changing parenting Would
for child outcome Multivariate include No longer evidence of in the outcome through a help only if it changes
including variables variables possible association mediator variable. the mediator, e.g., Home
conceptualized as involved in the causal environments
mediating variables pathway
Univariate E 
D Evidence of association Association is not affected for Parenting is possibly an
Multivariate include Association remains those potential mediator. The independent cause of the
variables possible after adjustment association is independent of outcome. The potential
involved in the causal the third variable. mediator is also possibly an
pathway independent cause.
collider bias (M. Porta, Vineis, and Bolumar, 2015). The collider is produced every time that a vari-
able is the common effect of an outcome and an exposure, is conditioned upon in the regression
(controlling, through stratification, restriction, or adjustment). In this adjusted analysis inclusion of
this variable can then induce an inexistent association (Munafo, Tilling, Taylor, Evans, and Davey
Smith, 2017; M. Porta et al., 2015).
Methods to Help Account for Residual Confounding
Negative Controls
A method that sometimes helps to deal with residual confounding in prenatal exposures is to test
the association with a negative control, for example comparing the exposure experienced by the
mother and the outcome in the child, with the association of same exposure in the father during
pregnancy and later behavioral outcome in the child. If the effects of interest are due to an intrau-
terine exposure, then maternal exposure during pregnancy should have a clearly greater influence
than paternal exposure (Smith, 2008). Using this approach in a cross-cohort study, Van Batenburg-
Eddes et al. (2013) found little evidence of a difference between the strength of associations of
maternal and paternal symptoms of depression during pregnancy with offspring- inattention in the
child. Therefore, the observed associations are likely to be driven by mechanisms shared by parents
(home environment, genetics or parenting) rather than maternal specific mechanisms such as in utero
effects. In contrast, there is evidence for associations between maternal depression during pregnancy
and offspring depression at age 18 but no such association with paternal depression during pregnancy
(Pearson, Evans, et al. 2013).
Family Designs
Twins, full siblings, and half-siblings are increasingly used as comparison groups in matched cohort
and matched case-control studies. The “within-pair” estimates of associations acquired through these
comparisons are free from confounding from all factors that are shared by the siblings (Frisell, Oberg,
Kuja-Halkola, and Sjolander, 2012). Thus, these methods are helping to solve problems with unmeas-
ured confounding. For example, prenatal exposure to maternal smoking has been associated with
increased risk of attention-deficit/hyperactivity disorder (ADHD), bipolar disorder, schizophrenia,
and other behavioral disorders in offspring. However, studies suggest this association might be due to
unmeasured familial confounding (Kuja-Halkola, D’Onofrio, Larsson, and Lichtenstein, 2014; Meier
et al., 2017; Quinn et al., 2017; Skoglund, Chen, D’Onofrio, Lichtenstein, and Larsson, 2014). In a
population-based cohort of 1.7 million Swedish offspring (Quinn et al., 2017), maternal smoking
during pregnancy showed increase risk of severe mental illness rates compared those unexposed off-
spring (moderate smoking during pregnancy: risk ratio (RR) 1.25; 95% CI, 1.19–1.30; high smoking
during pregnancy: RR, 1.51; 95% CI, 1.44–1.59). In sibling comparisons with within-family covari-
ates, associations were substantially weaker (moderate smoking during pregnancy: RR, 1.09; 95% CI,
0.94–1.26; high smoking during pregnancy: RR, 1.14; 95% CI, 0.96–1.35).
Sibling comparisons can be particularly powerful in parenting research but require careful meas-
urement of indices of parenting that differ across each parent-child relationship. There is evidence
that conflictual and negative parenting behavior directed specifically at an adolescent is strongly asso-
ciated with psychopathology for that adolescent but may be protective for their sibling (Reiss et al.,
1995). Such studies also demonstrate that parenting is key component of the nonshared environment
(i.e., that which is experienced uniquely by each individual). In contrast, other key confounding
variables of the association between parenting and child outcomes, such as maternal education,
income, age, marital status, and mental health, will be shared between siblings. Thus, these studies
364
help separate out the associations between the parenting experienced specifically by each child and
other sources of confounding.
Adoptive Studies
Adoptive studies are more common today in innovative ways, adoption at birth allows the influence
of the adoptive mother to be separated from adoptive mother genetic impact. However, the role of
the birth mother maybe through either in utero effects or direct genetic inheritance. Passive and
evocative gene/environment correlations have been well documented by twin studies. Genetically
influenced behavior can evocate or attract certain environments and canalize certain developmen-
tal pathways. Thus, the parenting of the adoptive parent may still be influenced by the genetic risk
passed from birth parents to child.
In vitro fertilization (IVF) studies offer a new way to deal with this problem. Children conceived
via assisted reproductive technologies can be genetically related to both parents (homologous IVF),
the mother only (sperm donation), the father only (egg donation), or to neither parent (embryo
donation; Golombok et al., 2002; Golombok, MacCallum, & Goodman, 2003). Using this natural
experiment, Harold and colleagues (2013) examined the association between maternal hostility and
child ADHD symptoms using mixed cross-sectional and longitudinal design in one sample of adop-
tion at conception and a second one of adoption at birth. They observed evidence of association
between maternal hostility in rearing mothers and child ADHD, and between mothers’ ADHD
symptoms and child ADHD in both samples. As all children were genetically unrelated to their rear-
ing mother, it is less plausible to attribute this association to a genetic correlation.
Randomized Controlled Trials

The field of observational epidemiology has vastly improved with regards to the causal inference
capacity. However, confounding and reverse causality remain difficult to fully overcome. Thus, evi-
dence from randomized controlled trials (RCTs), which eliminate all known/unknown and unmeas-
ured confounding, is essential because the exposure variable (i.e., the intervention) is randomly
allocated and therefore cannot be connected to the confounding variables (criteria of confounding
is an exposure to confounding link).
However, in the context of parenting research the exposure can only really be a parenting inter-
vention which is not the same as parenting. It is not possible to directly manipulate parenting,
thus RCTs only provide definitive casual evidence regarding whether parenting interventions work,
rather than whether parenting is causally related to child outcomes for three key reasons:
1. Not all parenting interventions change parenting behavior. If they do not change parenting,
then the fact that a parenting intervention does not child outcomes tells us nothing regarding
the relation between parenting and child outcomes.
2. Parenting interventions that change parenting may be subject to measurement bias. This is par-
ticularly the case because it is usually impossible to blind participants to the intervention they
receive when that intervention is a parenting intervention. Those parents in the intervention
arm are more likely to report and perform well in assessments of parenting, but at home nothing
may change.
3. Parenting interventions are complex and involve multiple components; thus, if a parenting inter-
vention changes a child outcome, it may be through a nonparenting mechanism or only one
specific components of parenting. Newer analytic techniques such as components network
meta-analysis break down complex interventions into component parts and compare which
component is driving intervention effectiveness (Caldwell and Welton, 2016).
365
Table 10.2 Sources of bias in different study designs
Epi approach/design Environmental confounding Genetic confounding Measurement Reverse causality Bias
Negative Control * *
Latent traits *
RCT * * * *
These caveats aside, RCT evidence from parenting interventions can provide further evidence to
support the role of parenting as a causal risk factor. This is because if parenting is causal then changing
parenting successfully should change child outcomes. Reviews suggest that parenting interventions
are associated with more positive child outcomes, but individual effects are mixed, and intervention
success is likely to be dependent on population, intervention and outcome (Lozano-Rodriguez and
Valero-Aguayo, 2017; Medlow, Klineberg, Jarrett, and Steinbeck, 2016; Mihelic, Morawska, and Filus,
2017; Vlahovicova, Melendez-Torres, Leijten, Knerr, and Gardner, 2017).
Chance
Across all study designs we can never rule out the role of chance, in other words that any association
is simply a product of sampling variation. P values give an estimate of this possibility, and increased
sample sizes increase the statistical power to detect associations unlikely to have occurred by chance.
The amount of power required depends on the effect size of interest. Pooling results across studies can
minimize the role of chance. However, just as selection bias can systematically bias the individuals who
take part in studies, systematic biases can operate that bias which studies are found in available parenting
literature. The most prominent of these is publication bias (Button et al., 2013) where studies that report
a positive association by chance are much more likely to be published than those that report a null
association. Reviews which include gray literature and unpublished studies, as well as greater journal
incentives to publish well-designed but null findings, will begin to minimize the role of publication bias.
Conclusions
Epidemiological methods systematically consider the role of threats to causality, which pose real
challenges for parenting research. As highlighted above and in Table 10.2 different study designs have
different strengths and limitations regarding these issues and in general each limitation of method is
orthogonal (i.e., unrelated) to another. Thus, triangulating evidence across study designs is likely to
be a most valuable tool. A practical guide for such an approach summarizes how different methods
overcome different bias (Lawlor, Tilling, and Davey Smith, 2016). Piecing together the epidemio-
logical puzzle and understanding the inconsistencies we observe with ever-progressing resources in
data availability, measurement, and analysis is the future of the epidemiology of parenting.
Acknowledgments
RP Is supported by the NIHR Biomedical Research Centre at the University Hospitals Bristol NHS
Foundation Trust and ERC Grant Agreements (Grant refs758813; MHINT). MC time for writing
of this chapter was funded by Becas Chile scholarship, Doctorado en el Extranjero program from the
National Commission of Scientific and Technologic Research of Chile (CONICYT–Chile). PR at
the time of writing this chapter was funded by the National Institute for Health Research (NIHR-
UK) via an Academic Clinical Fellowship.
366
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11
NEIGHBORHOODS
AND PARENTING
Introduction
From genetic contributions to the selection of contexts in which their children are reared, parents
play a fundamental role in shaping children’s development (Bornstein, 2015; National Research
Council and Institute of Medicine, 2000; Okagaki and Luster, 2005). These various influences are
not separable, but rather are mutually influential (Overton, 2015). Throughout childhood and ado-
lescence, as well as beyond, parents have a strong part in determining their children’s peer groups
(Collins, Maccoby, Steinberg, Hetherington, and Bornstein, 2000; Ladd and Korfenderfer-Ladd,
2019), the educational and extracurricular resources their children experience (Conger, Conger, and
Martin, 2010; Crosnoe and Ressler, 2019; Vandell, Simpkins, and Wegemer, 2019), the neighbor-
hoods where they live, and their degree of exposure to these various settings (Mazefsky and Farrell,
2005; Proctor, 2006). Neighborhoods provide a unique context in which to understand parenting
across multiple arenas, as peer groups and schools often are embedded in neighborhoods (Leventhal,
Dupéré, and Shuey, 2015). Moreover, research reveals that neighborhood conditions may have multi-
generational ramifications, with parents’ own neighborhood exposure during childhood being asso-
ciated with their children’s subsequent development (Sharkey and Elwert, 2011).
Growing policy interest in place-based initiatives and related attention to neighborhood-level
intervention strategies to promote children’s well-being underscore the need to understand more
specifically how, when, and for whom neighborhood features matter most (Komro, Flay, and Biglan,
2011). Developmentally focused neighborhood researchers have long recognized the importance of
parenting in helping to answer these questions (Klebanov, Brooks-Gunn, and Duncan, 1994; Simons,
Johnson, Conger, and Lorenz, 1997), and numerous studies have attempted to identify how the
overlapping contexts of neighborhoods and families matter for children (Burton and Jarrett, 2000;
Nettles, Caughy, and O’Campo, 2008).
This chapter synthesizes research on parenting and child development in the neighborhood con-
text to make theoretically and empirically based recommendations for future research. To achieve
this goal, we first situate research linking neighborhoods, parenting, and child development in histor-
ical context. Next, we briefly describe particular challenges for defining and studying the overlapping
contexts of neighborhoods and parenting, with a focus on research methodology around neighbor-
hoods. This discussion is followed by a literature review of neighborhood associations with parenting.
The review begins by addressing direct links between neighborhoods and parents’ well-being and
parenting behaviors before turning to mediated and moderated models that link neighborhoods and
371
parenting with children’s outcomes. The chapter concludes with a summary of findings and direc-
tions for future research.
Historical Considerations in the Changing Demography

of Neighborhoods for Children and Families
Of persistent and great concern to researchers and policymakers alike is how growing up in a neigh-
borhood marked by poverty and associated conditions, such as crime, violence, and disorder, might
harm child and family health and well-being. In this section, we track the changing demography
of neighborhoods over the past several decades and the implications for children and families. This
historical context provides a backdrop for the research presented.
The focus on neighborhood poverty—the percentage of poor people living in a neighborhood—
emerged because of demographic changes beginning in the 1970s and continuing through the 1990s
in which poverty became increasingly concentrated. As a result, the number of neighborhoods with
high concentrations (40% or more) of poor households grew ( Jargowsky, 2003; Reardon and Bis-
choff, 2011). By contrast, poverty rates at the family level did not increase during this period and
therefore could not explain the trend. Rather, a fundamental change in the spatial organization of
poverty occurred, with consequences for poor families in particular. Poor families became more
physically concentrated and isolated than in the past, shifting attention in their direction.
Despite declines in concentrated poverty during the 1990s, the Great Recession in 2008 again
led to rising numbers of high-poverty neighborhoods ( Jargowsky, 2015; Kneebone and Holmes,
2016) and continued concern about their implications for child and family well-being. Although
much attention is placed on the increasing concentration of poverty over the past several decades,
the concentration of affluence (i.e., percentage of high-income, educated, and professional residents)
grew even more extreme than the concentration of poverty over this same period (Massey and Fis-
cher, 2003; Reardon and Bischoff, 2011). Both factors contributed to rising social inequality and the
contemporary neighborhood context in which parents are rearing their children. As described in the
remainder of this chapter, parents play a central role in determining how the changing demography
of neighborhoods shapes children’s development.
Central Methodological Issues in Studying Neighborhoods and Parenting

Neighborhood socioeconomic conditions are a key facet of describing the neighborhood context of
families and among the most common aspect studied (Mihn, Muhajarine, Janus, Brownell, and Guhn,
2017); however, there are myriad other metrics that can be, and are, used to depict neighborhoods.
In this section, we describe how neighborhoods are defined, including their various dimensions, as
well as related methodological challenges in studying neighborhoods. This information sets the stage
for the subsequent literature review.
Defining Neighborhoods and Their Dimensions

U.S. census data are readily available and provide invaluable information on the structural or sociode-
mographic characteristics of bounded geographic areas. As such, census definitions of neighborhoods
(i.e., tracts or block groups) are most often used to define neighborhood boundaries in quantitative
studies. Fortunately, neighborhoods defined by census boundaries tend to overlap in size with par-
ents’ subjective reports of the area constituting their neighborhoods (Sampson, 1997).
A critical distinction to make in defining neighborhood dimensions is between neighborhood
structure and neighborhood processes. Neighborhood structure entails compositional or sociodemo-
graphic attributes, such as median income, employment rate, and racial composition. Neighborhood
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Neighborhoods and Parenting
processes include aspects such as social organization and institutional resources. Generally thought to
be a function of neighborhood structure, neighborhood social organization describes the capacity of
residents to work together towards common goals and values and to establish institutions that pro-
mote and enforce these goals by regulating behavior (especially that of youth; Sampson, Morenoff,
and Gannon-Rowley, 2002; Shaw and McKay, 1942). Institutional resources involve the presence of
services and organizations that promote health, well-being, and general social welfare. These neigh-
borhood processes may help address the questions of how, when, and for whom different neighbor-
hood features matter for diverse parents and their children.
Given the accessibility of census data, census-based measures of neighborhood structural charac-
teristics are employed in a majority of studies ( Jencks and Mayer, 1990; Leventhal and Brooks-Gunn,
2000). Neighborhood income or socioeconomic status (SES)—a combination of social and eco-
nomic indicators—is the most commonly studied structural dimension. Researchers often separate
measures of neighborhood SES into high-SES/affluence (e.g., indexing percent of high-income resi-
dents, percent professionals, and percent college-educated) and low-SES/poverty (e.g., assessing per-
cent poor, percent female-headed households, percent on public assistance, and percent unemployed).
This distinction is made because the presence of poor and affluent neighbors may have differential
associations with child and family well-being ( Jencks and Mayer, 1990).
Unfortunately, valid and reliable neighborhood-level measures of neighborhood social processes
and institutional resources are not as readily available as the demographic features included in the
census. There are several approaches to obtaining data on these neighborhood processes. One is
using individual parents’ or children’s ratings, collected from the same sample as the individual- and
family-level data; however, these ratings are problematic because they often are confounded with
child and family outcome measures also obtained by means of participant ratings, leading to problems
of shared method variance. In addition, the reliability of such measures may be questionable because
in most cases it relies on individual rather than ecological data and corresponding data-handling
methods. In other words, sampling is not done purposively to obtain large enough samples within
neighborhoods to truly measure neighborhood (and not individual) differences in processes. A sec-
ond approach to gathering survey data on neighborhood processes consists of conducting a com-
munity survey with a sample of nonparticipants in each study neighborhood (ideally at least 15 to 30
respondents per neighborhood), yielding measures of neighborhood processes that are independent
from those obtained by study participants. Raudenbush and Sampson (1999) lay out a compelling
argument for “ecometric” standards of gathering this type of data from multiple reporters (preferably
independent of study families) to enhance reliability of neighborhood measures and to use appro-
priate statistical tools to generate neighborhood-level reliability indices. Note that it is possible to
generate similar ecometric measures of neighborhood processes from sampled children and families
if the study design incorporates neighborhoods.
There are a number of other alternative methodologies to measure neighborhood processes.
Rather than interviewing a sample of neighborhood residents or study participants, neighborhood
expert surveys may be conducted with key community leaders such as prominent religious, politi-
cal, business, and social leaders about their neighborhoods (Sampson, 2012; Tobler, Komro, and
Maldonado-Molina, 2009). Yet another option is systematic social observations, which involve
trained observers using a structured format to characterize neighborhoods through videotaping, rater
checklists, or audiotaping (Sampson and Raudenbush, 2004). Recent innovations in technology such
as online mapping tools, notably Google Street View, can be a cost-effective approach to quantify-
ing readily visible aspects of neighborhoods (e.g., parks; Odgers, Caspi, Bates, Sampson, and Moffitt,
2012). Finally, administrative data sources are available from city, state, and federal agencies such as
vital statistics from health departments, school records from education departments, and child abuse
and neglect records from human and social service agencies (Coulton and Korbin, 2007). There are
also publicly available data sources often produced for commercial purposes. Again, innovations in
373
geographic information systems (GIS) allow researchers to use these data in new and creative ways
(e.g., families’ access to green space and healthy foods; Hillsdon, Panter, Foster, and Jones, 2006).
Methodological Challenges
The fact that individual family circumstances are not independent of families’ neighborhood cir-
cumstances (i.e., they are endogenous) presents a fundamental challenge for neighborhood research
because unmeasured factors related to both neighborhood selection and parenting and child out-
comes may explain their association. In addition, the challenges around consistently defining neigh-
borhoods and obtaining rigorous neighborhood-level measurements compound the difficulties of
conducting research on the importance of neighborhoods for parenting. Although it is beyond
the scope of this chapter to thoroughly address such challenges (see Duncan and Raudenbush,
2001; Duncan, Magnuson, and Votruba-Drzal, 2015; Jencks and Mayer, 1990; Leventhal et al., 2015;
Sampson, 2008; Wodtke, Harding, and Elwert, 2011, for discussions of methodological issues in
neighborhood research), we focus in this chapter on findings from studies that examine constructs at
both neighborhood and individual levels and make an effort to account for neighborhood selection
(e.g., incorporate measures of family income, ethnicity). By applying this focus, we aim to better
understand neighborhoods as an ecological construct in parents’ and their children’s lives.
Overarching Theoretical Framework for Studying

This section describes the overarching theoretical framework guiding this chapter. At the broadest
conceptual level, Bronfenbrenner’s seminal writings on the ecological model of human development
(Bronfenbrenner, 1979; Bronfenbrenner and Morris, 2006) were a clear departure from the strictly
psychological study of development, restricted to a focus on the individual. More recently, relational
developmental systems theories (see Overton, 2015) have come to the forefront in the study of
human development, providing a lens through which to recognize the complexities of individu-
als embedded within contexts from time to culture, including neighborhoods and families. These
theories emphasize the mutual influences of individuals and contexts on one another, and reject the
premise that it is possible to separate the various levels in the developmental system (Overton, 2015).
For the study of neighborhoods specifically these theories help to elucidate how the neighbor-
hood context affects parenting and children’s development, but also how these influences are shaped
by characteristics of both the child and the parent. For example, very young children’s contacts with
their neighborhood contexts may be largely controlled by their parents, whereas adolescents may
engage more closely with the people and physical aspects of their neighborhoods, potentially having
direct influence over neighborhood features through their actions and behaviors. This idea that indi-
viduals may influence their neighborhoods represents the bidirectional nature of associations inher-
ent in these systems theories: Parents may react to their neighborhoods and children to their parents,
but these links are dynamic, with each part of the system responding to alterations from other parts
(Overton, 2015). Thus, in defining the neighborhood context for children and their parents, we focus
on the elements of neighborhoods that help to describe and explain aspects of parenting and child
development beyond what is learned from characteristics of individual families, while recognizing
that neighborhood and family are dynamic parts of the developmental system.
There are multiple ways in which parent and family characteristics are likely to combine with
neighborhood characteristics that are relevant for children (Proctor, 2006). A range of potential
models for understanding these varying constellations of associations among neighborhood charac-
teristics, parenting, and child outcomes is presented in the following section as a framework for inter-
preting research covered in the literature review. In some instances, these models are at odds, giving
374
competing explanations for how parenting in neighborhoods relates to children’s development, but
in general the models are complementary. In reality, it is unlikely that one, or even a few, of the
models can capture the full complexity inherent in relationships between parenting, neighborhoods,
and children’s development. Thus, the goal of identifying and evaluating the specific models is to
know when and for whom the different theoretical perspectives may be most informative (Bornstein,
2017), while at the same time, acknowledging the complexity inherent in the developmental system
as described by the broader theories covered.
Review of Literature on Links Between Neighborhoods and Parenting

We begin this section by discussing direct links between parenting and neighborhoods with particu-
lar attention to studies that employ rigorous methods to address individual parenting in the neigh-
borhood context. Next, we expand on these direct links to consider parenting as a mediator between
neighborhoods and children’s outcomes before addressing the interactive ways in which parenting
and neighborhoods may combine to shape children’s development.
The vast majority of studies identified for this review look exclusively or primarily at moth-
ers (National Research Council and Institute of Medicine, 2000). Thus, we refer to “parenting”
throughout this section, but unless specified to include fathers, the research reported is limited to
female primary caregivers. In addition, with key exceptions, studies reviewed were conducted with
U.S. samples. This limitation of the literature facilitates comparison across studies, but confines our
insight about the role of neighborhoods for parenting. Finally, we acknowledge that an extensive
body of research suggests the importance of neighborhood features for adults in general (Casciano
and Massey, 2012a; Ludwig et al., 2012; Ross, 2000; Wilson, 1987, 1996). The literature on neigh-
borhood associations with adult well-being is broad (Mair, Roux, and Galea, 2008; Roux and Mair,
2010; Sampson, 2003; Truong and Ma, 2006), and thus we restrict our review to studies that look at
parents specifically, given potential risk and protective factors that may be unique to parents (Nelson,
Kushlev, and Lyubomirsky, 2014).
Direct Links Between Neighborhoods and Parenting

This section looks at two types of associations between neighborhoods and parenting: First, links
between neighborhood features and parental well-being, including physical and psychological health.
Next, we review the literature linking neighborhood characteristics with parenting behaviors, such
as warmth, monitoring, and approaches to discipline.
Parental Well-Being
Experimental evidence from the Moving to Opportunity for Fair Housing Demonstration (MTO)
indicates that parents who moved from high-poverty to lower-poverty neighborhoods experienced
both psychological and physical health benefits compared with parents who stayed in high-poverty
neighborhoods (Ludwig et al., 2012; Sanbonmatsu et al., 2011). MTO results are notable because this
program had a true experimental design. Families in public housing in high-poverty neighborhoods
in five cities across the United States were randomly assigned to one of three groups: (1) receive
housing vouchers (or subsidies) to move to lower poverty neighborhoods; (2) receive traditional
housing vouchers to move to any neighborhood of their choice; or (3) serve as an in-place control
group. This random assignment allows researchers to address issues of neighborhood endogene-
ity that typically arise in neighborhood research because neighborhood residence was randomly
assigned rather than selected by families. Despite the important methodological strengths of the
experimental design, results from mobility programs like MTO must be understood in the contexts
375
of limited program uptake (families did not necessarily comply with the conditions of their assigned
treatment group) and family residential moves. Moving can be a stressful process for families and may
place unique strains on parents as they attempt to adjust to their new circumstances and help their
children transition to a new home, neighborhood, and/or possibly school and peer group (Anderson,
Leventhal, Newman, and Dupéré, 2014). Moreover, families included in MTO were low income,
predominantly minority, and receiving housing assistance; thus, these experimental results should not
be assumed to generalize across a broader range of families and neighborhoods.
To this point, a quasi-experimental study found that parents who moved from poor, predomi-
nately African American neighborhoods to middle-class, predominately European American neigh-
borhoods reported less social support compared with parents who remained in poor neighborhoods
(Briggs, 1998; Fauth, Leventhal, and Brooks-Gunn, 2008). Similarly, in a national sample, greater
neighborhood socioeconomic advantage was associated with more parental alcohol use, suggest-
ing that, outside the context of housing mobility programs, greater neighborhood SES may not be
uniformly beneficial for parents’ well-being (Chuang, Ennett, Bauman, and Foshee, 2005). Given the
general focus on health risks associated with residence in low-SES neighborhoods for both parents
and children, high-SES neighborhoods have received less research attention (see Luthar, 2003; Luthar
and Barkin, 2012, for notable exceptions). Nonetheless, these findings highlight the need for better
awareness of the norms and behaviors typical in more affluent areas, and their implications for parents
and children.
Even so, nonexperimental studies generally suggest that higher neighborhood socioeconomic
disadvantage is associated with parents’ worse mental health, less social support, and lower overall
parenting quality through neighborhood social cohesion, parents’ perceptions of community risks,
and overall family stress (Barnes, Belsky, Frost, and Melhuish, 2011; Ceballo and McLoyd, 2002;
Kohen, Leventhal, Dahinten, and McIntosh, 2008; Paschall and Hubbard, 1998; Simons et al., 1997).
Along these lines, findings from the Project on Human Development in Chicago Neighborhoods
(PHDCN), a multilevel, longitudinal study of children and families sampled from 80 diverse neigh-
borhoods, revealed that living in a neighborhood with greater collective efficacy (a combination
of social cohesion among neighbors and their willingness to intervene on behalf of the neighbor-
hood to prevent problems) was associated with parents’ reports of greater social support 6 years later
(Tendulkar, Koenen, Dunn, Buka, and Subramanian, 2012). In addition to a longitudinal cohort
study providing data on children and families, PHDCN included an independent community survey,
which provides neighborhood-level data on social processes (Leventhal and Brooks-Gunn, 2003).
As one of only a few studies to use a multilevel framework to address questions of individual devel-
opment in neighborhood context, results from PHDCN are particularly useful for examining the
overlap of parenting and neighborhoods for child development.
However, findings from another notable study in the field of neighborhood research, the Los
Angeles Family and Neighborhoods Study (L.A. FANS), identify more nuanced associations
between neighborhood social processes and parenting. L.A. FANS was conducted in 65 diverse
neighborhoods using representative sampling within neighborhoods and provides aggregated reports
on neighborhood social processes from residents, including but not limited to parents. This study
found that higher neighborhood informal social control exacerbated the inverse relation between
parenting strain and parents’ personal control, such that in some instances elements of neighborhood
collective efficacy may be detrimental for parents (Carpiano and Kimbro, 2012). Conversely, feel-
ing a strong attachment to one’s neighborhood attenuated this same association between parenting
strain and personal control. The authors interpret these findings to suggest that for some parents,
high neighborhood-level informal social control may undermine personal agency, which is unlikely
to be the case when parents feel a strong connection to their neighborhoods (Carpiano and Kimbro,
2012). Qualitative research on parents’ sense of community paints a similar picture: in high-crime
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neighborhoods, a negative sense of community may contribute to mothers’ resilient functioning

(Brodsky, 1996).
Parenting Behaviors
Turning to direct links between neighborhoods and parenting behaviors, findings from a quasi-
experimental housing mobility program suggest that, compared with families who stayed in low-
income neighborhoods, parents who moved to middle-class neighborhoods reported less stringent
monitoring of their children and used less restrictive control and discipline because they were “not
struggling to isolate their kids from neighborhood risks” (Briggs, 1998, p. 208; Fauth, Leventhal,
and Brooks-Gunn, 2007). In contrast, using the New York MTO early evaluation, Leventhal and
Brooks-Gunn (2005) found that, compared with parents who remained in high-poverty neighbor-
hoods, parents who moved to lower poverty neighborhoods were observed to use harsher parenting
with their daughters. The authors proposed that this counterintuitive finding may be related to the
short-term stress of moving, with increasing conflict between girls and their parents as a normative
component of adolescence. This same study found no differences in parents’ monitoring behaviors or
in their engagement with their children and general family routines among families who remained
in high-poverty and those who moved to lower poverty. These null findings from a 3-year follow-up
with the New York MTO sample mirror results with the full sample and at later follow-ups, which
showed limited program effects on parenting (Orr et al., 2003; Sanbonmatsu et al., 2011). Yet, in a
smaller scale natural experiment, Casciano and Massey (2012b) found that children whose families
moved into a subsidized housing development in an affluent suburb reported their parents were
more involved in their school activities than children whose families applied for residence in the
housing development but were not selected.
Once again, such mixed findings from housing mobility programs raise important questions about
how these programs are associated with parenting behaviors; however, reporter differences in parental
monitoring (i.e., parent versus youth) may help to explain at least some of these disparities. Correla-
tional studies show similar patterns of mixed results: Some find that parental monitoring and involve-
ment are lower in more socioeconomically disadvantaged neighborhoods as well as in neighborhoods
rated as having lower collective efficacy (Beyers, Bates, Pettit, and Dodge, 2003; Byrnes, Miller, Chen,
and Grube, 2011; French and Kimbro, 2011; Liu, Lau, Chen, Dinh, and Kim, 2009; Zuberi, 2016).
Nevertheless, others report that parents engage in greater monitoring in lower-SES neighborhoods
as well as in neighborhoods with lower collective efficacy and those rated by community leaders as
having fewer strengths (Chuang et al., 2005; Rankin and Quane, 2002; Tobler et al., 2009; Zuberi,
2016). Qualitative studies further indicate stringent monitoring of children is a mainstay of parenting
in disadvantaged neighborhoods, including among fathers (Cruz-Santiago and García, 2011; Letiecq
and Koblinsky, 2004). These approaches to monitoring become harder to implement as children get
older and develop more friendships outside the neighborhood (Furstenberg, 1993). Thus, in addition
to reporter differences, the divergent findings in the literature may stem from the match between
monitoring strategies and children’s developmental status, and the aspects of neighborhoods under
consideration; dissimilarities across the studies, primarily with regard to sampling, make it difficult
even to speculate about the role of these differences.
Correlational studies are more consistent with regard to other parenting behaviors, aside from
monitoring. For instance, parents residing in more affluent neighborhoods tend to provide warmer,
safer, and more stimulating environments for their young children (Klebanov, Brooks-Gunn, McCa-
rton, and McCormick, 1998). Conversely, greater neighborhood disadvantage, parent and observer
ratings of more unfavorable neighborhood conditions (e.g., lack of safety, dissatisfaction with public
services), and lower neighborhood collective socialization are associated with parents’ fewer displays
377
of affection, reports of lower warmth and support, and harsher discipline (Gonzales et al., 2011; Kle-
banov et al., 1994; McDonell, 2007; Pinderhughes, Nix, Foster, Jones, and The Conduct Problems
Prevention Research Group, 2007; White and Roosa, 2012; Wickrama and Bryant, 2003).
At least one study finds that neighborhood differences account for observed differences in dis-
cipline strategies between African American and European American parents (Pinderhughes et al.,
2007); however, in a sample of Mexican American youth and their fathers, no association between
neighborhood danger and harsh parenting was found (White and Roosa, 2012). Moreover, the asso-
ciation between neighborhood danger and parental warmth may operate somewhat differently for
Mexican Americans: Quantitative findings with this population show that, in the context of high
perceived neighborhood violence, neighborhood disadvantage is positively associated with parental
warmth. Similarly, qualitative findings reveal that in such neighborhoods, Mexican American parents
expressed high levels of warmth and made active attempts to build strong, trusting relationships with
their children (Cruz-Santiago and García, 2011; Gonzales et al., 2011). However, these findings may
depend somewhat on mothers’ acculturation and specific beliefs. Among Mexican American moth-
ers who were less acculturated to U.S. norms (compared to their more acculturated counterparts),
perceived neighborhood disadvantage was associated with less supportive parenting. Furthermore,
among mothers who reported fewer beliefs consistent with familism, greater objective neighbor-
hood disadvantage was associated with less supportive parenting and cooperation in coparenting
(Barnett, Mortensen, Gonzalez, and Gonzalez, 2016).
Qualitative research predominantly with African American families highlights other behaviors
parents may use to support their children in the context of disadvantaged neighborhoods. For exam-
ple, parents in poor neighborhoods may need to go beyond their own neighborhoods to find activi-
ties and supports and may rely on extended family networks for help in doing so (Ceballo, Kennedy,
Bregman, and Epstein-Ngo, 2012; Elder, Eccles, Ardelt, and Lord, 1995; Jarrett, Jefferson, and Kelly,
2010; Jarrett, 1997, 1999; Jarrett and Jefferson, 2003; Richardson, Van Brakle, and St. Vil, 2014). Par-
ents who feel more efficacious are more likely to seek such opportunities for their children and to
find ways to encourage their children’s development despite the challenges associated with urban
poverty (Ardelt and Eccles, 2001; Furstenberg, 1993).
Some studies attempt to identify how neighborhood socioeconomic conditions contribute to
parenting behaviors. Looking at neighborhood institutional resources as a mediator, a study with
PHDCN found that greater availability of resources, such as mental health services and afterschool
programming, partially explained the link between greater neighborhood affluence and fewer harsh
parenting behaviors (Shuey and Leventhal, 2017); family use of such resources also may help lower
the risk of child maltreatment in otherwise similar neighborhoods (Garbarino and Sherman, 1980).
Furthermore, greater neighborhood cohesion and control are linked to parents’ perceived higher
social support, which is associated with more authoritative and less permissive parenting (Byrnes
and Miller, 2012). Maternal depression may play a central role in shaping parenting behaviors across
neighborhoods: One study found that the association between neighborhood poverty and mothers’
fear of letting their young children play outside was attenuated when mothers were not depressed
(Kimbro and Schachter, 2011).
Considering neighborhood conditions (i.e., police report of crime, parents’ perceived crime, and
neighborhood poverty) as a moderator, a study of African American parents found the association
between parents’ receipt of emotional support and their nurturance and punishment with their chil-
dren differed across neighborhoods. Overall, receipt of emotional support and nurturance were posi-
tively associated, whereas receipt of emotional support was negatively associated with punishment;
however, under worse neighborhood conditions, these links were attenuated. These findings suggest
that in challenging neighborhood contexts parents may need greater and more varied support to
help them in creating sensitive and nurturing environments for their children (Ceballo and McLoyd,
2002; Earls, McGuire, and Shay, 1994). Along these lines, a study using PHDCN found that living in
378
neighborhoods with higher percentages of foreign-born and Latin American residents was associated
with parents’ use of less aggression towards their children, and among Latin American families only,
the presence of dense networks of friends and kin in neighborhoods was an additional protective fac-
tor (Molnar, Buka, Brennan, and Earls, 2003). Finally, a small study of middle- to upper-class parents
in a single neighborhood suggests that availability of social support may contribute to greater use of
a range of parenting behaviors, including punitive parenting (Freisthler, Thomas, Curry and Wolf,
2016); again, this finding highlights the need to better understand parenting in more advantaged
neighborhoods.
Mediated Models: Parenting as a Link Between Neighborhoods

and Children’s Well-Being
Mediated models expand on the direct associations between neighborhood features and parents’
well-being and behaviors by adding a connection to children’s development. The formulation of
the mediated models linking neighborhoods and child development through parenting draws largely
from literature on the family stress model (Conger et al., 1992; Gutman, McLoyd, and Tokoyawa,
2005; McLoyd, 1990; Sampson and Laub, 1994). The family stress model posits that individual fam-
ily circumstances, notably poverty, are related to children’s outcomes through the mediating role of
parental perceptions (e.g., financial strain), parenting behaviors (e.g., less sensitivity), and other fam-
ily processes (e.g., marital conflict). That is, parents experiencing the strains of objective economic
hardship behave differently with their children (e.g., exhibit hostility) and employ different parenting
strategies (e.g., harsh discipline techniques) than parents without similar economic hardships (Con-
ger, Ge, Elder, Lorenz, and Simons, 1994). Rather than considering individual family characteristics,
such as poverty status, mediated models of neighborhood influence indicate that neighborhood-level
characteristics affect parent well-being and parenting behaviors, and thereby children (Figure 11.1).
A competing perspective that has received less attention in the neighborhood literature is the
family investment model. This model argues that parents with greater economic and social means are
simply able to invest more in childrearing, thereby contributing to children’s well-being (Becker and
Tomes, 1986; Duncan et al., 2015). Neighborhood affluence may facilitate investments in particular
aspects of childrearing, including access to stimulating resources (e.g., high-quality childcare pro-
grams) and reinforcement of parenting behaviors that effectively manage children’s time and activi-
ties. In addition, neighborhood affluence can be considered another type of investment parents make
in their children (e.g., moving to a better public school catchment area). These reciprocal processes
Neighborhood Structure
• SES
• Residential
stability
• Racial/ethnic
composition Parenting Behaviors
• Sensitivity &
Parent Well- responsiveness Child Outcomes
Being • Discipline
• Monitoring &
involvement
Neighborhood Social
Processes
Institutional Resources
Figure 11.1 Mediated model.
379
emphasize the difficulties in identifying the relative contributions of neighborhood and parenting
for child development.
The mediated model is likely relevant for understanding the connection between neighbor-
hood SES and children’s outcomes, but additional pathways should be considered in conjunction
with neighborhood social processes and institutional resources. For instance, neighborhood structure
(e.g., SES) can serve as an antecedent to neighborhood social processes and institutional resources,
which would in turn influence parenting and children’s outcomes (Leventhal, Dupéré, and Brooks-
Gunn, 2009; Tolan, Gorman-Smith, and Henry, 2003). In addition, it is likely that parents who are
more overwhelmed and stressed by their neighborhood contexts may be less adept at shielding their
children from the harmful aspects of their neighborhoods (Masten, Narayan, Silverman, and Osofsky,
2015). In this way, the general mediated model noted in Figure 11.1 may serve as both an indirect
pathway for the effects of neighborhoods on children’s development and a contributing factor in
children’s direct experiences of the neighborhood. Consistent with the family stress and investment
models (Conger and Donnellan, 2007), the general neighborhood mediated model is rooted in
measures of neighborhood structural conditions (most often socioeconomic status). Robust assess-
ment of these types of mediated pathways typically comes from longitudinal data, which allow
temporal sequencing of the proposed pathways to reduce the confounding influence of bidirectional
effects; however, numerous studies test mediational pathways using cross-sectional data.
Parental Well-Being as a Mediator

In a national sample of births in Britain clustered within neighborhoods, neighborhood disadvantage
and young children’s internalizing behaviors were linked through parental mental health (Flouri,
Mavroveli, and Tzavidis, 2012); similar findings come from a smaller U.S. study ( Jeon, Buettner, and
Hur, 2014). Other studies find that this pathway is mediated by more specific parenting behaviors: In
a nationally representative sample of Canadian families, in neighborhoods with less socioeconomic
disadvantage and greater social cohesion parents reported more consistent discipline strategies, which
in turn were associated with young children’s higher verbal ability scores roughly two years later.
Conversely, in neighborhoods with greater socioeconomic disadvantage and lower social cohesion,
parents reported more depressive symptoms, which in turn predicted greater use of punitive disci-
pline strategies and children’s subsequent behavior problems (Kohen et al., 2008). Furthermore, in
a British sample of families from disadvantaged neighborhoods, harsh parenting and parenting stress
accounted for the concurrent link between neighborhood crime exposure and children’s behavior
problems (Barnes and Cheng, 2006).
Parenting Behaviors as Mediators

In addition to studies that looked at both aspects of parental well-being and parenting behaviors
described in the previous section, many studies focus exclusively on parenting behaviors as media-
tors between neighborhood characteristics and children’s outcomes. One such study relied on the
Environmental Risk Longitudinal Twin Study (E-Risk), which includes a nationally representative
sample of British children and an independent mail survey of neighborhood residents. The study
finds that parental warmth and monitoring fully explained the link between neighborhood low
SES and children’s antisocial behavior at 12 years of age, as well as increases in antisocial behavior
from ages 5 to 12 (Odgers, Caspi, Russell, Sampson, Arseneault, and Moffitt, 2012). Similar results
from U.S. and Scottish samples indicate the importance of both monitoring on its own and com-
bined warmth and monitoring as mediators in the link between neighborhood disadvantage and
adolescent outcomes (Chuang et al., 2005; Chung and Steinberg, 2006; Hogan and Kitagawa, 1985;
Neumann, Barker, Koot, and Maughan, 2010). Furthermore, nurturance, family cohesion, provision
380
of more stimulating home environments, and overall parenting quality have support as at least partial
mediators of associations between neighborhoods and children’s behaviors and possibly achievement
(Deng et al., 2006; Dupéré, Leventhal, Crosnoe, and Dion, 2010; Gonzales et al., 2011; Hurd, Stod-
dard, and Zimmerman, 2013; Klebanov, Brooks-Gunn, Chase-Lansdale, and Gordon, 1997; Mrug
and Windle, 2009; Simons, Johnson, Beaman, Conger, and Whitbeck, 1996; White and Roosa, 2012;
Wickrama and Bryant, 2003). For the most part, the studies identified in this section align closely
with the family stress model, given their emphasis on neighborhood poverty rather than affluence;
however, the family investment model may be relevant when the home learning environment and
children’s achievement outcomes are a focus (Dupéré et al., 2010; Klebanov et al., 1997).
Interactive Models: Does Parenting Matter Differently

in Different Neighborhoods?
In addition to considering parenting as a potential pathway between neighborhood features and
children’s outcomes, the next models maintain that parenting may simply matter differently in dif-
ferent contexts (Figure 11.2). That is, parenting and neighborhoods may interact to influence chil-
dren’s development differently based on the specific combinations of parenting and neighborhood
attributes. For example, in neighborhoods with higher rates of crime and violence, parents may need
Parents provide… Parents provide…

• Low sensitivity and • High sensitivity and
responsiveness responsiveness
• Inconsistent or harsh • Consistent discipline
discipline • Strong monitoring and
• Little monitoring or involvement
involvement
Compensatory Compensatory
Disadvantaged Neighborhood
• High poverty
• Few resources
• Social disorganization
Amplification Evaporation
Evaporation Amplification
Advantaged Neighborhood
• Low poverty
• Resource rich
• Strong social organization
Proximal Other risk

context most factors more
salient salient
Negative child outcomes

Positive child outcomes
Figure 11.2 Interactive model.
381
to employ more stringent monitoring and control of their children to ensure safety; by contrast, in
safer neighborhoods these same parenting behaviors may undermine children’s growing need for
autonomy. These models are informed by a risk and resilience perspective (Cicchetti, 2010; Proctor,
2006; Werner, 1995), with both neighborhood and parenting features garnering consideration as risk
and protective factors.
This section considers two different formulations of how parenting and neighborhoods inter-
act to predict children’s development: the compensatory model and the evaporation/amplification
model. Each model is described in greater detail before we present relevant findings from the litera-
ture. Given the small body of work examining parental well-being in these models, we discuss this
aspect of parenting alongside parenting behaviors for each of the two interactive models.
Compensatory Model
Most similar to traditional risk and resilience models, the compensatory model indicates that parents
can provide a buffer against adverse neighborhood circumstances, resulting in healthy child devel-
opment despite disadvantaged neighborhood contexts (Roche and Leventhal, 2009; Simons, Lin,
Gordon, Brody, and Conger, 2002; Simons, Simons, Burt, Brody, and Cutrona, 2005; White, Roosa,
and Zeiders, 2012). Parenting behaviors such as high levels of emotional support, monitoring, and
involvement are often considered protective factors for children in neighborhoods high in poverty
and lacking in social organization and resources (Brody et al., 2001).
In addition to this traditional perspective, the compensatory model allows for the possibility that
parenting behaviors typically viewed as suboptimal for fostering children’s well-being in the U.S.
majority culture (e.g., restrictive control; Steinberg, 2001) may nonetheless serve a protective func-
tion in disadvantaged neighborhoods. In this case, restrictive and controlling parenting behaviors (for
example) may compensate for neighborhood disadvantage to promote children’s well-being (Roche,
Ghazarian, Little, and Leventhal, 2011). Regardless of the specific parenting behaviors and neigh-
borhood features considered, the compensatory model frames parenting as a moderator of the link
between neighborhood context and child development, proposing that parents play a protective role
for children’s outcomes in the context of neighborhood risk.
Analyses using the Family and Communities Health Study (FACHS; Brody et al., 2001) indi-
cate that child-reported nurturant parenting was most protective against African American youths’
affiliation with deviant peers in more socioeconomically disadvantaged neighborhood contexts. The
FACHS sample is notable because African American families were sampled in neighborhood clusters
in Iowa and Georgia, including nonurban as well as urban families, thereby allowing examination
of African American families outside the context of urban poverty. The sampling by neighborhood
clusters permitted aggregation of parent and youth neighborhood reports within neighborhoods to
create neighborhood-level measures. A study using the nationally representative Longitudinal Study
of Adolescent Health (Add Health) similarly found that greater closeness in the parent-adolescent
relationship was associated with boys having fewer sex partners in neighborhoods with a higher per-
centage of single-parent families (Cleveland and Gilson, 2004) (results for girls differed and are
addressed in the section on Evaporation and Amplification Models).
Studies with families living in higher poverty or riskier neighborhoods point to a compensatory
role for supportive parenting and parental involvement in learning activities and children’s achieve-
ment, although findings are somewhat uneven across racial/ethnic groups (Dearing, 2004; Gutman
and McLoyd, 2000; Klebanov et al., 1997; Shumow, Vandell, and Posner, 1999). Furthermore, with
African American and Latin American youth as well as in the Netherlands, parental monitoring can
be a protective factor in neighborhoods with higher disorder and lower collective efficacy ( Janssen,
Weerman, and Eichelsheim, 2017; Rankin and Quane, 2002; Roche and Leventhal, 2009). Qualita-
tive work with parents of African American adolescent boys in disadvantaged neighborhoods also
382
suggests that individual families can leverage social capital to support their sons and promote positive
outcomes (Richardson and Van Brakle, 2013).
In line with the hypothesis that more restrictive parenting can be protective, or at least not detri-
mental, in more disadvantaged neighborhood contexts, three studies find that these parenting behav-
iors can serve a compensatory function for children (Dearing, 2004; Roche et al., 2011; Roche et al.,
2005). These findings suggest that the adverse associations between restrictive parenting behaviors
and children’s development reported elsewhere (see Steinberg, 2001) may not hold among families
living in more dangerous and disordered neighborhoods. This idea is further supported with a body
of qualitative work that indicates greater parental control may be protective for minority youth in
high-risk settings (Furstenberg, 1993; Jarrett, 1999; Jarrett and Jefferson, 2003). Findings with Mexi-
can American youth suggest that children’s age may further moderate the compensatory function of
fathering in challenging neighborhood contexts, with authoritative control being more protective
among younger youth whereas additional harshness of a “no-nonsense” fathering style may be pro-
tective as youth get older (White, Liu, Gonzales, Knight, and Tein, 2016). Overall, the compensatory
model is most well-researched for minority families, and African American families in particular.
Evaporation and Amplification Models

In contrast to the compensatory model, the evaporation model posits that neighborhood features
can overwhelm parenting behaviors, resulting in the “evaporation” of parenting effects on children’s
development. Simons and colleagues (2002) originally described this model in the context of greater
neighborhood disadvantage overwhelming any advantages of typically beneficial parenting behaviors
(e.g., high responsiveness) such that children displayed deleterious outcomes. This idea is in line with
Jarrett’s argument (1999) that high-risk neighborhoods demand “super-parents” to achieve even
average outcomes for their children. Findings from behavioral genetics support the premise of evapo-
ration, suggesting that genetic influences are stronger in less disadvantaged neighborhoods; whereas,
genetic contributions in the youth-parent relationship evaporate in the context of more disadvan-
taged neighborhoods (Yun and Lee, 2016). Nonetheless, it is equally plausible in the evaporation
framework that the link between ineffective parenting behaviors (e.g., low monitoring) and adverse
child outcomes might evaporate in the presence of more supportive neighborhoods. For instance,
neighborhoods with stronger collective efficacy could deter youth from engaging in risky behaviors
even when parents provide low monitoring (Browning, Leventhal, and Brooks-Gunn, 2005).
Of course, the conceptualization behind the evaporation model assumes that there must be some
association that can evaporate, or that under alternative neighborhood conditions parenting would
have a different association with children’s outcomes. The complementary hypothesis, then, is that
parenting behaviors and neighborhood characteristics can work in the same direction, resulting in
amplification, for better or for worse. That is, harsh, ineffective, or uninvolved parenting strategies
may confer greater threats to children’s development in the context of high-risk neighborhoods
(Roche, Ensminger, and Cherlin, 2007), resulting in amplification of negative outcomes for children,
whereas highly supportive neighborhood contexts may contribute to the evaporation of deleterious
associations between children’s outcomes and these same parenting behaviors. On the contrary, par-
enting strengths (e.g., warmth, consistent monitoring, and involvement) may be even more beneficial
for children’s outcomes in the presence of neighborhood assets (e.g., high collective efficacy; Simons
et al., 2005), amplifying positive outcomes for children, but in high-risk neighborhoods, the link
between these same parenting behaviors and children’s outcomes may evaporate.
Taking the case of parenting strategies typically considered as beneficial first, findings from
FACHS lend support to these models: parental authoritative control was associated with children’s
fewer behavior problems in safer neighborhoods, but the positive association between authoritative
parenting and youth behavior problems evaporated in the face of more dangerous and disordered
383
neighborhoods (Simons et al., 2002). The distinction between the compensatory role of nurturant
parenting (see previous section) and the potential for parental authoritative control to evaporate in
adverse neighborhood circumstances, both found with the FACHS sample, highlight the complex-
ity of understanding the interplay of parenting and neighborhoods. Moreover, these findings suggest
the need for strong theory and conceptualization to meaningfully investigate interactive models of
parenting and neighborhoods as predictors of children’s outcomes.
In a study using Add Health, support also emerged for the evaporation/amplification model,
but only when parents reported on their own behavior: Parent reports of acceptance towards their
adolescents were associated with adolescents’ lower reports of depressive symptoms in the context
of neighborhoods with high collective socialization, but this link was attenuated in neighborhoods
with low collective socialization. Yet, adolescent reports of parental support (versus parental reports)
were associated with adolescents’ fewer depressive symptoms regardless of neighborhood context
(Wickrama and Bryant, 2003). Findings from another study with this sample are also in line with the
evaporation/amplification model, in this case with parents’ reported knowledge of their children’s
friends: Greater knowledge of children’s friends was associated with youth’s delayed sexual initia-
tion in more socioeconomically advantaged neighborhoods only (Roche et al., 2005). Moreover,
findings from this sample reveal that better-quality parent-adolescent relationships were associated
with girls having fewer sex partners, suggesting positive amplification (results for boys were discussed
in the Compensatory Model section; Cleveland and Gilson, 2004). Conversely, in a sample of pre-
dominantly middle-class European American families, lower parental monitoring was found to be
particularly detrimental for children’s behavior in neighborhoods with higher rates of residential
instability (Beyers et al., 2003). Further research is needed to better understand the reporter differ-
ence with respect to the nature and importance of neighborhoods for adolescent well-being and if
this difference extends to outcomes beyond depression, such as sexual behavior, and whether the
evaporation/amplification models are equally meaningful for both boys and girls.
In addition to potential reporter differences, how parents’ involvement in learning activities mat-
ters for their children in various neighborhood contexts may vary by families’ race or ethnicity.
Among young children in disadvantaged neighborhoods, European American, but not African
American, children benefited from more stimulating home learning environments (Klebanov et al.,
1997). This finding, from two samples, including one nationally representative sample, supports
the compensatory model for European American children, but the evaporation model for African
American children. Overall, African American families’ neighborhoods were more disadvantaged
than European American families’ neighborhoods, suggesting that given neighborhood-level chal-
lenges, more stimulating home learning environments may not have been enough to compensate
for neighborhood conditions experienced by African American families (Klebanov et al., 1997). In
direct contrast to results for their African American peers, Dearing (2004) found that among Euro-
pean American youth, more restrictive parenting was most detrimental to academic performance in
low quality neighborhoods, again suggesting that racial or ethnic differences may be at play.
Turning to parenting strategies that are typically considered less beneficial for children’s out-
comes, findings from PHDCN show that the combination of physically aggressive parenting and low
neighborhood cohesion is especially harmful for children’s internalizing behaviors as they move from
early childhood to adolescence (Riina, Martin, and Brooks-Gunn, 2014). Research based on the
FACHS sample also endorses the negative amplification model: In more disadvantaged neighbor-
hoods, harsh and inconsistent parental discipline practices were more strongly associated with chil-
dren’s symptoms of conduct disorder (Brody et al., 2003). Again, the fact that findings from FACHS
support an array of different interactive models does not mean the results are at odds, but rather likely
reflects the different aspects of parenting under consideration. Other findings from L.A. FANS and
PHDCN suggest that deleterious associations between negative parenting behaviors and child out-
comes may evaporate in the context of supportive neighborhoods; however, this model may be more
384
relevant for adolescents than for younger children (Delany-Brumsey, Mays, and Cochran, 2014), and
for girls rather than for boys (Browning et al., 2005). In these studies, positive neighborhood social
processes attenuated adverse associations between parents’ well-being or parenting behaviors and
their children’s development.
Finally, neighborhood-level parenting values and norms may moderate the association between
parents’ use of discipline and children’s outcomes by conferring different meanings to these behav-
iors. For example, using the FACHS sample, Simons and colleagues (2002) found that parents’ greater
use of corporal punishment was associated with their children displaying more conduct problems in
neighborhoods where the overall use of corporal punishment was low; however, in neighborhoods
where corporal punishment was normative, there was no such association, consistent with evapora-
tion. Along these lines, how much families connect with their neighborhoods may contribute to
amplification: Research with African American families in a broad range of neighborhoods in Balti-
more found that young children’s internalizing behaviors were lower when mothers had a low sense
of community in more impoverished neighborhoods, but in less impoverished neighborhoods chil-
dren’s internalizing behaviors were lower when mothers had a strong sense of community (Caughy,
O’Campo, and Muntaner, 2003). These findings fit with the amplification model, suggesting that,
when children have greater connections to their neighborhood contexts (i.e., through mothers’
stronger sense of community), neighborhood SES may have a stronger association with their well-
being whereby in more advantaged neighborhoods connections are beneficial, and in less advantaged
neighborhoods they may be detrimental. White and colleagues (2016) drew on this idea of compara-
tive norms to explain their finding that less involved fathering is associated with Mexican American
youth greater externalizing over time, but only in more advantaged neighborhoods, where fathers’
engagement with their offspring may be more expected than in less advantaged neighborhoods.
Summary of Findings
Overall, the studies reviewed lend support for the general mediated model presented in Figure 11.1;
specifically, in line with the family stress model, disadvantage in neighborhood structural and social
features appears to be associated with compromised parental well-being and, in turn, children’s devel-
opment. Conversely, and more consistent with the family investment model, more advantaged neigh-
borhoods appear to be associated with adaptive parenting behaviors (e.g., warmth and monitoring)
that have favorable links to children’s outcomes. Direct tests of this full pathway are scare. Further,
contradictory findings on the direction of associations between neighborhood SES and parent well-
being, as well as some null findings (e.g., Klebanov et al., 1994), hint at the potential for a curvilinear
relation between neighborhood SES and parent well-being. Clarification from additional research
is needed. The family stress model presupposes that economic hardship will have deleterious effects
on parent well-being, but the role of neighborhood socioeconomic circumstances may be somewhat
more complicated, with individual, family, and other neighborhood features contributing to associa-
tions between neighborhood SES and parent well-being. In addition, specific neighborhood features
(i.e., social cohesion versus informal social control) may matter differentially for specific aspects of
parental well-being (i.e., depression versus feelings of personal control), pointing to the important
role theory and careful hypothesis testing could play in elucidating these various pathways (Born-
stein, 2017).
Studies examining interactions between parenting and neighborhoods lend support to each of
the interactive models. Once again, the specific parenting behaviors and aspects of neighborhood
context under consideration may be pivotal. The popular hypothesis that restrictive control may be
appropriate, providing a compensatory function, in particular neighborhood settings is supported
primarily by studies of African American and Latin American families; only one study testing this
model directly compared minority and European American families (Dearing, 2004). Moreover,
385
findings relevant to harsh parenting behaviors demonstrate a pattern consistent with the amplifica-
tion model: These types of parenting behaviors may be particularly detrimental in the context of
neighborhood risk. Taken together, these studies argue for considering certain types of discipline as
protective factors, contributing to children’s resilient functioning in otherwise disadvantaged con-
texts, but also highlight some discipline strategies as potentially harmful to children’s well-being
regardless of neighborhood context. In the case of warm and supportive parenting behaviors, and
even authoritative control, findings are consistent with the evaporation model, suggesting that neigh-
borhood disadvantage can overwhelm the typically positive association between these parenting
approaches and children’s outcomes. In general findings are qualified by family race and ethnicity—
although it is unclear if racial and ethnic differences are distinct from socioeconomic differences—
and in some cases, suggest boys and girls may be differentially affected by parenting behaviors.
An Integrated Model of Neighborhoods and Parenting

Despite numerous differences among the studies reviewed, there are themes that can be drawn from
the literature. In this section, we build on these themes to propose a general model linking neighbor-
hoods, parenting, and child outcomes that can be used to guide future research.
Support for each of the models reviewed suggests that the associations of specific parenting
behaviors and neighborhood characteristics for children’s outcomes can best be explained by dif-
ferent combinations of models. Thus, we propose the model in Figure 11.3 as a framework for
testing and interpreting research on the overlapping contexts of neighborhoods and parenting. The
overarching framework of this model is rooted in a relational developmental systems perspective,
Child Age
Neighborhoods Child Gender
Socioeconomic Conditions Race/Ethnicity &

Immigrant Status
Other Structural Characteristics
Institutional Resources
Social Processes
Proactive & Supportive Parenting Parent Well-Being

Behaviors
• Sensitivity & Support
• Consistent/Restrictive Discipline
• Involvement (developmental Reactive Parenting Behaviors
stimulation, discussion-based • Harsh/Punitive Discipline
monitoring) • Rule-Based Monitoring
1. 2. 3b.
3a.
Child Outcomes
Figure 11.3 Integrated model; 1. Primary mediated pathways; 2. Compensatory; 3. Evaporation/Amplification.
386
assuming that all constructs included in the model are mutually influential so that there are no uni-
directional direct effects. The proposed model is consistent with multiple theoretical frames: Collec-
tive efficacy theory informs direct links between neighborhoods and parenting; the family stress and
investment models are applied to indirect or mediated relations; and risk and resilience perspectives
contribute to identifying when each of the interactive models may be most appropriate. We describe
the theoretical rationale for the proposed model in greater detail throughout this section.
The proposed model integrates all the models discussed, adding a few important specifications to
help refine our understanding of these overlapping contexts in ways that comport with the literature
reviewed and can guide development of theory, research, practice, and policy (Bornstein, 2017). First,
neighborhood socioeconomic conditions are noted separately from other aspects of neighborhood
structure because of their unique role as one of the most oft-investigated constructs in the neighbor-
hood literature. Second, specific parenting behaviors are considered along two dimensions: proactive/
supportive and reactive. These dimensions are rooted in past work on constellations of parenting
behaviors (Furstenberg, Cook, Eccles, and Elder, 1999), and, in contrast to traditional parenting typol-
ogies, the two dimensions are viewed as clusters of behaviors rather than types of parents: Parents
may at different times use different or multiple strategies rather than having a static, overall parenting
style. We use the terms proactive/supportive and reactive to avoid conflating past research on racial
and ethnic differences in parenting styles with the current focus on neighborhoods and parenting.
In the proposed model, proactive/supportive parenting includes sensitivity and support, consistent
and possibly restrictive discipline, and involvement, such as provision of developmental stimulation
and discussion-based monitoring, akin to what Stattin and Kerr (2000) refer to as “child disclosure”;
as noted in studies reviewed, parental sensitivity and monitoring often work in tandem to promote
children’s development. We include restrictive discipline and control in the constellation of sup-
portive parenting behaviors given evidence that these strategies may foster children’s development
in the context of neighborhood risk (Roche et al., 2011). Together, these strategies are considered
proactive/supportive in the proposed model because they demand consistency and follow-through
from parents, and at high levels are most likely to be associated with children’s adaptive function-
ing. Reactive parenting, in contrast, is conceived of as harsh and punitive approaches to discipline
and involvement characterized primarily by rule-based strategies for monitoring. The term reactive
implies that parents may at times use these strategies in response to specific challenges or threats to
child well-being. Furthermore, these strategies need not be applied in the same sort of consistent
manner as the proactive strategies to have implications for children’s outcomes. We argue that reac-
tive parenting behaviors are distinct from low levels of proactive/support parenting behaviors, and
vice versa.
We propose that proactive/supportive parenting behaviors act as mediators between neighbor-
hood conditions and children’s outcomes (Figure 11.3 number 1), consistent with the family invest-
ment model. In neighborhoods where parents are supported, through socioeconomic advantages
as well as through institutional resources and social organization, proactive/supportive parenting
behaviors are likely to be high, transmitting neighborhood benefits to children. Yet, consistent with
the family stress model, in less advantaged neighborhoods, proactive/supportive parenting behaviors
appear to become less robust, conferring neighborhood risk to children’s outcomes; parental well-
being likely further mediates this path, at least partially explaining links between neighborhoods and
parenting behaviors. In addition, we anticipate that compromised parental well-being will contrib-
ute to greater use of reactive parenting behaviors. Greater reliance on reactive parenting behaviors,
particularly harsh and punitive discipline strategies, is also anticipated to transmit neighborhood
disadvantage to children’s outcomes.
Turning to the interactive models, we hypothesize that the compensatory model will be most
common in the context of more disadvantaged neighborhoods and higher levels of proactive/sup-
portive parenting (Figure 11.3 number 2). We anticipate that evaporation/amplification is possible
387
for both dimensions of parenting behaviors (Figure 11.3 numbers 3a and 3b), although the neighbor-
hood conditions required to produce evaporation are expected to be extreme. For example, for reac-
tive parenting behaviors to evaporate, neighborhood social organization and institutional resources
will need to be highly supportive of children, and for proactive/supportive parenting behaviors to
evaporate, neighborhoods would need to pose serious risks for children, again most likely through
weak social organization and limited institutional resources. Conversely, amplification may occur in
less extreme neighborhood contexts. In more advantaged neighborhoods with higher social organi-
zation and numerous institutional resources, amplification is most likely to occur with proactive/
supportive parenting behaviors, leading to favorable child outcomes (Figure 11.3 number 3a). In the
context of somewhat more disadvantaged neighborhoods, amplification is most likely with reactive
parenting behaviors, leading to adverse child outcomes (Figure 11.3 number 3b).
The integrated model also recognizes the central importance of children’s age, gender, and race/
ethnicity and immigrant status in determining the specific pathways that are most relevant for indi-
vidual outcomes. We expect that these individual characteristics influence every aspect of the model,
from selection into neighborhoods to additional factors beyond neighborhoods and parenting that
are likely to matter for children’s development. Given this broad importance of individual charac-
teristics, but also the limited literature examining how these characteristics shape the meaning and
interplay of neighborhoods and parenting, we do not indicate specific moderated pathways based on
these characteristics in Figure 11.3. However, the existing literature suggests that the compensatory
and evaporation models may be highly relevant for African American and perhaps Latin American
families. Minority families in the United States, especially African Americans, continue to be more
likely live in poor neighborhoods than European American families (Pendall, Davies, Freiman, and
Pitingolo, 2011). Given this inequality, demands on minority parents can be extreme (Leyendecker,
Harwood, Comparini, and Yalcinkaya, 2005; Spencer, 2007), contributing to the potential for sup-
portive/proactive parenting behaviors to evaporate in the face of neighborhood risk. Families in
extremely disadvantaged neighborhoods, nonetheless, demonstrate resilient functioning, compen-
sating for contextual risk to promote children’s well-being (Brody et al., 2001; Jarrett et al., 2010;
Roche and Leventhal, 2009).
Individual characteristics (i.e., age, gender, race/ethnicity, and immigrant status) are likely to oper-
ate in different ways from one another at different times. Although the research reviewed here notes
that all of these individual factors are relevant, there is simply not enough research to draw firm con-
clusions about when each may be more or less salient in the overlapping contexts of neighborhoods
and parenting. Thus, in the next section, we address these gaps in the literature, outlining areas for
future research with attention to individual differences.
Finally, this model is not intended to be all-encompassing, but rather to guide theory and research
on the ways parenting and neighborhoods may overlap to influence children’s development. There
are notable gaps in the model. For instance, it does not explicitly address the nuanced role of neigh-
borhood social capital and parents’ sense of community. Still, the model provides a framework for
testing competing hypotheses regarding the nature of the associations among neighborhoods, parent-
ing, and child outcomes.
Future Directions in Research on Neighborhoods and Parenting

In addition to examining the specific hypotheses generated by the model in Figure 11.3, the research
reviewed suggests three general directions for future research: consideration of developmental timing
and individual differences; greater attention to neighborhood diversity; and examination of the role
of specific neighborhood social processes and institutional resources in supporting families. Each of
these directions is discussed in turn.
388
Developmental Timing and Individual Differences

Parenting behaviors change as children get older (Lamborn, Dornbusch, and Steinberg, 1996; Molnar
et al., 2003), but other research suggests that underlying parenting strategies may remain stable even
as parents adapt their behaviors to their children’s level of development (Aunola and Nurmi, 2005).
Furthermore, length of neighborhood exposure is important for children’s outcomes (Wodtke et al.,
2011), and changes in neighborhood context matter even for adult well-being (Casciano and Mas-
sey, 2012a; Ludwig et al., 2012). Moreover, different aspects of neighborhoods are likely to be most
salient for parents when children are at different developmental periods (Leventhal et al., 2015). For
example, parents’ connections to local resources and supports, particularly quality childcare, may
be most important during early childhood, whereas neighborhood social organization and norms
may become increasingly central as children age and parents try to balance adolescents’ need for
autonomy with the safety of the surrounding neighborhood.
These differences may relate to the neighborhood and parenting models that best explain child
outcomes, with mediation potentially being most important in early childhood and the interactive
models gaining significance as children get older. The relevance of different models at different
developmental periods may explain the varying conclusions about when neighborhoods matter
most for children: Research support exists for the significance of neighborhood context in early
childhood (Anderson, Leventhal, and Dupéré, 2014; Dupéré et al., 2010; Wheaton and Clarke,
2003), middle childhood (Ingoldsby and Shaw, 2002), and adolescence (Leventhal et al., 2009;
Wodtke, 2013). Future research could address some of these questions about timing by using cohort
samples that enable direct comparison of children and their parents at different developmental
periods.
Gender differences may become more pronounced during adolescence (Leventhal et al., 2009),
making gender a key characteristic to consider alongside developmental timing. Many studies
reviewed did not examine (or report) differences by children’s gender, despite suggestive evidence
that this individual characteristic is important for both parenting and neighborhood experiences
(Bornstein, 2015; Leventhal et al., 2015), and that boys and girls may be differentially affected by the
combination of neighborhood characteristics and parenting behaviors (Browning et al., 2005; Cleve-
land and Gilson, 2004). Boys generally may be more susceptible to neighborhood conditions than
girls, and this differential sensitivity is often presumed to be due to lower parental monitoring of boys
than girls and their greater freedom outside the home (Shuey, Leventhal, Elliott, and Dupéré, 2016);
however, given the relative dearth of information, the pathways underlying any gender differences in
the joint roles of neighborhoods and parenting remain hypothetical. Future research on neighbor-
hoods and parenting should routinely examine gender differences, at least in adolescent samples, and
include mention of even null findings to generate a stronger knowledge base.
Finally, addressing differences across race/ethnicity and immigrant status for the various models
of overlap between neighborhoods and parenting is also key to clarifying the meaning of divergent
research findings. Yet, these individual characteristics are difficult to disentangle from neighborhood
features, particularly in U.S. samples where minority families, especially African Americans, continue
to be poor and to live in poor neighborhoods (Duncan, Brooks-Gunn, and Klebanov, 1994; Pendall
et al., 2011). Race/ethnicity and immigrant status in relation to parenting is discussed more fully
elsewhere (Bornstein and Cote, 2019; Bornstein, Hahn, Suwalsky, and Haynes, 2003; Bradley and
Corwyn, 2002; Halgunseth, 2019; Leyendecker et al., 2005; Magnuson and Duncan, 2019; McLoyd,
2019; National Research Council and Institute of Medicine, 2000; Ng and Wang, 2019; White et al.,
2012), but greater empirical attention to the role of neighborhoods in shaping parenting may deepen
our knowledge of the function and strengths of various childrearing behaviors across diverse families,
potentially offering insight to interventions that could support parents within their neighborhoods.
389
Research efforts tackling these questions will need to consider the different dynamics of diverse
neighborhoods (see Pattillo, 2013), a point we return to in the next section.
Neighborhood Sampling
Overall, research with more diverse neighborhoods is needed: Both research and theory on neigh-
borhood associations with parenting and child outcomes tend to focus on urban areas, notably
disadvantaged ones. Parents in rural areas are generally less emotionally supportive than parents in
urban and suburban areas (Pinderhughes et al., 2007; Sheridan, Koziol, Clarke, Rispoli, and Coutts,
2014), suggesting urbanicity may partially determine the joint roles of parenting and neighborhoods
for children’s outcomes. However, even with the use of national samples and FACHS, which allow
for comparisons of diverse neighborhoods, research on how family functioning may differ across the
spectrum of urbanicity is scarce.
Similarly, most neighborhood research focuses on poor neighborhoods, but a few studies find
that more socioeconomically advantaged neighborhoods may present some challenges to parents
(Chuang et al., 2005; Fauth et al., 2008). Thus, research is needed in affluent communities as well to
address community norms around parenting and the availability and role of support systems in these
contexts. Such research would provide greater insight into the function of neighborhood SES across
the full distribution: The difference between absolute and relative neighborhood disadvantage is
likely important for interpreting different conclusions from different studies. Despite policy recogni-
tion that neighborhoods matter for children’s well-being and a desire to improve neighborhoods to
support children and families, there is still a relatively weak grasp of the mechanisms through which
to improve child and family well-being vis-à-vis neighborhoods. As such, greater knowledge of social
processes and parenting behaviors in neighborhoods typically assumed to be “low-risk” (i.e., more
affluent ones) would provide an informative point of comparison for understanding when and how
neighborhoods confer risk versus promoting children’s development.
In addition to stratifying neighborhood samples across a broader range of urbanicity and socio-
economic status, neighborhood racial/ethnic composition must be considered. Growing awareness of
the importance of studying nonpoor minority neighborhoods notwithstanding, such neighborhoods
are less frequently the focus of research on parenting and child development than are disadvantaged,
urban minority neighborhoods (Burton and Jarrett, 2000; Pattillo, 2005). However, at least in Chicago,
African American middle-class neighborhoods are more likely to be contiguous with areas of con-
centrated poverty than are European American middle-class neighborhoods, meaning that parents and
children in middle-class neighborhoods likely have different experiences depending on the neighbor-
hood racial/ethnic composition (Pattillo, 2013; Pattillo-McCoy, 2000). Furthermore, disadvantaged
European American neighborhoods do not have the depth of concentrated poverty present in dis-
advantaged African American neighborhoods (Sampson, 2012). These well-documented distinctions
between European and African American neighborhoods do not speak to the neighborhoods of the
growing Latin American population in the United States, nor to the neighborhood context of the
many other races/ethnicities represented in the American population (see Leventhal et al., 2015, for
a more detailed discussion).
With regards to parenting, these neighborhood structural features (urbanicity, socioeconomic status,
racial/ethnic composition) likely work together to influence parents’ well-being and their approaches to
childrearing. Nationally representative data can contribute to understanding how these dimensions of
neighborhoods matter for parenting and child development, although comparisons of various types of
neighborhoods must address the qualitative differences in neighborhood experiences for individual fam-
ilies. More varied data sources are needed to identify similarities and differences in parenting approaches
within specific types of neighborhoods (e.g., predominantly Latin American, rural neighborhoods).
390
Neighborhood Social Processes and Institutional Resources

In addition to neighborhood sampling, measurement of neighborhood social processes and institu-
tional resources is a key issue for study design. Despite the challenges of measuring neighborhood-
level social organization and resources discussed previously, numerous techniques for assessing these
constructs are available and should be employed to rigorously test the processes through which
neighborhoods may matter for parents and children (Leventhal et al., 2015). Beyond understanding
nuances around the meaning of constructs like social cohesion and enforcement of norms, future
studies should further investigate the potential importance of neighborhood-level parenting norms,
which some research indicates help to explain how and when specific parenting behaviors matter for
children (Fletcher, Darling, Dornbusch, and Steinberg, 1995; Simons et al., 2002).
Compared with neighborhood social processes, neighborhood institutional resources have
received scant attention in relation to parenting. Some qualitative work alludes to the processes
involved in low-income parents accessing resources in more disadvantaged neighborhoods, includ-
ing seeking resources beyond the neighborhood ( Jarrett et al., 2010; Jarrett, 1999; Richardson et al.,
2014), with some acknowledgment of neighborhood institutions supporting parents (Kissane, 2010;
Small, 2006). However, studies have not addressed questions about variability across neighbor-
hoods in access to resources, or how lack of resources within a neighborhood may be related to
parenting behaviors and children’s development. Further research is needed to better understand
specific neighborhood contexts that promote the use of high-quality resources, and the ways in
which neighborhood resources contribute to or interact with specific parenting behaviors. Again,
neighborhood socioeconomic conditions may not be linearly associated with the availability and
quality of resources; for instance, resources like Head Start that are targeted to low-income fami-
lies are more available in lower than higher income neighborhoods (National Survey of Early
Care and Education Research Team, 2016). Improving neighborhood resources is a key tenet of
neighborhood-based intervention strategies, and thus developing an empirically grounded assess-
ment of how parents use local resources is imperative (Lareau, 2011). In sum, we see these three
areas—exploring individual neighborhood variability, expanding neighborhood diversity, and cap-
turing neighborhood processes and resources—as important next steps for understanding parenting
in neighborhood contexts.
Conclusions
Child development takes place at multiple levels and across many contexts (Bronfenbrenner and
Morris, 2006). Parents influence the neighborhoods where their children live and children’s peer
groups, school experiences, and access to other contexts. In fact, one study suggests that, when family
functioning is “exceptional,” young adolescents demonstrate the best outcomes regardless of neigh-
borhood context (Gorman-Smith, Henry, and Tolan, 2004). Even given this backdrop, the bulk of
the research reviewed reveals that parenting matters for how children experience and respond to
their neighborhood contexts. Conclusions about the specific directions of these associations remain
tentative given the complexity of the questions and the limited neighborhood research that has
used rigorous strategies to explicitly address issues of selection and omitted variable bias. Also of
importance to acknowledge is that a majority of the research on neighborhoods and parenting is
U.S.-based, with several exceptions (e.g., Canada), and its generalizability to other countries remains
to be tested. Nonetheless, enough evidence has amassed to compel further work on neighborhoods
and parenting, with attention to methodological issues and both individual and neighborhood vari-
ability. Our goal in developing a novel integrated model is to lay the groundwork for this next phase
of research, taking into consideration what we already know and where we need to go.
391
Acknowledgments
Tama Leventhal would like to acknowledge the support of the Foundation for Child Develop-
ment, the John D. and Catherine T. MacArthur Foundation, and the William T. Grant Foundation.
Elizabeth Shuey’s views are her own and do not necessarily reflect those of the Organisation for
Economic Co-operation and Development or its member countries.
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12
PARENT EDUCATION
ATTAINMENT AND PARENTING
Pamela E. Davis-Kean, Sandra Tang, and Nicholas E. Waters
Introduction
When examining the topic of parenting, the focus is often on the behaviors of the parent toward
their children and, indeed, is often defined in this way. Parents, however, are individuals (generally
adults) who derive the behaviors (parenting) toward their children based on the resources they have
available to them. These resources can be quite complex, such as their self-regulation and personality
traits (Bornstein, Hahn, and Haynes, 2011; Smith et al., 2007) but also their own problem solving and
cognitive ability (Anger and Heineck, 2010; Rueter and Conger, 1998). In much parenting research,
a parent’s cognitive ability has been measured by proxy variable using the demographic information
on the parent’s education attainment (e.g., years of formal schooling). In this chapter, we review why
this one variable is important for understanding both parenting beliefs and behaviors and subsequent
influence on child development. This chapter begins with an overview of the important role parent
education provides as a demographic variable that relays information on the social resources that
may be available for parenting children. This is followed by a discussion of the important relation
that parent educational attainment has with constructing the home environment in which children
develop. In the next section, we outline the literature on parent and school interactions and how
parent educational attainment relates to parental educational involvement in school and home. This
chapter concludes with suggestions for the future direction of research in understanding the com-
plexity of the family environment and child development.
Parent Versus Maternal Educational Attainment

Throughout the chapter the term parental educational attainment is used to note that the educa-
tion of all of the caregivers of the child is important in parenting. One curiosity about the parenting
education literature is that it often only considers mothers’ education as a predictor of children’s
outcomes. One explanation for this practice is that, typically, mothers are the primary caregivers and
as such their characteristics are more likely to influence children’s well-being (Murray and Richards,
2019). Another explanation is that with increasing rates of single parenthood, a substantial propor-
tion of families may not provide information about paternal or other caregiver education, but will
be able to provide information about maternal education (Duncan and Brooks-Gunn, 1997). Finally,
individuals often choose partners with similar levels of education, so it is difficult to disentangle
effects of either partner’s education in a family. Studies using genetically informed designs have
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Parent Education Attainment and Parenting
attempted to isolate the unique contribution of dual parent, heterosexual families. Higher levels of
both maternal and paternal education benefit children’s academic outcomes (Neiss and Rowe, 2000;
Plug and Vijverberg, 2003). This research suggests that, to understand the role of parent education
on child development, more attention is needed to the unique contribution that each parent’s years
of education may have on children’s development and outcomes. At the moment, however, the
parenting literature continues to be dominated by the examination of maternal education or the
female caregiver of children in the household and children’s positive academic achievement (Have-
man and Wolfe, 1995; Sirin, 2005; White, 1982), school readiness (Christian, Morrison, and Bryant,
1998; Dotterer, Iruka, and Pungello, 2012; Raviv, Kessenich, and Morrison, 2004; Seefeldt, Denton,
Galper, and Younoszai, 1999), school grades (Sirin, 2005; Smith, 1989), tests of academic achieve-
ment (Alexander, Entwisle, and Bedinger, 1994; Alwin and Thornton, 1984; Dauber, Alexander, and
Entwisle, 1996; Davis-Kean, 2005; Mercy and Steelman, 1982; Smith, 1989), and educational attain-
ment (Alwin and Thornton, 1984; Ensminger and Slusarcick, 1992; Mare, 1980; Rumberger, 1983;
Sewell, Haller, and Ohlendorf, 1970). There is also evidence that obtaining additional years of educa-
tion for mothers is associated with increases in children’s cognitive skills and academic achievement
(Gennetian, Magnuson, and Morris, 2008; Harding, Morris, and Hughes, 2015; Magnuson, 2003;
Magnuson, 2007; Magnuson, Sexton, Davis-Kean, and Huston, 2009; Tang, Davis-Kean, Chen, and
Sexton, 2016). Thus, the majority of research reviewed in this chapter is related to how female car-
egivers’ beliefs and behaviors are influenced by their years of formal schooling. Although this focus
gives a selected and perhaps incomplete overview of the role that parent educational attainment may
play in why we see the variety of parenting styles in families, it provides a good starting point for
thinking about how additional years of education can relate to the behaviors of any caregiver and
their children’s outcomes, and provides guidance on the direction of future research that is needed to
fully understand this important predictor of parent and child outcomes.
Parental Educational Attainment or Socioeconomic Status?

It is clear that parental educational attainment is important in understanding how parents form their
parenting styles, but education attainment is also considered part of an expanded set of variables
(most notably family income and occupational status) that define the socioeconomic status or social
class of the family. Thus, a problem that arises in examining parent’s years of schooling is how it plays
a unique role in the larger construct of family socioeconomic status. Conceptually, SES refers to eco-
nomic resources and/or the social positioning within society that derives from economic resources
or “the capacity to create or consume goods in our society” (Hauser and Warren, 1997, p. 179).
However, as Bollen and colleagues (2001) argued, operationally it serves as shorthand for social and
economic characteristics that are “believed to be important, but the rationale or meaning of which is
not always made clear” (Bollen, Glanville, and Stecklov, 2001, p. 157). Because it may be difficult to
measure a family’s access to economic resources or their position in a social hierarchy directly, indi-
cators of parents’ education, occupation, and household income typically are used to measure SES.
When researchers seek to examine social class or gradient, they generally approach it in two
ways (Bollen et al., 2001). The first approach seeks to obtain a single indicator or variable that sug-
gests more of a latent representation of SES. In this approach, one of the indicators of SES is used
(e.g., income), which is used to represent the larger construct of socioeconomic status or class.
Reviews of the most common way that SES is measured across studies suggest that, when a single
indicator of SES is used, it is almost always educational attainment (Bollen et al., 2001; Elo and Pres-
ton, 1996; Ensminger and Fothergill, 2003). Elo and Preston (1996) suggested that this dominance
of educational attainment as the primary SES indicator is due to the fact that educational attainment
can be determined for most individuals but large parts of the population do not have occupations
or income. Thus, it is a more reliable indicator of SES. Similar to the single indicator approach is the
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Pamela E. Davis-Kean et al.
index approach whereby multiple indicators of socioeconomic status are combined into a single scale
that is then used as an additive index of social class or social standing. The most common and well-
known index is the Hollingshead Four-Factor Index (Bornstein, Hahn, Suwalsky, and Haynes, 2003).
Although the single indicator or index approach has some advantages including ease of measure-
ment, reduction in complexity of variables, and less burden on the individual in providing informa-
tion, it has been criticized for not adequately taking into account the role that each indicator of
socioeconomic status has in understanding parenting or child development (Duncan and Magnuson,
2003). Hence, the alternative approach is to recognize the unique contribution of each SES indi-
cator (education, occupation, and income) as a source of social inequality and stratification and to
model the individual prediction of each indicator (Duncan and Magnuson, 2003). That is, educa-
tion, income, and occupation, although related constructs, exert independent and unique effects on
parents and children through differing mechanisms (Corwyn and Bradley, 2003; Davis-Kean, 2005;
Duncan and Magnuson, 2003). Davis-Kean (2005), for example, demonstrated that parental edu-
cational attainment and income both related to the resources available in the home environment,
but that parental educational attainment had as stronger influence on the home environment espe-
cially related to beliefs and behaviors and also directly to children’s achievement. Similarly, Duncan
and Brooks-Gunn (1997) replicated analyses across 12 groups of researchers working with different
developmental data sets to determine the effects of household income on children and adolescents’
well-being. Analyses conducted on eight of these data sets that contained measures of maternal edu-
cational attainment demonstrated that maternal education was significantly associated with better
cognitive and educational outcomes for children after statistically controlling for income and other
important demographic correlates. Thus, to understand the role of any socioeconomic indicator on
parenting, each indicator and the role it may have on parenting beliefs and behaviors need to be care-
fully considered and then appropriately modeled for where they may have their strongest influence.
How to Measure Parent’s Educational Attainment?

Another challenging issue for examining parents’ educational attainment is how to measure the con-
struct. It is important to understand what the questions are that need to be answered regarding the
role of parents’ educational attainment. As with all studies, it is important to understand how meas-
urement fits the theory or questions that are being examined in a study. In 2012, the National Center
for Educational Statistics (NCES) convened a group of scholars to review the issue of the measures of
multiple socioeconomic indicators to try and standardize the way these were ascertained in surveys
and interviews (U.S. Department of Education, 2012). The way that this construct is measured varies
widely across disciplines, and understanding these differences is important for understanding how
parent educational attainment relates to parenting.
There are researchers, for example, who are interested in the category of schooling (whether or
not parents have completed high school or received a Bachelor’s degree). In this situation, individuals
are characterized by the highest degree or level of education that they have completed. In some stud-
ies, this amounts to using a set of dichotomous variables (e.g., 1 = yes, parent received a high school
diploma; 0 = no, parent did not get a high school diploma). In other studies, categories are placed on
an ordinal scale (e.g., 1 = less than a high school diploma; 5 = graduate degree). This approach has
the benefit of distinguishing between different types of post-secondary education, but also assumes
that schooling that does not result in a degree does not affect children’s well-being.
Another approach that is often used by researchers is the continuous measurement of years of
schooling or amount of schooling parents have received by a certain age in their development. In this
instance, the measurement assumes that an additional year of education at the low end of the scale
has the same effect as an additional year of school at the high end. This method can be problematic
402
when examining the role of educational training that happens after high school but prior to a col-
lege degree (e.g., vocational or community college). All of these are equated as the same post-high
school experience, and it is not clear that the assumption of increased education is appropriate for all
developmental outcomes. It is important, then, for the choice of measurement for studying parental
educational attainment to be guided by the theoretical or conceptual framework of the role that edu-
cational attainment may play in explaining parenting. Is the researcher interested in how additional
years of education relate to a parent’s behavior, or is there is a categorical threshold of education that
matters for observing certain parenting? These are critical differences to be aware of when reading
the literature and understanding the role of educational attainment on parenting.
The Role of Parent’s Educational Attainment in Parenting

Reviews by Goodnow (1995) and McGillicuddy-De Lisi and Sigel (1995) highlighted all the con-
tribution of parents’ cognitive skills, beliefs, and schemas to parenting practices and child outcomes.
These cognitions and schemas interact with each other and other contextual influences, such as
social support, economic constraints, and children’s behaviors, to produce parenting behaviors
(McGillicuddy-De Lisi and Sigel, 1995). Given that education does not explicitly teach adults how
to be better parents, what do adults learn in school that makes them good parents? Research by
Bronfenbrenner (1958) and Hoffman (2003) suggests that parental educational attainment influences
parenting by facilitating parents’ ability and willingness to seek out expert advice about rearing their
children. Michael (1982) also argued that higher levels of education increase adults’ abilities to seek
out and synthesize information. He further suggested that this synthesis of information leads to bet-
ter decision making and greater efficiency in meeting goals. Thus, higher levels of education might
improve parenting by increasing parents’ proclivity to seek out and evaluate information about chil-
drearing, their child’s well-being, and other important aspects of family life.
Parental educational attainment appears to also increase positive parenting behavior across a wide
spectrum of child outcomes. Compared to parents with lower levels of education, more highly
educated parents are more likely to explicitly define higher levels of education as desirable, encour-
age their children to do well in school, and have higher expectations for their children’s academic
achievement (Alexander et al., 1994; Dauber et al., 1996; Davis-Kean, 2005; Grolnick and Slowiaczek,
1994; Lee and Croninger, 1994; Neitzel and Stright, 2004). Parents with higher levels of educational
attainment have teaching styles that promote children’s development (Bee, Van Egeren, Pytkowicz
Streissguth, Nyman, and Leckie, 1969; Carr and Pike, 2012; Harris, Terrel, and Allen, 1999), engage
their children in higher quality verbal interactions (Hoff, 2003; Raviv et al., 2004; Richman, Miller
and Levine, 1992; Tapia Uribe, LeVine and LeVine, 1993; Tamis-LeMonda, Kuchirko, Escobar, and
Bornstein, 2019), provide cognitively stimulating learning environment and literacy activities in the
home (Davis-Kean, 2005; Kohl, Lengua, and McMahon, 2000; Linver, Brooks-Gunn, and Kohen,
2002; Tamis-LeMonda, Shannon, Cabrera, and Lamb, 2004), and are more comfortable and involved
with their children’s education, teachers, and educational institutions (Brody and Flor, 1998; Steven-
son and Baker, 1987). Finally, higher levels of parental educational attainment are associated with
higher levels of warmth and sensitivity in parent-child interactions (Bradley et al., 1989; Davis-Kean,
2005; Klebanov, Brooks-Gunn, and Duncan, 1994; Tamis-LeMonda, Briggs, McClowry, and Snow,
2009), and lower levels of hostility (Fox, Platz, and Bentley, 1995; Dotterer et al., 2012). Thus, the
role of parents’ education in parenting and child development is important to understand. Similar
to other context variables, such as income, the pathway from parents’ years of education obtained to
child development are indirect rather than direct. That is, parental educational attainment influences
children’s behaviors through parenting. The next sections of this chapter review these important
pathways.
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Parenting and the Home Environment

The relation between parents’ educational attainment and child development has been established
in the parenting literature (for reviews, see Conger and Donnellan, 2007; Hoff, Laursen, and Tardif,
2002). More recently, parenting researchers have begun to investigate the pathways through which
education may be important to parenting (e.g., parental beliefs and behaviors) and that may explain
important variation in a host of children’s developmental skills (Bradley and Corwyn, 2002; Davis-
Kean, 2005; Hart and Risley, 1995; Hoff, 2003; Klebanov et al., 1994). In general, family process
models of parenting have examined how parental behaviors and aspects of the home environment
influence children’s development (Conger et al., 2002; Linver et al., 2002), whereas other models
have expanded the focus and examine the role of parental beliefs regarding childrearing and edu-
cational expectations that influence parent behavior and, subsequently, child outcomes (Alexander
et al., 1994; Davis-Kean, 2005; Davis-Kean and Sexton, 2009).
This section reviews the literature on the relation between parental educational attainment and
parents’ beliefs and expectations regarding their children’s development and schooling. Next, the
impact of parental educational attainment on parenting behaviors and the structure of the home
environment is explored with an emphasis on the link between parents’ beliefs and behavior. This
section concludes with evidence from studies demonstrating how increases in maternal education
relate to subsequent changes in parenting behaviors and children’s academic performance.
Parenting Beliefs and Educational Expectations

Parental beliefs include parental goals for children’s development and the role they see for themselves
in helping children achieve those goals. Research has provided evidence that parents from different
educational backgrounds have different beliefs concerning their children’s early development and
learning (Hoff et al., 2002). As children grow older and enter school, parental educational attainment
also predicts the accuracy of the beliefs and expectations they hold about their children’s academic
performance (Alexander et al., 1994), and these educational expectations are an important predictor
of children’s achievement (Davis-Kean, 2005) as well as the change in their achievement over time
(Davis-Kean and Sexton, 2009; Tang et al., 2016).
Childrearing Beliefs
Differences in parents’ beliefs and expectations regarding their children’s development have been
demonstrated as early as toddlerhood. For example, Tardif, Au, Wellman, and Nakamura (2000)
conducted a cross-national study of mothers in the United States, China, Hong Kong, and Japan to
assess whether there were socioeconomic-related differences present in mothers’ estimates of the age
at which toddlers achieve a range of developmental milestones. Across all cultures, socioeconomic-
related differences were present in mothers’ estimates. Mothers with higher levels of education esti-
mated that their children were able to produce their first sounds and say their first words earlier than
mothers with lower levels of education; in contrast, mothers with lower levels of education expected
their children to be toilet trained and exhibit formal manners earlier than mothers who had com-
pleted more years of education. These findings demonstrate distinctions in parental expectations and
values regarding children’s achievement of early cognitive and social skills.
The role parents envision for themselves in cultivating these early skills has also been found to
vary across different educational groups. Parents from higher education backgrounds have been
shown to view time spent with children as an investment, and are more likely to view problem-
solving contexts as an opportunity for their children to learn important strategies and skills (Guryan,
Hurst, and Kearney, 2008; Neitzel and Stright, 2004). These early education-based distinctions in
404
parental beliefs are fundamental in determining the ways in which parents’ approach children’s learn-
ing and the support they provide in doing so. As children grow older and encounter the schooling
environment, these beliefs may also translate into the priority parents place on their children’s aca-
demic success and the behaviors they engage in to cultivate an environment in which children can
achieve the goals set forth by their parents.
Educational Expectations
In addition to parents’ perception of children’s skills, parents’ level of educational attainment is an
important indicator of the expectations that they hold for their children’s academic success. As chil-
dren enter school, socioeconomic differences have been observed in parental beliefs and expectations
concerning their children’s academic performance such that parents from higher educational back-
grounds are more accurate in estimating their children’s abilities than their less educated counterparts
(Alexander et al., 1994). The authors suggest that the accuracy of parents’ representations of their
children’s academic skills may be essential in their ability to provide home and learning environments
that fit their children’s needs. Theoretically, these demographic differences in parents’ representations
of children’s abilities also influence the expectations that parents hold for their children’s academic
success.
Indeed, several studies demonstrated that more highly educated mothers have higher expectations
for their children’s school success, including long-term beliefs such as whether their children will
attend and complete college (Carneiro, Meghir, and Parey, 2013; Suizzo and Stapleton, 2007). Meta-
analyses indicate that these educational expectations are one of the best predictors of children’s aca-
demic success (Fan and Chen, 2001; Jeynes, 2007). Additionally, research conducted by Davis-Kean
and colleagues suggests that these expectations are indirectly related to children’s academic skills
through the resources parents provide children and the behaviors (e.g., reading to children, educa-
tional involvement) they engage in within and outside of the home environment, highlighting the
pathways whereby these expectations ultimately lead to variation in children’s academic outcomes
(Davis-Kean, 2005; Davis-Kean and Sexton, 2009; Tang et al., 2016).
Parenting Behaviors and the Home Environment

Parenting behaviors are broadly defined as the ways in which parents interact with their children and
structure their environments, both inside and outside of the home, in an effort to shape children’s
development (Bornstein, 2015; Hoff et al., 2002). Researchers have used various methods, including
direct observations of natural interactions and facilitated scenarios as well as self-report instruments,
in an effort to better understand various socialization practices and priorities that parents across
socioeconomic strata engage in, and how they are, in part, responsible for variation in children’s
development (Conger and Donnellan, 2007; Lugo-Gil and Tamis-LeMonda, 2008). This section
summarizes the extant literature on the early home language environment and its association with
children’s emergent language skills. Then, the impact of cognitively stimulating resources and activi-
ties parents use to foster their children’s cognitive and behavioral development is discussed. Finally,
how variation in parental educational attainment produces distinct profiles in the emotional climate
as well as the management and discipline strategies parents use and the ways in which they affect
children’s developmental trajectories is explored.
Language
There is a large body of literature documenting socioeconomic-related variation in both the quantity
and quality of parent-child verbal interactions and its relation to children’s language development.
405
In a seminal study of socioeconomic variation in the early home language environment, Hart and
Risley (1995) observed striking variability in the amount of language children were exposed to in
the context of the home as a function of their socioeconomic background. Since then, considerable
work has been conducted to further understand how various aspects of socioeconomic status are
related to the amount and complexity of children’s language exposure and, subsequently, to children’s
language and broader cognitive development (for review, see Hoff et al., 2002).
In one such study, Hoff-Ginsberg (1998) found that college-educated mothers spoke more to
their 2-year-olds, asked them more questions, used fewer directives, and exhibited more responses
that were contingent to their children’s utterances than less educated mothers. In a follow-up inves-
tigation, Hoff (2003) found that children from more highly educated parents also grew more in
the size of their productive vocabularies than the children of less educated parents. These patterns
of growth were fully mediated by properties of maternal speech, explaining the differential growth
patterns observed in children’s language skills. Raviv and colleagues (2004) built on these findings,
using a large-scale dataset and incorporating additional parenting behaviors into the model of chil-
dren’s language development. Findings from this study indicate that both parenting measures, cogni-
tive stimulation and maternal sensitivity, mediate the association between maternal education and
children’s language skills, providing evidence for additional pathways whereby parents’ educational
background and other sociodemographic variables are intricately linked to children’s emergent lan-
guage skills (Raviv et al., 2004).
Cognitive Stimulation
One way in which parents invest in their children’s development is through the provision of educa-
tional materials and the use of practices intended to facilitate children’s cognitive growth. Generally,
mothers with higher levels of education have more economic and social resources to invest in their
children’s learning and therefore have the opportunity to acquire resources and generate learning
opportunities for their children both within and outside of the home (Bornstein, 2015; Bradley and
Corwyn, 2002; Bradley, Corwyn, McAdoo, and García Coll, 2001). Simply spending more time with
one’s child has been found to vary as a function of maternal years of education, such that more edu-
cated mothers spend more time with their children (Carneiro et al., 2013; Guryan et al., 2008) and
they spend it performing activities that are more likely to enhance their children’s development (Carr
and Pike, 2012; Huston and Aronson, 2005; Kalil, Ryan, and Corey, 2012). Studies across various eth-
nicities, ages, and skill domains have established that one way in which parental educational attainment
influences children’s developmental outcomes is through stimulation in the home environment (Brad-
ley and Corwyn, 2003; Huston and Aronson, 2005; Raviv et al., 2004; Tamis-LeMonda et al., 2004).
Moreover, parents who have completed more education themselves also tend to have high aspira-
tions for their children’s education, which in turn is associated with using more effective strategies to
help children excel academically (Conger and Donnellan, 2007). Several studies have been conducted
to better understand the unique association between parents’ educational backgrounds (i.e., over and
above the influence of income), the expectations they hold for their children, and the behaviors
they engage in that influence their children’s academic skills across various periods of development.
Halle, Kurtz-Costes, and Mahoney (1997) found that, in a sample of low-income African American
families, mothers of higher educational backgrounds held higher expectations for their children and
that these beliefs predicted the use of more achievement-enhancing behaviors in the home, which
in turn predicted children’s higher math and reading performance. Using a nationally representative
cross-section of 8- to 12-year-olds, Davis-Kean (2005) found that parents’ expectations for children’s
school success were related to parental provision of cognitive stimulation. For both European Ameri-
can and African American children, the presence of reading resources and parenting behaviors in the
home environment predicted children’s achievement. In a follow-up investigation, Davis-Kean and
406
Sexton (2009) tested whether these pathways replicate when including expectations and behaviors
at kindergarten in predicting children’s academic achievement at third grade. This was found to be
the case. Both prior parental educational expectations for their children and the investments they
made to foster that success in kindergarten were predictors of children’s academic skills at third grade.
These findings underscore the relation among parental educational attainment, parental beliefs, and
cognitive stimulation in the home, but other aspects of parental practices are affected as well. In the
next section, the literature on variation in the emotional climate and managerial practices parents
engage in when interacting with their children, and the role parental education plays in shaping these
profiles, are reviewed.
Emotional Climate and Management

Parental education is also associated with differences in the emotional climate of the home (e.g., sen-
sitivity, warmth, responsivity) throughout childhood, with more highly educated parents generally
engaging with children in a more supportive manner (Bradley et al., 1989; Klebanov et al., 1994;
Tamis-LeMonda et al., 2004), whereas less educated mothers have been found to use more direct
control practices and discipline with their children (Hoff et al., 2002). Studies have demonstrated
associations between parental educational attainment and early language skills through sensitivity
(Raviv et al., 2004). Others have produced mixed findings in predictions to academic achievement
via warmth, especially across various ethnic groups (Davis-Kean, 2005; Davis-Kean and Sexton, 2009;
Linver, Brooks-Gunn, and Cabrera, 2004). The reason for these incongruent findings may be related
to the way in which warmth is conceptualized. That is, what might be considered “warm” may vary
from one ethnic group to the next. Lending support to this idea is the fact that ethnic differences
emerge in investigations of the use of management and discipline strategies by parents of various
educational backgrounds. For example, Dotterer and colleagues (2012) found support for a moder-
ated mediation such that sensitive parenting mediated the association between socioeconomic status
and children’s school readiness for European American families only, whereas negative and intrusive
behaviors mediated this association for both European American and African American families.
These results underscore the importance of considering cultural variation when conceptualizing and
measuring parenting (Dotterer et al., 2012).
Increases in Maternal Education and Changes in Parenting

Parental educational attainment has often been measured as if it were static, even though it is increas-
ingly common for adults to pursue additional education following the birth of their child, particu-
larly for adults from low-income households (Love et al., 2002). Additional schooling may shape
mothers’ expectations for their children’s education and enable parents to create better home learn-
ing environments (Alexander et al., 1994; Davis-Kean, 2005). Several studies have been conducted to
isolate the effect of increases in parents’ educational attainment on aspects of parenting and the home
environment as well as their children’s academic skills. Results indicate that, in general, participation
in additional education after the birth of a child yields positive results, with a few exceptions. Some
studies have observed benefits for all families (Gennetian et al., 2008; Harding et al., 2015), whereas
others have observed effects limited to only those at the lower end of the educational distribu-
tion (Magnuson et al., 2009). Magnuson (2007), for example, found that for young parents with
low initial levels of education, increases in parental education were associated with improvements
in their children’s reading skills from ages 6 to 10 as well as increases in the quality of the home
environments. In comparison, these benefits were not observed for older and more highly educated
parents. Other investigations have employed various quasi-experimental methods, using instru-
mental variables (Gennetian et al., 2008) and principal scores (Harding, Morris, and Hill, 2017)
407
to make casual inferences about the effect of participation in education on aspects of parenting, the
home environment, and children’s cognitive and academic skills. Thus, additional years of education
appear to play an important role in how parents provide both language skills and an enriched home
environment to their children. Another area where educational attainment has a profound effect and
is more explicit is in parenting behaviors toward their children’s schooling. The next section details
the role of parents’ educational attainment and their explicit behaviors toward advancing the educa-
tion of their children.
Parent Educational Attainment and Children’s Schooling

A substantial proportion of parenting behaviors is focused on promoting their children’s educational
achievement. Parents’ educational involvement has garnered attention as an important contributor
to students’ educational success (U.S. Department of Education, 2001), and educational policies, such
as No Child Left Behind, have highlighted the important role of parents in their children’s school-
ing outcomes. The emphasis on increasing all parents’ educational involvement stems from the idea
that children learn in homes as well as in schools, thus recognizing the value of parents as important
contributors to children’s educational outcomes.
Syntheses of the literature support the notion that parents play an important role in their chil-
dren’s educational success. In general, when parents are highly engaged in children’s education, chil-
dren exhibit better academic outcomes (Fan and Chen, 2001; Jeynes, 2005, 2007; Hill and Tyson,
2009; Hill and Taylor, 2004). High levels of parent involvement are associated with improved aca-
demic outcomes in mathematics, science, and positive socioemotional development (Domina, 2005;
Englund, Luckner, Whaley, and Egeland, 2004; Grolnick and Slowiaczek, 1994; Hoover-Dempsey
and Sandler, 1995; McNeal, 1999). The strongest effects have been found on literacy, a domain that
is strongly linked to positive achievement in other academic domains (Lesaux, Hastings, Kelley, Mar-
letta, and Russ, 2010).
What Is Parent Educational Involvement?

Parent educational involvement is a multidimensional construct that can be defined in a variety of
ways (Epstein, 1995; Fan and Chen, 2001; Fantuzzo, Tighe, and Childs, 2000). Christenson and Sher-
idan’s framework (2001) is used in this chapter because of its conceptually clear distinction between
two general types of parent engagement, involvement in the school and home.
School-Based Involvement
School-based involvement is the most visible and recognizable form of parent involvement because it
encompasses parents’ interactions within the school and with school personnel. In the United States,
the most common forms of school-based involvement include attending parent-teacher conferences,
volunteering at school, and attending school events (U.S. Department of Education, 2006). In gen-
eral, school-based involvement is linked to higher student achievement and better behavioral out-
comes across ethnicities and socioeconomic statuses (Fan and Chen, 2001; Jeynes, 2005). However,
research highlights variability in the value of parents’ school-based involvement for certain subgroups
of students. Benner and colleagues (2016), for example, found that school-based involvement was
most beneficial for struggling students from families with a low-SES background (Benner, Boyle, and
Sadler, 2016). Tang (2015), however, found no association between school-based involvement and
children’s achievement for newcomer immigrant families, who tend to have low levels of parental
education. These differences in the value of school-based involvement for children’s achievement
are likely related to the context of the family created by more distal factors such as differences in
408
socioeconomic status. As such, what parents do in the home may matter more than what they do at
school for children’s achievement.
Home-Based Involvement
Home-based involvement encompasses a broad range of education-related activities that families
engage in outside the school (e.g., communicating about school, helping with homework) and
activities not directly related to education but focused on creating an intellectually stimulating learn-
ing environment for the child (e.g., visiting the library, museums, and attending plays). Prior literature
on home-based involvement demonstrates mixed associations with child achievement, which may be
in part due to the wide range of activities that families can engage in to support their child’s educa-
tion that are classified as home-based involvement. Hill and Tyson’s meta-analysis (2009) found that,
when parents created educationally stimulating and supportive home environments, children had
better academic outcomes, whereas parents’ home-based involvement via homework help, in gen-
eral, was associated with more negative child achievement. Research on specific academic-focused
parenting strategies used by parents in the home found that certain types of strategies were associated
with lower achievement whereas some strategies had no relation to children’s achievement (Tang
and Davis-Kean, 2015).
Homework Help
Helping with homework is a common home-based involvement activity which parents engage in
and which demonstrates mixed associations with children’s academic outcomes (Cooper, Lindsay
and Nye, 2000; Epstein, 1988; Levin et al., 1997; Patall, Cooper, and Robinson, 2008; Pomerantz and
Eaton, 2001). One potential explanation relates to the quality of involvement. For example, parents
may be undermining children’s achievement when they engage in home-based involvement activi-
ties because they are teaching skills in a way that may counter how they are taught in the classroom
or in a way that is emotionally frustrating and fosters a negative attitude towards education (Cooper
et al., 2000). Pomerantz and colleagues (2007), for example, found that involvement is most benefi-
cial when it is characterized by positive affect, whereas involvement characterized by negative affect
can be detrimental to children’s outcomes (Pomerantz, Moorman, and Litwack, 2007).
An alternate explanation for the negative association between homework help and students’ aca-
demic outcomes is that parents are responding to children’s poor school outcomes by increasing their
involvement at home. Parent involvement at home, for example, may be prompted by the teacher
reaching out to parents to elicit their help in monitoring the child’s school work and helping with
homework. In some instances, the child may initiate parents’ involvement at home.
Response to Poor Grades

Parents’ academic-focused parenting strategies are another type of home-based involvement that
parents engage in on a consistent basis. In a study using nationally representative data, Tang and
Davis-Kean (2015) examined parents’ endorsement of using different parenting strategies in response
to their child bringing home a report card with grades or progress lower than expected. When
parents endorsed punitive parenting strategies in response to their child’s poor grades (e.g., lecture
or punish the child, limit/reduce the child’s nonschool-related activities), students exhibited lower
levels of literacy and math achievement 5 years later (Tang and Davis-Kean, 2015). The authors
speculated that punitive parenting strategies are likely ineffective in promoting achievement when
the parenting response does not directly address the underlying problem causing academic under-
performance, and thus, it is important for parents to understand the cause of the poor grade prior
409
to using a punitive response. In alignment with research on use of punitive parenting in general
(Gershoff, 2002), low-SES parents in the Tang and Davis-Kean (2015) study were also more likely
to endorse more punitive academic-focused parenting strategies in comparison to their counterparts
with higher levels of SES.
Despite some of the negative associations between certain types of home-based involvement
and student achievement, extant literature supports the importance of fostering families’ educa-
tional involvement at home, particularly for those parents who may not be comfortable engaging
in traditional school-based involvement activities (Moll and Greenberg, 1992; Suárez-Orozco, and
Suárez-Orozco, 1995; Tang, 2015). That is, even if parents have limited schooling themselves, parents’
home-based involvement activities (e.g., providing an enriching home learning environment, hav-
ing high educational expectations) still have benefits for their children’s achievement. In fact, several
aspects of parent involvement in the home have been identified to be associated with positive child
outcomes: alignment between content and skills taught at home and school, provision of develop-
mentally appropriate materials and instruction, and presence of positive affect during family-child
interactions around education (Pomerantz et al., 2007).
Direct and Indirect Mechanisms of Parent Educational Involvement

The theoretical pathways for explaining how parent educational involvement leads to children’s
educational success include both direct and indirect mechanisms (Bandura, Barbaranelli, Caprara and
Pastorelli, 1996; Hoover-Dempsey et al., 2001). Specifically, direct mechanisms are theorized to be
skill-based, and indirect mechanisms are motivation-based (Pomerantz et al., 2007). When parents,
for example, provide homework assistance or cognitive stimulation they can directly transmit aca-
demic skills to their child and model learning practices (Grolnick and Slowiaczek, 1994).
Indirectly, when parents are involved in their children’s education, they convey the message that
school is important, which leads children to value education more highly and foster their motiva-
tion for doing well academically (Grolnick, Ryan, and Deci, 1991). Furthermore, when parents are
involved in their children’s education through attendance at school-based activities (e.g., parent-
teacher conferences) and communication with teachers and/or other students’ parents who are vol-
unteering in the schools, they have opportunities to learn more about their child’s academic strengths
and needs as well as learn more about the culture of school, more generally. In turn, understanding
their child’s academic progress and the culture of schools better allows parents to align home prac-
tices with school practices, and places parents in a better position to intervene to remedy problems
(Domina, 2005).
The strength of these theoretical pathways, however, may vary depending on parents’ educational
attainment. Direct mechanisms, which are skill based, may be particularly contingent on parents’
education. Parents who have experienced success in school and who have completed more years of
education are likely to be better equipped to provide academic support to their children (Magnuson,
2007; Magnuson et al., 2009; Tang et al., 2016). For example, parents with low levels of education
may not be familiar with what are effective study skills and may not be able to scaffold their chil-
dren’s learning strategies. Parent education becomes a more prominent factor for effective parent
educational involvement given the advanced nature of students’ school work as children progress
through school and the knowledge about course selection necessary to prepare students for college
and higher education (Choy, 2001; Crosnoe, 2001).
In comparison, indirect mechanisms are less responsive to parents’ educational attainment. Parents
do not need to have attended formal schooling or have a college degree, for example, to convey that
they value school and have high educational expectations for their children. Even when families are
not directly assisting with homework, children’s motivation increases as long as families maintain
a positive affect; this pattern is particularly true for those children who demonstrate high levels of
410
helplessness (Pomerantz, Wang, and Ng, 2005). Furthermore, there is a robust body of work on immi-
grant families demonstrating that parents of low socioeconomic status are still able to successfully con-
vey the importance of schooling and high educational expectations to their children (Delgado-Gaitan,
2004; Portes and Rumbaut, 2001; Suárez-Orozco, Suárez-Orozco, and Todovora, 2008).
Given the various mechanisms through which family involvement can affect children’s edu-
cational success, subgroups with “at-risk” characteristics for academic under-achievement such as
children from families with low parental education, may benefit uniquely from parents’ educational
involvement (Dearing, Kreider, Simpkins, and Weiss, 2006; Domina, 2005; Tang, Dearing, and Weiss,
2012). In other words, if parent educational involvement promotes academic skills, positive motiva-
tional attitudes towards learning, and family-school synchrony, then children (and families) at greatest
risk for lacking in these areas may benefit the most from involvement. Research findings indicate
that school-based involvement matters more for the literacy achievement of students with the least
educated parents than child ethnicity, maternal education, or family income, for example (Dearing,
McCartney, Weiss, Kreider, and Simpkins, 2004).
Antecedents of Parent Educational Involvement

Despite the benefits of educational involvement for students, not all parents are involved. For
example, in comparison to more highly educated parents, parents with low levels of educational
attainment generally engage in fewer parent involvement activities (Englund et al., 2004; Lareau,
2011). These differences may be related, in part, to between-family differences in ecological context
(e.g., knowledge of what to do, working conditions). That is, families’ investments in their children’s
education may assume different forms based on complex interactions between parents’ priorities
(e.g., whether parents value certain types of involvement activities) and aspects of their ecological
context (e.g., whether parents have flexible work schedules that allow them to volunteer in schools
or whether schools solicit parents’ involvement).
Hoover-Dempsey and Sandler (1997) proposed a theoretical model highlighting three factors
related to why and how families become involved in their children’s education: parents’ perception
of their role in their children’s education, feelings of self-efficacy in helping their children succeed in
school, and perception of opportunities and invitations to become involved. First, parents are more
likely to be involved when they believe it is their duty to be involved in their children’s education.
Second, parents need to believe that they have the ability and/or skills to help their children with
their school work or that they can positively affect children’s academic trajectory. Third, parents are
more likely to be involved when they believe that teachers, schools, and their children want parents
to be involved in children’s education and value their input.
Role Construction
The first factor, role construction, is one barrier that may be important for understanding differences
in involvement between-families because it is highly dependent on parents’ ethno-theories, which
are “cultural models that parents hold regarding children, families, and themselves as parents” (Hark-
ness and Super, 2002, p. 62). Individuals who attended schools that did not elicit parents’ involvement
or whose own parents were uninvolved in their schooling may not believe that it is the parent’s role
to monitor children’s academic progress, but rather the teacher’s, because schooling is in the teacher’s
domain (Huntsinger and Jose, 2009). Further, some parents view asking the teacher questions about
their child’s progress as being disrespectful to the teacher who is considered the expert (Valdés, 1996),
or parents may assume that school personnel like the guidance counselor will know how to help
their child select courses so that they are on track to attend college and so parents take a hands-off
approach. This perspective may be amplified for parents with lower levels of education.
411
Self-Efficacy
The second factor, parents’ feelings of self-efficacy, may be a particularly salient barrier for parents
with low levels of education. Parents high in self-efficacy are more likely to actively engage in their
children’s academic development (Bandura et al., 1996). Parents who have not received much formal
schooling or are unfamiliar with the American school system may hesitate to become involved in
school-based activities because they feel uncomfortable in such a setting and may not know how
to navigate the school system to best advocate for their children (Finders and Lewis, 1994; Turney
and Kao, 2009). Alternatively, parents who dropped out of school prior to graduating may have the
advantage of being familiar with a formal school system, they may be less motivated to be involved if
they faced negative school experiences when they were younger, a concern voiced by parents from
marginalized groups (Lightfoot, 1978; Moles, 1993; Ramirez, 2003). Thus, parents’ self-efficacy is an
important factor that may be a particularly salient barrier to involvement for parents who have low
levels of education or who may not have any experiences with formal schooling. This issue is fur-
ther compounded for parents who have low levels of education and who may not have the English
language proficiency or cultural capital to communicate effectively with their children’s teachers
and school administrators to advocate on their child’s behalf (Delgado-Gaitan, 2004; Peña, 2000;
Ramirez, 2003). Indeed, teachers and schools can promote parent involvement when they facilitate
communication (Henderson and Mapp, 2002). For example, low-SES Mexican American parents
who spoke Spanish increased their school-based parent involvement when their children consistently
had bilingual Spanish-speaking elementary school teachers. In turn, increased parent involvement
predicted children’s higher literacy over time (Tang et al., 2012).
Opportunities
The third factor impacting parent educational involvement relates to whether parents perceive that
there are opportunities for them to become involved in their children’s education and whether
parents feel that their help at home or at school is of value. For example, if parents perceive that
school-based activities are the only way that they can be involved in their children’s activities, they
may feel that there are not enough opportunities for them to become engaged in their children’s
education. Parents with low socioeconomic backgrounds are more likely to experience long work
days, unstable work schedules, and work places that are located far from their homes (Moles, 1993;
Presser, 2004), which makes it difficult for these parents to be highly engaged in their children’s
schooling, particularly with school-based activities. Low-SES parents experiencing financial strain
may not have the financial resources to hire tutors for their children, purchase additional educational
materials, enroll their children in enriching camp programs, or know how to access these resources.
Moreover, the type of involvement that parents engage in is shaped by the value that parents place
on those activities. Some parents, for example, are less likely to volunteer in schools because they do
not view that type of involvement as directly beneficial to their child’s educational success (Hill and
Torres, 2010; Hwa-Froelich and Westby, 2003; Sy and Schulenberg, 2005).
Beyond the practical issue of lack of resources (e.g., time, transportation, financial) that may
impede parents’ involvement (Williams and Sánchez, 2013), relational issues may be another reason
for some parents’ limited engagement in school-based activities (Carreon, Drake, and Barton, 2005).
Low-SES and ethnic-minority parents have reported finding the school climate to be intimidating
and unwelcoming (Ramirez, 2003; Valdés, 1996); for example, parents have reported feeling uncom-
fortable and disrespected due to teachers’ judgmental attitudes and style of communication (i.e., they
often feel that teachers talk down to them; Henderson and Mapp, 2002; Peña, 2000). These types
of negative interactions create an unwelcoming school climate that gives parents little motivation
to participate in activities at school (Trueba, 1998). In turn, limited school involvement hampers
412
parents’ awareness of and access to resources that may benefit their children’s academic development.
Thus, the consequences of school-based involvement may be moderated by the family-school fit,
which may be less than optimal for certain parents, such as parents with low levels of education.
Together, these three factors are important in understanding how and why parents adopt certain
practices in relation to their children’s schooling. Specifically, it is important to understand how these
factors enhance or impede parents’ proclivities to engage in certain types of educational involvement
activities, and how those activities, in turn, are associated with students’ academic outcomes. In addi-
tion, it is important to recognize that each of these three factors—how parents’ construct their roles,
their levels of self-efficacy, and perceived barriers and opportunities to involvement—are themselves
embedded within a larger social context, including parental educational attainment.
Conclusions
This chapter highlights the important role that education plays in promoting positive parenting
beliefs and behaviors that then influence of the positive development of children. In many studies
of children’s behavior, parental educational attainment is used as a covariate to remove variance that
might be associated with the child variable of interest. This type of analysis loses the important role
that parental educational attainment may play in parenting beliefs and behaviors that lead to the child
outcome. Much of the research to date examining socioeconomic status has examined the role of
income in promoting positive parenting. Indeed, the research that has examined the negative influ-
ence of income deprivation made a strong contribution toward understanding factors that influence
parent mental health and subsequent behaviors (McLoyd, 1998), but less research has focused on the
possibly stronger and more persistent influence of low education. This chapter reviewed some of the
avenues by which education may exert influence on family and child well-being through behaviors
in the home and toward involvement in schooling, but there are more avenues to be tested and
explained. Future research should continue to examine the pathways and the unique contributions
that being educated afford families both in their ability to use often very limited resources wisely, and
as a potential agent for helping families function better as a system.
There are challenges ahead in considering various way to measure educational attainment but also
the unique role it may play above genetic contributions that parents transfer to their children (Noble
et al., 2015). Continuing to employ rigorous methods (e.g., quasi-experimental and experimental
techniques), as used by Magnuson et al. (2009) and Gennetian et al. (2008), will help to address these
issues and account for potential confounding influences which may aid the design of effective inter-
ventions. Indeed, relative to other countries, education is an easily attained commodity in the United
States, and a very important one to consider investing in for the future. Understanding how educa-
tion may help with the success of family functioning, health, mental health, and general well-being
can help with the design of future interventions that need to target the distribution of resources and
ability of recipients to understand and use these resources for the benefit of themselves and others.
Acknowledgments
The first author thanks Katherine Magnuson for her many conversations and observations that led to
thinking about parent educational attainment and child development over the years.
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13
SOCIOECONOMIC STATUS
AND PARENTING
Introduction
Socioeconomic status (SES) is a pervasive predictor of child development; parenting is a large part of
the reason why. Across different populations and different developmental domains, there is evidence
that parenting mediates the relation between SES and child outcomes (Bornstein and Bradley, 2003;
Fernald, Kariger, Hidrobo, and Gertler, 2012; Kağıtçıbaşı, 2007; Paxson and Schady, 2007) and is part
of the process by which social inequality is transmitted across generations (Conger and Donnellan,
2007; World Bank, 2015). The aim of this chapter is to elucidate one link in this causal chain from
SES to child outcomes—the link between SES and parenting. The chapter begins with a historical
introduction to research on SES and parenting, followed by a discussion of definitions of SES and
approaches to its measurement. The third section summarizes the literature on differences in parent-
ing cognitions and parenting practices associated with SES. The fourth and fifth sections ask how
SES-related differences in parenting come about. The fourth section traces pathways of influence
from SES to parenting; the fifth unpacks SES and identifies the separable effects of its constitu-
ents, income, education, and occupation. The penultimate section discusses remaining questions and
future research directions in the study of SES and parenting, followed by a brief conclusions section.
Historical Considerations and Classical Research

in the Study of SES and Parenting
The history of research on SES and parenting has been shaped by the waxing and waning of con-
cern for the welfare of low-SES children, by social and political attitudes toward admitting and
discussing social inequality, by shifting foci of research attention within developmental science, and
by developing interest among economists in children as national resources and in parenting as the
means by which that resource is developed. National concern for child welfare in the United States
was the impetus for a 1936 White House conference on child health and protection; the resulting
report described SES-related differences in aspects of parenting as diverse as the likelihood that
parents served their children spinach and read books on child care (Anderson, 1936). For the next
two decades, the child health and protection motive remained a minor theme in research on SES
and parenting. In the 1960s, a heightened awareness of social inequalities across North America and
Western Europe provided impetus for new research on disadvantaged children. In the United States,
the War on Poverty was launched to eradicate some of the worst consequences of low-income living
421
conditions and was accompanied by an infusion of funds for research aimed at better understanding
the developmental sequelae of SES.
By the end of the 1970s, research on SES, including SES and parenting, was on the wane. There
was a general unease with the very notion of social stratification. Findings of SES-related differences
in parenting and the attendant interpretation that lower-SES parents were to blame for their chil-
dren’s difficulties were controversial and unpopular (Ensminger and Fothergill, 2003). The topic was
additionally sensitive because SES and ethnicity are confounded, and thus negative descriptions of
low-SES parenting were sometimes negative descriptions of minority groups. Independently, scien-
tific interest turned from seeking explanations of variation in development to seeking explanations
of universal aspects of development. Research on normative growth and developmental trajectories
supplanted research on individual and group differences.
In the 1990s, the child welfare motive regained currency among research scientists (Garcia Coll
et al., 1996; Collins, Maccoby, Steinberg, Hetherington, and Bornstein, 2000). The view that impli-
cated lower-SES parents in their children’s difficulties gave way to a view that included forces outside
the family (e.g., poverty, prejudice, schools, neighborhoods) as sources of SES-related differences in
child development. SES came to enjoy legitimacy as a contextual variable in studies of both parent-
ing and child well-being (DeGarmo, Forgatch, and Martinez, 1999; Glasgow, Dornbusch, Troyer, and
Steinberg, 1997; Gutman and Eccles, 1999).
Another strand in the early research on SES and parenting, which was dominant during the
1940s and 1950s, consisted of studies that sought to document the differences associated with SES
in a manner similar to anthropologists’ descriptions of cultural differences (e.g., Mead, 1928, 1930;
Whiting and Child, 1953). In Middletown, Lynd and Lynd (1929, 1937) carefully documented social
stratification in a typical U.S. town and concluded that one aspect of life that varied with social status
was childrearing. Davis and Havighurst (1946) described differences in feeding, weaning, and toilet
training practices associated with class and ethnicity. Ethnographic approaches to studying parenting
in different social strata have more recent, influential examples in the work of Brice Heath (1983)
and Lareau (2011). Heath (1983) documented differences in the communicative styles of parents in
three communities: a poor, rural African American community, a working-class European Ameri-
can community, and a mainstream community, all in the Piedmont region of the southern United
States Lareau (2011) documented differences in the organization of daily life, in language use, and in
interactions with social institutions among families from three social strata—middle class, working
class, and poor, in the U.S. Midwest and Northeast. These ethnographies not only documented SES-
related differences in parenting, but they also sought to describe the contexts that shape parenting
differences and to explain the dislocation that children from lower-SES backgrounds experienced
when they entered mainstream schools.
Shifts in the scientific focus of developmental science also have caused changes in the focus of
research on SES and parenting. Early work in developmental science focused on parents as agents of
socialization and on aspects of parenting thought to influence social and personality development. In
the 1930s and 1940s, weaning and toilet training were of great concern, as were aggression and sex
play, the latter of which often fell under the heading of “impulse control” (Bronfenbrenner, 1958).
Through the 1950s, “responsibility training,” or the demands for independence that parents placed
on children, was the focus of attention. These aspects of parenting were considered important to
socialization, and it is clear that socialization was of interest because it was thought to hold the key
to explaining SES-associated differences in children’s achievement. For example, Davis and Havi-
ghurst (1946, p. 707) concluded that “middle-class children are subjected earlier and more consist-
ently to the influences which make a child an orderly, conscientious, responsible, and tame person.”
Scientific trends shifted in the 1960s, and interest in specific behaviors, such as weaning and toilet
training, gave way to interest in more global dimensions of parenting, in particular parenting style.
Also in the 1960s, the federal War on Poverty, and its associated funding, resulted in a new research
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Socioeconomic Status and Parenting
focus on how experiences with parents prepared children for schooling (Slaughter-Defoe, 1995).
The HOME inventory was developed to assess aspects of household environments thought to be
related to cognitive outcomes, and it has subsequently been used in a great deal of research examin-
ing SES-related differences in children’s experiences (Caldwell and Bradley, 1984; Gottfried, 1985).
SES-associated differences in mother-child interaction (and their consequences for cognitive devel-
opment), along with differences in the strategies mothers used in teaching and conversing with their
children, received the greatest attention (Williams, 1970).
Work by economists has gained prominence in the field of SES and parenting (Foster, 2002). To
economists, skills and knowledge are human capital. Economists’ research on SES and parenting inves-
tigates how parents across the range of socioeconomic strata invest their resources of time, effort, and
money in the development of the human capital of their children (Becker and Tomes, 1986; Foster,
2002; Kalil, Ryan, and Corey, 2012; Lugo-Gil and Tamis-LeMonda, 2008; Paxson and Schady, 2007).
The psychologists’ study of parenting as an influence on child development is recast in the economists’
approach as the study of parenting as a means of human capital formation (Heckman, 2006).
Most recently, a cognitive neuroscience framework has influenced research on SES and parenting.
That research has focused on language, memory, and executive function because these skills appear
to have distinct underlying neurocognitive systems. The research aims to link SES both to children’s
skills and to the neural structures that support those skills ( Johnson, Riis, and Noble, 2016; Noble
et al., 2015). Findings from neuroscience have directed attention to parenting as a source of SES-
related differences because other influences, such as schools, begin too late. The correlates of SES in
brain structure are observable in infancy, and early appearing differences in skill may not easily be
ameliorated later in childhood (Nelson and Sheridan, 2011; Noble et al., 2015).
In summary, multiple sources of influence, from science and from politics, have combined to
shape the history of research on SES and parenting.
Definitions and Measures of SES

The foregoing historical introduction ignored questions of the definition and measurement of SES.
But inadequate attention to measurement is a criticism that has been leveled at much of the research
in this field (Bornstein et al., 2003; Duncan and Magnuson, 2003), and measurement questions are
integral to the research goals of identifying the factors with causal impacts and describing processes
by which those factors affect parenting.
The traditional definition of SES is that it is an individual’s or household’s relative position in a
social hierarchy based on access to, or control over, wealth, prestige, and power (Mueller and Parcel,
1981; Willms, 2003). Other definitions have been proposed, however, and there is not a current
consensus on the best definition or measure of SES (Oakes and Rossi, 2003, Bornstein and Bradley,
2003). One conceptualization that has influenced work on child development comes from Coleman
(1988), who identified financial capital, human capital, and social capital as the components of SES.
Financial capital is the source of material resources—food, clothing, housing, and everything else
that depends on money; a typical index of financial capital is income. Human capital is the source
of nonmaterial resources, such as knowledge and skills, a frequent index is parent education. Social
capital is a less transparent concept; it is an individual’s or household’s connections to a larger social
group, in the form of access to the expectations of that group, the norms of that group, and the
channels of information that group provides (Coleman, 1988). Occupational status may index social
capital (Conger and Donnellan, 2007; Entwisle and Astone, 1994), although social capital has also
been argued to reside in schools and neighborhoods as well (Caughy and O’Campo, 2006).
Operationalizing SES has proven challenging. One approach is to create a composite measure that com-
bines the multiple variables that define SES. The best known and most widely used composite measures of
SES in the United States are the Hollingshead Four-Factor Index of Social Status and the Socioeconomic
423
Index of Occupations (SEI; Bornstein et al., 2003). The Hollingshead is based on the education and
occupation of each employed householder in the home; the SEI is a measure of occupational prestige that
makes use of data on the educational requirements and income associated with occupations (see Bornstein
et al., 2003, for a more complete discussion). Other composite indices also exist (e.g., Willms, 1999).
Another approach to measuring SES is to combine traditional indicators with measures of other,
often correlated factors, to create an index of cumulative risk (Burchinal, Roberts, Hooper, and Zei-
sel, 2000; Mistry, Benner, Biesanz, Clark, and Howes, 2010; Sameroff, Seifer, Baldwin, and Baldwin,
1993; Wachs, Cueto, and Yao, 2016). The cumulative risk index used in the Family Life Project
(a longitudinal study of over 1,000 children in poor rural communities) combines the defining
indictors of SES, education, income, and occupational prestige with work hours, household density,
neighborhood safety, and continuity in interparental relationships (Vernon-Feagans and Cox, 2013).
Other composite indices include factors such as poor nutrition, exposure to environmental toxins,
and maternal depression (Wachs et al., 2016).
A final approach to characterizing the diverse experiences associated with different levels of SES is
to use a person-centered statistical approach to identify clusters of individuals (i.e., types) who share
similar SES-related characteristics and who differ meaningfully from each other (see Laursen and
Hoff, 2006). Of course, the types that emerge from such an analysis depend on the sample studied
and the characteristics entered into analysis. When latent class analysis was applied to the Family Life
Project sample, described in terms of 10 measures of family structure, income, and other risk factors,
six distinct groups were identified, including Poor and Married, Poor and Unmarried, Poor, Unmar-
ried and No Partner, for example (Rhoades, Greenberg, Lanza, and Blair, 2011).
Arguments have been made against the use of any composite indicator of SES. It has been argued
that the effects of SES can be accounted for in the separate effects of its components and thus SES “is
not more than the sum of its constituent parts” (Bornstein et al., 2003, p. 66). It has also been argued
that composite indices may obscure the source of some SES effects, hiding unique pathways of influ-
ence from different sources (Conger and Donellan, 2007; Duncan and Magnuson, 2003; Entwisle
and Astone, 1994). Finally, it has been argued that the effects of single components can be studied in
experimental or quasi-experimental designs, whereas effects of a global measure cannot, and policies
can be implemented to target specific components whereas “there are no treatments for enhancing
overall SES” (Duncan and Magnuson, 2012, p. 377).
Often, researchers select a single component to serve as a measure of SES. Where financial capital
is the variable of interest, the measure is typically income or income-to-needs ratio. Where human
capital is the variable of interest, the measure is typically maternal education (Ensminger and Fother-
gill, 2003). There are advantages to maternal education as a measure. Participants may be more will-
ing to report their level of education than their income, and their reports may be more accurate.
Also, depending on how the research is conducted, inquiries about income may adversely affect the
relationship between the researcher and the participant (Hoffman, 2003). Maternal education is more
stable than occupation or income (Duncan and Magnuson, 2003). Finally, maternal education is the
most robust predictor of parenting among SES components (Bornstein et al., 2003).
Use of the term SES, however it is measured, implies a continuous dimension. Another currently
used term, inequalities, similarly entails the notion of a single scale on which households differ. The
more old fashioned term, social class, entails a different conceptualization. As the sociologist Kohn
put it in his work on social class and parent-child relationships,
social class has proved to be so useful a concept because it refers to more than simply
educational level, or occupation, or any of the large number of correlated variables. It is so
useful because it captures the reality that the intricate interplay of all these variables creates
different basic conditions of life at different levels of the social order.
(Kohn, 1963, p. 471)
424
That argument has not prevailed. Research in psychology tends to use the term SES and one or more sin-
gle indicators. Research by economists tends to use the term inequalities and to use income as the indica-
tor. Sociologists sometimes use the term social class and to use multiple indicators—at least descriptively;
but the utility of the concept of social class is debated even within the field (Lareau and Conley, 2008).
In summary, SES is a complex, multifaceted variable that scholars have conceptualized and meas-
ured in different ways at different times. There is no single agreed upon best measure of SES. Rather,
the explanatory power of SES as a composite and of the constituent components of SES with respect
to different domains of parenting are part of what scholars study when they study SES and parenting.
Parenting Differences Associated With SES

Parenting encompasses cognitions, styles, and practices. Parenting cognitions include expectations,
values and goals, and the beliefs, attitudes, and knowledge about child development that parents have.
Parenting style consists of the attitudes about children that parents communicate to their children
and the emotional climate in which these attitudes are expressed. Parenting practices are the behav-
iors parents produce in interactions with their children, the home environments parents create for
children, and the connections to the world outside the home that parents both enable and permit.
There are SES-related differences in all domains.
Parenting Cognitions
Expectations
Parents from different socioeconomic levels expect different developmental timetables. In general,
and across cultures, higher-SES mothers give earlier age estimates for children’s attainment of devel-
opmental milestones and higher estimates of young children’s capacities than lower-SES mothers
(Hess, Kashiwagi, Azuma, Price, and Dickson, 1980; Mansbach and Greenbaum, 1999; Ninio, 1988;
von der Lippe, 1999). SES-related differences of this sort most reliably appear with respect to time-
tables of language development and estimates of cognitive capacities. In contrast, lower-SES mothers
have been found to have higher expectations with respect to when children will be toilet trained and
the politeness behavior they should show (Tardif, Au, Wellman, and Nakamura, 2000).
Values and Goals

In general, and across cultures, lower-SES parents emphasize proper behavior, which includes being
obedient, respectful, and quiet, as goals for their children. Higher-SES parents want their children
to be self-directed (Alwin, 1984; Harwood, 1992; Harwood, Miller, and Lucca Irizarry, 1995; Kohn,
1979; Luster, Rhoades, and Haas, 1989; Pearlin and Kohn, 1966; Tudge, Hogan, Snezhkova, Kula-
kova, and Etz, 2000; Wright and Wright, 1976).
Beliefs, Attitudes, and Knowledge

Lower-SES parents feel they have less control over child developmental outcomes than do higher-
SES parents (Elder, Eccles, Ardelt, and Lord, 1995; Luster and Kain, 1987). In her ethnography of
U.S. childrearing, Lareau (2011) described the characteristic middle-class attitude toward parenting
as one that assumes parents have the power to influence child outcomes. She terms the approach
concerted cultivation. Middle-class parents organize how their children will spend their time out of
school, with lessons and organized sports. Middle-class parents intervene in their children’s school
experiences as well, in an effort to create an environment that will cultivate their children’s talents. In
contrast, blue collar parents see their role as allowing the natural accomplishment of growth. They do not
425
assume responsibility for the planning of their children’s leisure time, nor do they seek to influence
their children’s school experience. In her ethnographic study of communicative style, Heath (1983)
found similar differences with respect to children’s language development. In contrast to middle-class
mothers who actively label objects and explain the world to their children, adults in the lower-SES,
African American group do not, believing that children have to learn such things for themselves.
A clear finding from developmental science is that early experience matters. Thus, SES-related
differences in beliefs about the malleability of child outcomes—and perhaps also in expectations
for developmental timetables—reflect an awareness of current knowledge about child development.
Parents from higher-SES backgrounds show greater knowledge of child development and recom-
mended parenting practices, and they seek to acquire and update that knowledge more readily than
do parents from lower-SES backgrounds (Bornstein, Cote, Haynes, Hahn, and Park, 2010; Rowe,
2008; Rowe, Denmark, Harden, and Stapleton, 2016; Stevens, 1984).
Parenting Styles
Different parenting styles create different emotional climates in the home (Darling and Steinberg,
1993). Scholars have long argued that parenting style varies as a function of SES, and considerable
evidence supports this assertion. As early as 1958, Bronfenbrenner concluded that “parent-child
relationships in the middle-class are consistently reported as more acceptant and equalitarian, while
those in the working-class are oriented toward maintaining order and obedience” (p. 420). Parenting
styles in higher-SES homes have been described as democratic (Hoffman, 1963) and child-centered
(Sears, Maccoby, and Levin, 1957), in contrast to the authoritarian and parent-centered style that
characterizes lower-SES homes.
Research using Baumrind’s (1967) typology of parenting styles has found that the prevalence of
authoritative, authoritarian, and permissive styles varies as a function of social strata. Authoritative
parenting involves high levels of control over children combined with high levels of responsiveness
to children. Authoritative parents reason with their children and encourage their children’s develop-
ment of autonomy. Authoritarian parenting involves high levels of parent control over children and
low levels of responsiveness to the children. Authoritarian parents require obedience, use punish-
ment, and may display little warmth. Permissive parenting involves little control and few require-
ments for mature behavior (Holden, 2010). Across cultures and ethnicities, families with higher
parental education tended to be lower in authoritarian and permissive parenting and higher on
authoritative parenting than families with lower parental education (Bluestone and Tamis-LeMonda,
1999; Chen, Dong, and Zhou, 1997; Dornbusch, Ritter, Leiderman, Roberts, and Fraleigh, 1987;
von der Lippe, 1999). Fewer studies of SES-related differences in parenting style have used Maccoby
and Martin’s (1983) expanded typology of parenting styles, in which permissive parenting is termed
indulgent, and a fourth category of neglectful or uninvolved is added (Holden, 2010). The research
that has been conducted suggests that the prevalence of those categories also varies as a function
of SES. Reports from several thousand U.S. adolescents indicate that parent education is positively
correlated with authoritative parenting and negatively correlated with authoritarian parenting, while
also revealing that parent education is positively correlated with indulgent parenting and negatively
correlated with neglectful parenting (Glasgow, Dornbusch, Troyer, and Steinberg, 1997).
Parenting Practices
Time Spent With Children

More educated and affluent parents spend more time with their children, even though they also
spend more time at work. Results from the 2003–2007 American Time Use Survey, in which over
426
6,000 mothers reported on the time they spend in different activities (Guryan, Hurst, Kearney, 2008;
Kalil et al., 2012), also indicated that, as children get older, more educated mothers are more apt to
change how they spend their time with children. Thus, the differences between more and less edu-
cated mothers in basic care and play time are greater when the children are infants and toddlers than
they are later, whereas the difference in time spent in management activities is greater when children
are school age (Kalil et al., 2012).
Warmth and Sensitivity

Higher-SES parents are more sensitive, demonstrate higher levels of positive regard, and are less
intrusive in interaction with their children than lower-SES parents (Raviv, Kessenich, and Morrison,
2004; Tamis-LeMonda, Shannon, Cabrera, and Lamb, 2004; Vernon-Feagans et al., 2008). That is,
the higher-SES parents more frequently take the child’s perspective, accurately perceive the child’s
signals, and promptly and appropriately respond to these signals, and they more frequently demon-
strate love, respect, and admiration for the child and they are not overcontrolling. Children in higher-
SES homes hear more affirmations (i.e., encouragements) from others and few prohibitions (Hart
and Risley, 1995). Higher-SES mothers show more warmth and sensitivity in interaction with their
children than lower-SES mothers (Burchinal, Vernon-Feagans, Cox, and Key Family Life Project
Investigators, 2008; Mistry et al., 2010).
Direct Control Practices

In interacting with young children, lower-SES mothers have consistently been found to be more
controlling, restrictive, and disapproving than higher-SES mothers. Across cultures, lower-SES moth-
ers devote proportionately more talk to directing child behavior than do higher-SES mothers (Hart
and Risley, 1992, 1995; Hoff-Ginsberg, 1991, 1998; Reger, 1990). Lower-SES mothers grant their
children less autonomy, and they are more restrictive, punitive, and intrusive than higher-SES moth-
ers (Bayley and Schaefer, 1960; Caldwell and Bradley, 1984. These findings extend to the use of
physical forms of discipline: Lower-SES parents practice corporal punishment to a greater degree
than higher-SES parents (Clarke-Stewart, VanderStoep, and Killian, 1979; Lareau, 2011; Straus and
Stewart, 1999). During middle childhood, harsh punishment is more frequently found in the lower
socioeconomic strata than in the upper strata (Bronfenbrenner, 1970; Simons, Whitbeck, Conger,
and Wu, 1991; Straus, Gelles, and Steinmetz, 1980). Financial strain and economic hardship exacer-
bate these trends (Gutman and Eccles, 1999).
Managerial Control
For parents of infants and preschool children, education and, to a lesser degree, paternal occupation,
are positively related to the provision of appropriate play materials, the variety of daily stimulation,
and the organization of the environment (e.g., regular outings and trips to the doctor and to the
grocery store; Caldwell and Bradley, 1984). Families with higher incomes are more likely to send
to their children to early care and education programs (Magnuson, Meyers, and Waldfogel, 2007).
As children get older, indices of SES predict how children spend their time in and out of home.
Maternal education was positively related to the amount of time 9-year-olds spent in skill activities,
such as homework and reading for fun, and negatively related to the amount of time spent watch-
ing television (DeGarmo et al., 1999). Higher-SES parents also may coordinate participation in the
larger community through activities such as clubs, private lessons, sports teams, and church-related
activities (Kalil et al., 2012; Lareau, 2011). SES, not employment, predicts the degree to which moth-
ers volunteer in these out-of-home activities (O’Donnell and Stueve, 1983).
427
Language and Communication

A substantial body of evidence indicates that both the amount and nature of communicative interac-
tion between parents and children differ as a function of SES (Hoff, 2003a, 2003b, 2006; Schwab and
Lew-Williams, 2016). The differences begin early. Higher-SES mothers use more communicative
gestures with their 14-month-old children than lower-SES mothers (Rowe and Goldin-Meadow,
2009). Studies of mothers’ interactions with their children spanning the age range from 2 years to
adolescence have found that higher-SES mothers address more speech to their children than lower-
SES mothers (Brody, 1968; Dunn, Wooding, and Herman, 1977; Field and Pawlby, 1980; Feiring and
Lewis, 1981; Greenberg and Formanek, 1974; Heath, 1983; Hess and Shipman, 1965; Hoff-Ginsberg,
1992, 1998; Jacob, 1974; Ninio, 1980; Rowe, 2008, 2012; Schachter, 1979; Tulkin and Kagan, 1972,
and see Hoff, 2006, for a summary). Relatedly, the responsivity subscale of the HOME inventory,
which is heavily loaded with indices of verbal responsivity, is positively related to parental education,
and, less robustly, to occupation (Caldwell and Bradley, 1984; Raviv et al., 2004). SES-related differ-
ences emerge on the language and literacy subscale of the HOME inventory as well (Mistry et al.,
2010). The most dramatic difference in the quantity of talk addressed to children was documented by
Hart and Risley (1995). Extrapolating from measures of the number of words addressed to children
in hour-long recordings made on a monthly basis from child age 9 months to 36 months, Hart and
Risley estimated that children of high-SES parents have been exposed to 30 million more words than
children of low-SES parents by the age of 4 years.
With respect to the content of verbal interaction, higher-SES children who hear more speech
also hear speech that contains a greater variety of words (Hoff, 2003b; Rowe, Pan, and Ayoub, 2005),
more frequent rare words (Rowe, 2012), and is more syntactically complex (Hoff, 2003b; Hutten-
locher, Vasilyeva, Waterfall, Vevea, and Hedges, 2007). The child-directed speech of higher-SES
mothers contains a larger proportion of conversation-eliciting questions (Hoff-Ginsberg, 1991) and
a smaller proportion of directives than the child-directed speech of lower-SES mothers (Hart and
Risley, 1995; Hoff-Ginsberg, 1991; Rowe, 2008). Both the quantity and quality of speech have posi-
tive relations to language development (Bornstein, Haynes, and Painter, 1998; Huttenlocher, Haight,
Bryk, Selzer, and Lyons, 1991; Huttenlocher, Waterfall, Vasilyeva, Vevea, and Hedges, 2010), which
suggests that higher-SES children are provided a better data base from which to learn language than
lower-SES children.
More educated mothers also provide more explicit information when they talk to their children
than less educated mothers. Findings from several different ethnic groups reveal that more educated
mothers engage in more explicit teaching of object labels than less educated mothers (Brophy, 1970;
Hammer and Weiss, 1999; Lawrence and Shipley, 1996; Ninio, 1980; Reger, 1990). In addition,
middle-class mothers also have been found to talk about causality with their children more than
working-class mothers (Brophy, 1970; Sigel, McGillicuddy-DeLisi, and Johnson, 1980). Higher-SES
mothers also provide more cognitive stimulation than lower-SES mothers, actively teaching and
attempting to expand the child’s knowledge and abilities (Raviv et al., 2004; Tamis-LeMonda et al.,
2004; Vernon-Feagans et al., 2008). Higher-SES mothers talk to their children about things and
events that are not in the immediate context more than do lower-SES mothers; the talk of lower-SES
mothers to their children is more about the here-and-now (Rowe, 2012).
In general, there are more SES-associated differences on language measures than on nonverbal
measures of how parents interact with their children. For example, college-educated mothers who
talk more to their children than high-school-educated mothers do not spend more time in joint
(verbal and nonverbal) attention with their children (Hoff-Ginsberg, 1991). More educated African
American mothers who talk more and engage in more verbal games with their children do not differ
from less educated African American mothers in nonverbal measures of involvement in play (Ham-
mer and Weiss, 1999). A comparison of caregiving across 28 low-income countries found national
428
differences associated with national economic indicators and found that cognitive caregiving prac-
tices (e.g., book reading, counting) varied more than socioemotional caregiving practices (e.g., sing-
ing songs, playing; Bornstein and Putnick, 2012).
Literacy Practices
Compared to higher-SES families, low-income and less educated families are less likely to have
children’s books in the home (Raikes et al., 2006), and they are less likely to read to their children
on a regular basis (Bradley, Corwyn, McAdoo, and Garcia Coll, 2001; Raikes et al., 2006; Whitehurst
et al., 1994). This difference is not only a difference in children’s exposure to books. Differences in
the amount of book reading create differences in the oral language children hear because adults’
child-directed speech during book reading contains a richer vocabulary and more complex syntax
than speech in other contexts (Hoff-Ginsberg, 1991; Snow et al., 1976).
Parenting Cognitions as a Source of Influence on Parenting Practices

Although parenting cognitions and parenting behaviors can both be thought of as parenting outcomes
influenced by SES, only the parenting behaviors are proximal variables shaping child experiences and
outcomes. Thus, part of the task of tracing paths of influence from SES to parenting involves asking
whether cognitions are part of what links SES to parenting behavior. There is evidence that they are.
The attitude of concerted cultivation that Lareau (2011) described among middle-class parents
makes sense only if middle-class parents believe that their efforts will have effects. As Lareau describes
them, middle-class parents sign their children up for sports to foster their children’s athletic ability.
They talk to their children in ways that will advance their children’s knowledge and ability to express
themselves. They intervene with the schools, camps, and other institutions their children may attend
to secure the best circumstances for promoting their children’s development. Concerted cultivation
actually rests on two beliefs: a belief that child outcomes can be influenced and a belief that they, the
parents, have capacity to exert that influence.
The belief that outcomes can be influenced, termed mindset in much of the research on the topic,
shows clear relations to parent behavior. In survey data, parents who report believing that children’s
abilities are malleable also report spending more time on math and reading activities with their
children compared to parents who believe children’s abilities are fixed (Muenks, Miele, Ramani,
Stapleton, and Rowe, 2015). An experimental manipulation of this mindset has also shown effects
on parents’ constructive involvement with their children on a laboratory based task (Moorman and
Pomerantz, 2010). A longitudinal study of the relation of mothers’ parenting cognitions and practices
found that the degree to which the mothers attributed success in parenting tasks to their own ability,
effort, or mood in questionnaire responses when their children were 20 months old was significantly
related to direct assessments of how supportive their parenting behaviors were when their children
were 4 years old (Bornstein, Putnick, and Suwalsky, 2017). There is little research on SES-related
differences in mindset.
The belief that one can have influence as a parent is the self-efficacy belief that earlier research has
found to differ as a function of SES. In one study, lower-SES parents were less likely than higher-SES
parents to believe they have influence over their children’s outcomes, and were therefore less likely to
endorse positive outcomes as childrearing goals and less likely to engage in “competence promoting
parenting” (Brody, Flor, and Gibson, 1999). Another example is the finding that childrearing beliefs
are more closely linked to childrearing behaviors among middle-class mothers than among working-
class mothers (Tulkin and Cohler, 1973).
Other beliefs may also help explain SES-related differences in parenting practices. Some have
suggested that the high level of verbal activity that characterizes middle-class Western mothers’
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interactions reflects the belief that babies are “separate individuals with whom communication is
possible and necessary” (Snow, De Blauw, and Van Roosmalen, 1979, p. 270). SES-related differences
in mothers’ warmth and involvement may reflect the greater value lower-SES mothers place on
conformity and the greater value higher-SES mothers place on self-direction as an outcome (Luster
et al., 1989). Knowledge of child development constitutes another set of beliefs that can influence
parenting. Parents who endorse the view that infants are “cognitive beings” provide more cognitively
enriching activities (Ninio, 1979). In one study, parents with overall higher scores on a standardized
test of knowledge of child development talked more to their children, showed more word diversity,
and talked to their children in longer, more complex utterances than parents with lower scores, and
knowledge of child development accounted for all of the SES-related differences in these properties
of talk to children (Rowe, 2008).
In summary, there are SES-related differences in the beliefs and attitudes parents hold about par-
enting and in a wide range of parenting practices. Studies of parenting differences related to SES in
the United States and studies of parenting differences related to national income across developing
countries both find that verbal interaction and cognition-related aspects of parenting vary more as
a function of SES or national income than do the nonverbal interaction and socioemotional com-
ponents of parenting (e.g., nonverbal interaction, play, singing songs; Bornstein and Putnick, 2012;
Hammer and Weiss, 1999; Hoff-Ginsberg, 1991).
Pathways of Influence in Relations Between SES and Parenting

This section continues the effort to trace SES-related differences in parenting back to their sources
by laying out, in data analytic terms, models of the different pathways through which SES might
influence parenting. Examples will be provided (see also Hoff, Laursen, and Bridges, 2012). Most of
the models, applied here to the relation between SES and parenting, are offshoots of theories whose
goal is to explain how SES is linked to child development outcomes. They include SES in the role
of predictor of parenting and in the roles of moderator, mediator, and spurious correlate of relations
between SES and parenting.
Models in Which SES Is a Predictor and Cause of

Associated Differences in Parenting
Most of the literature on SES and parenting can be subsumed under a model in which SES is the
predictor and parenting the outcome. Such models are termed social causation models. They come in
two forms: direct and indirect effects models.
Direct effects models of the relation between SES and parenting. In direct effects models, SES is causally
and directly linked to parent attitudes and behaviors. The extended investment model is an example of
a direct effect model. It posits that increases in resources translate into increases in parents’ ability to
provide material goods, information, and experiences that promote child development (Conger and
Dogan, 2007). The function that relates SES to its consequences can be linear or nonlinear. In linear
direct effects models, increasing levels of SES are hypothesized to be associated with constant, propor-
tional, corresponding changes in parent attitudes and behaviors, and these effects are hypothesized
to operate across the entire range of the variable that indexes SES. In nonlinear direct effects models,
the effects of SES are not assumed to be constant across its range. Change in the dependent vari-
able (parenting) either accelerates or decelerates as levels of the independent variable (SES) change
incrementally. Some direct effects models treat SES as a continuous variable and others treat SES as
a categorical variable. For example, income or income-to-needs ratio are measured as continuous
variables and have been treated that way in analyses as well (Davis-Kean, 2005; Raviv et al., 2004).
These measures have also been dichotomized using a threshold to divide the sample into those in
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poverty and those not in poverty (Bradley et al., 2001) or into those with and without an income-
to-needs ratio greater than 1.5 (Rhoades et al., 2011). Similarly, education can measured and entered
into analyses as a continuous variable (i.e., years of education, Davis-Kean, 2005, Raviv et al., 2004;
LeVine, LeVine, Schnell-Anzola, Rowe, and Dexter, 2012), and it has also been dichotomized, divid-
ing the sample into those with and without a high school diploma (Rhoades et al., 2011) or into
those with and without at least a 4-year degree (Hoff, Burridge, Ribot, and Giguere, 2018).
Indirect Effects Models of the Relation Between SES and Parenting

In a mediated or indirect effects model, SES or its constituents create other conditions that, in turn,
affect parenting. Models in this category include family stress models (Conger and Elder, 1994) which
describe the debilitating effects of financial challenges on the well-being of parents—as individu-
als and as a couple—that eventually take a toll on parenting. Originating in research indicating that
economic hardship interferes with family functioning (Angell, 1936), the model posits that economic
pressure on parents adversely effects the marital relationship and parent mental health, which, in
turn, interfere with parenting. The environmental chaos model, another indirect effect model, posits
pathways from low-SES conditions to household disarray, which limit opportunities for positive par-
enting (Wachs and Corapci, 2003). These models both assume nonlinear threshold effects—income
levels or economic changes are harmful to parenting only when they trigger financial hardship.
Curvilinear models of indirect effects, another nonlinear model, posit that both low-income and
high-income are associated with stressors that, in different ways, diminish the capacity for positive
parenting (Luthar and Latendresse, 2005). The effects at the high-SES end of the continuum arise
because successful parents may be too busy to emotionally and physically engage with children and
because the achievements of these parents may foster unrealistically high expectations for their chil-
dren’s achievement.
Evidence of Causal, Predictive Relations Between SES and Parenting
Evidence of Direct Effects of SES on Parenting

A clear example of direct effects of SES on parenting is the effect of income on parents’ ability
to provide a stimulating environment for their children (Yeung, Linver, and Brooks-Gunn, 2002).
Analyses of two national data bases, the Consumer Expenditure Survey and the Early Childhood
Longitudinal Study, Kindergarten Cohort, indicate that families with higher incomes spend more
money on enrichment items (toys, books, sports equipment) and enrichment activities (music les-
sons, sports classes, tutoring) than low-income families (Kaushal, Magnuson, and Waldfogel, 2011).
Direct effects of education have been proposed in which the language skills and style of language
use mothers acquire in the classroom are carried over into their interactions with their children
(Hoff et al., in press; Hoff-Ginsberg, 1991; LeVine, LeVine, and Schnell, 2001; LeVine, LeVine,
Schnell-Anzola, Rowe, and Dexter, 2012). Education-related verbal skills also directly influence
parents’ ability and confidence to interact with their children’s teachers and schools (Lareau, 2011).
Direct negative effects of low levels of education are evident when low-SES parents report that their
own limited academic skills keep them from helping with their children’s homework (Lareau, 2011).
Direct effects of high levels of parental education are abundantly evident in the projects displayed at
middle school science fairs.
Direct effects of SES are not necessarily linear. The meaningful increases in ability to buy things
and experiences for children that are associated with increases in income may asymptote. With
respect to effects of education among mothers in the United States, the distinction between having
and not having a college education seems to make more of a difference than differences in levels of
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education on either side of that boundary. Kalil et al. (2012) found most of the significant differences
in how mothers spent time with their children were between the college educated and noncol-
lege educated mothers; there were few differences between the mothers with some college and
those with a high school degree. Similarly, education-related differences in lexical and grammatical
properties of child-directed speech have been found for different levels of education up to a college
a degree, whereas years of education past college made little difference (Huttenlocher et al., 2007).
The threshold at which education differences matter may well depend on the domain of parenting
under consideration. Parental involvement in their children’s homework may be a domain in which
differences at the very high end of the distribution matter—as when adolescents come home with
assignments in calculus or physics.
Evidence of Indirect Effects of SES on Parenting

Examples of indirect effects of SES on parenting are most abundant in the literature on the negative
consequences of low levels of income and education. Three categories of intervening variables are
particularly prominent: parent well-being, social relationships, and home environment.
Parent well-being encompasses physical and mental health. Chronic health problems, such as dia-
betes and hypertension, and health risks, including obesity and substance abuse, are more prevalent
among low-income adults (Adler and Stewart, 2010). Health risk behaviors, such cigarette smoking
and sedentary lifestyle, are more frequent among less educated adults (Conti, Heckman, and Urzua,
2010). Health troubles are compounded when financially distressed individuals forego medical treat-
ment. Health disparities translate into parenting disparities because parents who are unwell and
unhealthy are limited in the time and energy they can devote to parenting. Parents who are physi-
cally incapacitated may be unable to perform more than minimal parenting duties. Parents who must
devote time and attention to treating their own ailments may have less of each to devote to parenting.
Mental health also has been found to vary as a function of income, of education, and of compos-
ite indices of SES (Hudson, 1988; Yu and Williams, 1999). Low educational attainment is related
to subsequent depression, antisocial personality disorders, and substance use disorders (Dohrenwend
et al., 1992) Low income predicts a heightened incidence and prevalence of depression and anxi-
ety disorders (Kessler and Bromet, 2013) as well as antisocial behavior and criminality (Farrington,
1993). SES disparities in parent mental health should translate into disparities in parenting behaviors.
Interactions with depressed parents lack warmth and, like those with their anxious counterparts, are
marked by negative affect, which reduces child engagement (Lovejoy, 1991). Depressed mothers talk
less to their children (Breznitz and Sherman, 1987; Rowe, Pan, and Ayoub, 2005) and are less respon-
sive to their children (Bettes, 1988) than nondepressed mothers. Furthermore, the negative effects
of depression on parenting are greater among low-income mothers (Lovejoy, Graczyk, O’Hare, and
Neuman, 2000). Troubled parents may withdraw from children, preoccupied with their own feel-
ings. For different reasons, depressed and antisocial parents may minimize parental monitoring and
supervision, the former because they don’t feel like it and the latter because they don’t appreciate the
need for it (Chilcoat, Breslau, and Anthony, 1996; Patterson and Capaldi, 1991).
Social relationships encompass parents’ relationships with each other, their larger social networks,
and the quality of the parent-child relationship, all of which have been hypothesized to be interven-
ing mechanisms in the relation between SES and parenting. Low-SES parents face special challenges
establishing and maintaining romantic relationships. Low-SES parents are less likely than higher-SES
parents to be married at the birth of a child and more likely to separate afterward (Stanley, Amato,
Johnson, and Markman, 2006). More low-SES parents report dissatisfaction with romantic partners
than do higher-SES parents (Amato, Booth, Johnson, and Rogers, 2007). Low-SES is associated
with poorer couple quality and stability (Conger, Conger, and Martin, 2010). In turn, poor couple
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or marital quality is a significant stressor that dampens the rate and quality of engagement between
parents and children (Belsky, 1984). Relationship dissolution has an equally profound impact on par-
enting; when one parent is physically removed from the child, the other must assume responsibilities
previously shared. Conflict and relationship instability can increase parent negative affect and depres-
sion, which undercut positive parenting (Forehand, Thomas, Wierson, Brody, and Fauber, 1990).
Some impoverished parents have social networks that are, in several key dimensions, also impov-
erished. Lower-SES parents have access to fewer sources of social capital than higher-SES parents;
their networks differ in terms of the influence, expertise, and information available from members
(Horvat, Weininger, and Lareau, 2003). Assuming that network members both inform and buttress
parenting, lower-SES parents may be disadvantaged from the moment a child is born. Transience
also undercuts social network support. Low-SES parents change jobs, residences, and the schools
their children attend more frequently than higher-SES parents, making it difficult to maintain sus-
tained ties with network members (Evans, 2004). Others have suggested, however, that some of the
social capital disadvantages outlined above are offset by (1) the density of ties (especially family ties)
between members of low-SES social networks and (2) a culture of sharing and communal assistance
among members of low-SES networks (Burton and Jarrett, 2000). Few would argue, however, that
the ameliorative effects of high quality, dense support networks can fully overcome the stressors that
low-SES environments place on parents’ social relationships.
Low-SES parents may also face special challenges when it comes to the quality of relationships
with offspring. For example, rates of disorganized attachment relationships in low-SES families are
nearly twice that in middle-SES families (van IJzendoorn, Schuengel, and Bakermans-Kranenburg,
1999), and part of the reason for this is likely the effect of environmental stressors associated with
poverty, which disrupt the establishment of trusting bond between parent and child (Cassidy and
Mohr, 2001). Violence, hunger, forced separation, and disease are but a few examples of environ-
ments that impinge on relationship quality. The indirect path from SES to parenting via parent-child
relationship quality starts from the premise that low-SES households experience more traumatic
events that, because of their unpredictability and lack of resolution, create an atmosphere of distrust
that permeates the parent-child relationship. Parents who have poor quality relationships with their
children are apt to alter their parenting practices, redirecting emotional and material resources toward
more successful and satisfying interpersonal outlets.
Home environments encompass the level of chaos in the household and the opportunities afforded
by the physical environment. Low-SES parents are apt to live in physical environments that are
challenging, both for what they contain and for what they lack. Environmental chaos describes
household crowding, traffic (i.e., frequency and number of people coming and going), noise, dis-
order, and an absence of routines (Wachs, Cueto, and Yao, 2016). Each has been linked to poverty
(Evans, Eckenrode, and Marcynyszyn, 2010). Poverty forces families to share small rooms that may
be located in neighborhoods with considerable ambient noise. Many of these neighborhoods are
unsafe, keeping people indoors. Physical disarray often reigns in cramped, highly trafficked quarters.
Overwhelmed parents, seeking respite, withdraw and avoid the company of children; chaos-induced
stress and fatigue also inhibit positive parent-child interactions and increase the likelihood of dis-
tracted and debilitating parent behaviors (Wachs, 1992). Household chaos is related to higher levels of
negative parenting (including intrusiveness and negative regard) and lower levels of positive parent-
ing (including sensitivity, positive regard, stimulation, animation, and engagement; Vernon-Feagans,
Garrett-Peters, Willoughby, Mills-Koonce, and the Family Life Project Key Investigators, 2012). This
finding obtained when the parenting assessment was based on a semistructured play activity that was
conducted in the same way across households, suggesting that the effects of chaos endure beyond
chaotic moments. Environmental disruptions that are out of parent control lead to diminished effi-
cacy, a sense that one cannot effect change (Corapci and Wachs, 2002). Low parent self-efficacy may
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extend to low parenting efficacy, including a lack of confidence in competence as a parent. Crowding
and noise also preempt private time between parents and children, which is an important oppor-
tunity for sharing, intimacy, and positive affect. Residential density adversely affects the amount of
talk addressed to children (Evans, Maxwell, and Hart, 1999). Finally, household chaos interferes with
the establishment and maintenance of routines, activities, and practices, bulwarks against the destruc-
tive force of unpredictability (Weisner, 2010). Daily routines give meaning to life, meaning that is
reinforced by ceremonial routines that articulate values and priorities. Routines make it easier to
accomplish the mundane tasks of childrearing, freeing up parent time, energy, and cognition needed
to identify and implement superordinate parenting goals.
Compared with their higher-SES counterparts, lower-SES parents rear their children in physical
environments that afford fewer opportunities for positive parent engagement. Poor parents can ill-
afford educational toys, games, and books that provide cognitive scaffolding opportunities (Bradley
and Corwyn, 2005; Bornstein and Putnick, 2012). Parent education plays a role too. Parents who
rarely read and never received musical instruction are unlikely to live in homes populated by books
and musical instruments. Absent games, books, sports equipment, and musical instruments, how do
parents and children spend their time together? Most low-SES households have a TV; far fewer have
a desk for schoolwork. A physical environment that makes it easy to watch TV and hard to read,
play, and do homework is one that presents obstacles for parents and children to engage in sustained,
shared cognitive and instructional activities. Social interactions will suffer too. Play and book reading
are important vehicles whereby parents and children share positive affect and create mutual interests
that provide a foundation for future exchanges (Ginsburg, 2007).
SES as a Moderator of the Relation Between

Predictors of Parenting and Parenting
SES functions as a moderator variable when the relation of other predictors to parenting differ
depending on SES. Child-driven effects (i.e., parent responses to child attributes) are strong candi-
dates for associations that may be moderated by parent education and by cultural expectations about
child behavior that differ between social classes (Laursen and Collins, 2009). Models that incorporate
SES as a moderator have often used social class as a grouping variable, on the assumption that cat-
egorically derived social class groups constitute qualitatively different environments (Hoffman, 2003).
A related, but distinct approach entails the use of continuous measures of SES to moderate associa-
tions between predictor variables and parenting variables.
Moderated models have been advanced that differ with respect to whether SES serves as a devel-
opmental asset or a developmental liability. In social risk models, social class is hypothesized to (1)
amplify the potential for the predictor variable to provoke adverse parenting or (2) inhibit positive
effects. The usual formulation posits risks arising at lower levels of SES (or lower class group mem-
bership). For example, maternal depressive symptoms lead to a deterioration in parenting among
less educated, but not more educated, mothers (Hoffman and Youngblade, 1998). In social benefits
models, social class is hypothesized to (1) inoculate against conditions that would otherwise motivate
detrimental parenting or (2) amplify positive effects. The usual formulation posits benefits arising at
higher levels of SES (or upper class group membership). For example, better educated mothers profit
more than less educated mothers from interventions designed to encourage responsiveness and stim-
ulating parent-child interactions (Walker et al., 2011). Another example of SES as a moderator may
be the findings that middle-class parents who hold the gender stereotyped belief that boys are better
in math than girls provide their daughters with more unsolicited help with math homework than
parents who do not hold that view (Bhanot and Jovanovic, 2005). In contrast, in lower-SES Latino
families, mothers with gender stereotypical beliefs are less involved in their daughters’ homework (8).
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SES as Mediator of the Relation Between

SES may be construed as a mediator or intervening variable that functions as an explanatory link
connecting a predictor variable to changes in parenting. Here too the clearest examples come in
the form of child-driven effects, wherein child characteristics are a cause of changes in SES, which,
in turn, affect parenting via previously discussed mechanisms. For example, child ill-health may be
indirectly linked to changes in parenting through changes in household income. Expensive, time-
consuming medical care could have a detrimental impact on family financial resources which, in
turn, can directly and indirectly alter the ways in which parents interact with children. Other child
attributes that provoke extraordinary expenses, such as special needs, handicapping conditions, gift-
edness, accidents, or legal difficulties, may be similarly linked to parenting through their impact on
household resources.
It is also possible that parent characteristics bring about changes in SES, which, in turn, affect
parenting. The argument that individual characteristics cause SES was first developed to explain the
fact that individuals with mental illness or very low intelligence tend to be socioeconomically disad-
vantaged as adults, regardless of the SES of their family of origin. The movement of such individuals
down the SES scale was termed “social drift” (Dohrenwend and Dohrenwend, 1969). The social drift
hypothesis also applies to individual characteristics such as temperament, personality, and intelligence
that have sources other than SES but may contribute to SES achieved in adulthood (Rowe and
Rodgers, 1997). Although these models were not devised to describe parenting outcomes, it is but a
short step to see how links from parent characteristics may be indirectly tied to changes in parenting
behaviors and attitudes, through a debilitating decline in socioeconomic conditions. In this case, SES
functions as a mediator, linking characteristics of parents that caused SES to parenting behaviors that
are caused by SES. It can be difficult, however, to disentangle characteristics of parents that predict
SES from characteristics of parents that are confounded with SES (Conger and Donnellan, 2007).
SES as a Spurious Correlate in the Relation Between

Because SES is confounded with many variables that are themselves predictors of parenting or
mediators or moderators of relations between other variables and parenting, it is possible that some
associations between SES and parenting are spurious. That is, there may be cases in which SES does
not actually contribute to the parenting outcome with which it is correlated, but rather there is a
third variable at work, influencing both SES and parenting. Examples of a third variable would be
intellectual or personality characteristics of individuals, genetically based or acquired, that give rise to
economic and educational success and may also, independently, be responsible for positive parenting
behaviors (Corcoran and Adams, 1997). Contextual factors such as family structure are also possible
spurious correlates. For example, single parenthood is associated with SES, and single parenthood is
thought to involve more permissiveness than parenting in two-adult families because single parents
have less time to invest in supervision and are more prone to elevate children as equals to serve as
friends or confidants (Astone and McLanahan, 1991).
In summary, multiple pathways potentially connect SES to parenting. SES can influence parent-
ing directly, as when education changes how mothers talk to their children (LeVine et al., 2012),
or indirectly, as when the stresses of poverty lead to depression and depression results in less verbal
interaction with children (Rowe et al., 2005). SES can moderate the influence of other factors, as
when higher-SES parents suffer less deterioration in their parenting as a result of depression than
do lower-SES parents (Hoffman and Youngblade, 1998). SES may also function as the mediator of
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other influences, if the expenses of child illness, for example, result in inadequate financial resources
to meet expenses, with the ensuing negative effects of financial stress on parenting.
The Contributions of Income, Education, and

Occupation to Relations Between SES and Parenting
Income, education, and occupation are correlated, but the kinds of capital they index, financial
capital, human capital, and social capital, are different things with different potential influences
on parenting. This section reviews evidence that helps to identify the constituent of SES at work
in the observed relations between SES and parenting. This unpacking of SES is relevant for
understanding the processes that shape parenting, and it is crucial for formulating public policy.
A persistent public policy question concerns the returns that should be expected from interven-
tions that increase household income versus interventions that directly target parenting behavior
(Berger, Paxson, and Waldfogel, 2009; Duncan, Morris, and Rodrigues, 2011; Gertler et al., 2014;
Leffel and Suskind, 2013; Noble, 2017). It has also been argued that education, particularly the
education of women, is the most cost effective intervention to affect parenting and children’s lives
(LeVine et al., 2001).
Income Effects
When families are poor, parenting suffers. The effects of poverty on parenting are largely indirect,
mediated by the multiple consequences of poverty, such as parental stress, household chaos, maternal
depression, high residential density, and food insecurity. Degrees of poverty matter. Differences in
financial resources within the low-income range are associated with differences in parenting qual-
ity (Lugo-Gil and Tamis-LeMonda, 2008), and increases in income through intervention programs
that provide cash transfers to poor families have been associated with increases in positive parenting
behaviors (Knauer et al., 2016). Outside of poverty, income has larger effects on material aspects
of the home environment than on parenting behaviors (Berger et al., 2009). Together the findings
suggest that income or financial capital affects parenting via two different pathways. One pathway
is a direct influence of financial resources on things parents can buy for their children. The rela-
tion is likely to be linear to a point, after which increments in financial capital may yield diminish-
ing returns. The second pathway is indirect and also nonlinear. That is, low levels of income have
negative consequences for parenting, mediated by the many negative effects of poverty on parent
well-being, parents’ social relationships, and the home environment. A curvilinear function relating
income to parenting is also possible if there are negative effects mediated by stresses and unrealistic
expectations that are associated with very high incomes (Luthar and Latendresse, 2005).
Education Effects
Effects of maternal education on parenting are often stronger and more reliability observed than
the effects of income (Davis-Kean, 2005; Menaghan and Parcel, 1991;Tamis-LeMonda et al., 2004;
Raviv et al., 2004; Wright and Wright, 1976). Effects of education have also been observed across
ethnic groups within the United States and across countries in the developed and developing world
(Kelley, Sanchez-Hucles, and Walker, 1993; Laosa, 1980; LeVine et al., 2012; Rubio-Codina, Attana-
sio, and Grantham-McGregor, 2016). Parental education affects a range of outcomes including the
nature of talk to children, the nature of discipline practices, and, more globally, the style of parent-
ing. The pathways through which education affects parenting are both direct effects of language
and literacy skills on how parents interact with their children and societal institutions (Lareau, 2011;
436
LeVine et al., 2012) and indirect effects mediated by parents’ knowledge of child development, and
parents’ own health and well-being (Bornstein et al., 2017; Rowe, 2008).
Occupation Effects
Parents’ occupations are statistically related to the other defining constituents of SES, education and
income, but some unique effects have been attributed to occupation per se. The clearest articulation
of occupation effects is the Kohn hypothesis (Kohn, 1963; Kohn and Schooler, 1982), according to
which the nature of their jobs shapes parents’ childrearing values and, thereby, their practices. Kohn
argued that blue collar jobs foster authoritarian childrearing because those jobs require obedience
and conformity, whereas white collar occupations foster authoritative parenting because those occu-
pations require initiative and independent thinking. More recent work has added to this the idea
that job characteristics also shape cognitive skills, with consequences for parenting (Duncan and
Magnuson, 2003). Consistent with this argument, Menaghan and Parcel (1991) found that the effects
on the home environment of mothers’ beginning employment depended on the nature of the job.
Beginning a job that was low in complexity was associated with a worsening home environment.
More broadly, if occupation is construed as one’s adult role in society, the Kohn hypothesis is the idea
that parents act to prepare their children for the same sort of adult role that they themselves play. That
idea remains current in theorizing about the source of differences in parenting behaviors across both
SES and culture (Kağıtçıbaşı, 2007; World Bank, 2015).
In summary, although the defining constituents of SES, income, education, and occupation are
correlated, it is possible to identify unique effects of each on parenting.
Remaining Questions and Future Directions

for Research on SES and Parenting
The research summarized in this chapter describes multiple ways in which parenting differs as a
function of SES and also identifies, to a degree, the sources of SES effects on parenting and the path-
ways of influence. Questions remain, however.
Capturing SES With Person-Centered Approaches

One open question concerns what measure best captures the differences in parents’ lives that derive
from their positions in the social hierarchy and that shapes their parenting. The current literature
reflects a near consensus that social class and SES are equivalent constructs and that SES is the pre-
ferred term (Conger and Dogan, 2007; Lareau, 2008). There also appears to be substantial agreement
that the three indictors of SES should not be combined into single composite scores (Conger and
Dogan, 2007). The most frequently used indicators are income and education; some studies use
one of these, some use both and analyze their effects separately. One motive for this approach is to
identify sources of influence that can be targets of intervention. If, however, the effects of SES in
parenting are not traceable to independent effects of education and/or income, then that approach
will miss its target.
Person-centered approaches have the potential to identify categories of individuals, defined by
constellations of SES components and other risk factors, who parent in meaningfully different ways.
For example, Rhoades et al. (2011) found membership in the categories Married, Stress and Depressed,
and Poor and Married were better predictors of parenting quality than household income. Such con-
stellations of variables, rather than single indicators or composite measures, may capture the “different
basic conditions of life” that Kohn (1963) referred to in talking about social class. Person-centered
437
approaches, such as the latent class analyses that yielded categories in Rhoades et al. (2011) have the
potential to find clusters of individuals who share the same basic condition of life (see Laursen and
Hoff, 2006).
Using Genetically Informed Designs

Another unresolved question is the degree to which genes account for SES, its correlated character-
istics in parents, and the parenting behavior that SES predicts. Genetically informed designs can test
the hypothesis that heritability accounts for much of the variance in models assessing associations
between parent characteristics and parenting (McGuire, 2003). The same models can examine effects
mediated through SES. To extend the example, there may be a shared genetic component that is
responsible for SES, parent mental health, parent marital status, and parenting attitudes and behaviors.
Especially likely, in this context, are gene-environment correlations. Inherited traits may exacer-
bate socioeconomic drift, as low-functioning parents seek less challenging occupational contexts
than high-functioning parents. Parents may also structure households that encourage or discourage
opportunities for parent-child contact, consistent with inherited traits that govern social stimula-
tion needs and social interaction preferences. Gene-environment interactions are also possible. That
is, SES-related differences in environments may alter the likelihood that genetic liabilities will be
expressed (Krueger, South, Johnson, and Iacono, 2008). Three processes have been described through
which differences among environments in norms, in opportunities, and in contextual triggers, affect
the expression of genetic liabilities (Rutter, Moffitt, and Caspi, 2006; Shanahan and Hofer, 2005).
Applied to SES and parenting they suggest that SES-specific norms may suppress the expression of
genetic liabilities for disapproved parenting behaviors; that SES-specific opportunities may allow the
expression of genetic liabilities which would go unexpressed in absence of those opportunities; and
that SES-specific contextual triggers (for example, stressors in poverty environments) may exacerbate
the expression of some genetic liabilities.
Cross-Cultural and Cross-National Generalizability

of Relations Between SES and Parenting
Also unresolved is the question of the cross-cultural generalizability of relations between SES and
parenting. Although the bulk of the research on SES and parenting has been carried out in high-
income, Western countries (Wachs and Cueto, 2016), a concern for child welfare and for the devel-
opment of children as the human capital of nations is an international concern (Bornstein and
Putnick, 2012; Engle et al., 2011; World Bank, 2015). The relation of income (or wealth) and educa-
tion to parenting has been studied in low and middle income nations across the globe.
Some generalizations are possible. Across countries that vary in income, parent behavior that is
related to children’s cognitive outcomes (e.g., reading books) varies more than does parent behavior
related to children’s socioemotional well-being (e.g., not leaving the child alone) (Bornstein and
Putnick, 2012). And within countries outside of the high-income Western countries usually studied,
income (or household wealth) and education have observable effects on parenting. Income effects
are observed when cash transfer programs result in changes in how much parents play with and read
to their children (Knauer et al., 2016). Education effects are observed when increases in maternal
schooling and resultant literacy, unaccompanied by changes in income, are related to changes in the
manner in which mothers talk to their children (LeVine et al., 2012; Rogoff, 2003). In both cases,
increases in income and education move parenting to be more like the parenting of parents with
higher incomes and education levels in Western, high-income countries. The parallels in effects of
income and education across different cultures and mean levels of income and education speak to a
remarkable generality of effects of SES on parenting. Another generalization, which is suggested by
438
the similarity of conclusions reached in a study of rural families in the United States and in a study of
poor families in Ecuador, is that both income (or wealth) and parent education affect child outcomes,
but the effect of education on outcome is transmitted more by parenting than is the effect of income
(Paxson and Schady, 2007; Vernon-Feagans et al., 2008).
It is less clear to what extent and in what ways the effects of SES on parenting may be different
across cultures. Culture itself is a strong influence on parenting, and one might expect that cultural
influences to create differences in the consequences of higher incomes or education for parenting.
There is evidence that culture can moderate the effects of SES (Gutierrez, Sameroff, and Karrer,
1988; Harwood et al., 1996). Among Anglo-Americans, lower-SES mothers were more concerned
with their children’s proper demeanor and higher-SES mothers were more concerned with their
children’s self-maximization (Harwood et al., 1996). Among Puerto Rican mothers, in contrast,
mothers were primarily concerned with proper demeanor regardless of SES.
Future research that makes use of new methods and that extends its reach to a wider range of
environments should shed further light on the processes by which SES shapes parenting and the
outcomes of those processes of influence.
Conclusions
Open questions notwithstanding, the existing research makes it clear that parenting differs across
socioeconomic strata. On average, higher-SES parents provide their children with experiences that
better equip them to be emotionally, academically, and occupationally successful than do lower-SES
parents. The reasons parenting differs across different levels of income, education, and occupation
are many. Providing a safe, stable, and cognitively stimulating environment for children costs money.
Being sensitive, responsive, and engaged in interaction with one’s children requires internal resources
that may be difficult to muster for parents who are stressed by financial hardship, depressed, ill, or
without social support. For parents with minimal education and low prestige jobs, an authoritative
(rather than authoritarian) parenting style and a great deal of conversation that includes the child’s
contribution may not come naturally. Parents with little control over their own lives may not see
themselves as shapers of their children’s development. The folk knowledge about parenting available
in lower socioeconomic strata may be better for preparing children for their parents’ roles in society
than in preparing their children for upward mobility. Parents with low levels of education are not
likely to read books on childrearing that will give them different advice. Parents with low levels of
education may feel (and be) poorly equipped to help their children in school—either at home with
assignments or in school as their children’s advocates.
One goal of research on SES and parenting is to inform public policy regarding how to address
inequality. This can be a sensitive topic. Some have and may continue to object that scientists and
policy makers have no business intervening in family life (Carneiro and Heckman, 2003). When the
lower-SES families are also members of minority groups, charges of cultural insensitivity can and
have been leveled. The counter argument to those concerns is that most parents truly want their
children to achieve and succeed. The research on SES and parenting makes it clear that SES-related
differences in parenting often serve to continue social inequalities and to limit the life chances of
children born into the lower levels of the social hierarchy.
Acknowledgments
Erika Hoff received support for the preparation of this chapter from the Eunice Kennedy Shriver
National Institute of Child Health and Human Development, grant HD068421. Brett Laursen
received support for the preparation of this chapter from the U.S. National Science Foundation,
grant 1248598. We are grateful to Thomas Adkins and Jordan McAdams for bibliographic help.
439
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14
CULTURE AND PARENTING
Xinyin Chen, Rui Fu, and Wai Ying Vivien Yiu
Introduction
Research has shown considerable cross-cultural similarities as well as differences in individual soci-
oemotional and cognitive functioning. For example, relative to their Western counterparts, children
and adolescents in many African and Asian societies tend to be less expressive of their emotions and
engage in less active interactions in challenging social settings (Chen, DeSouza, Chen, and Wang,
2006; Edwards, 2000; Oakland, Pretorius, and Lee, 2008). Chinese, Korean, and Mexican children
display more controlled and cooperative-compliant behavior than North American children in task-
oriented or limited-resource situations (Kagan and Knight, 1981; Oh and Lewis, 2008; Orlick, Zhou,
and Partington, 1990). Given this background, theorists and researchers have been interested in how
socialization factors, particularly parenting, play a role in explaining children’s socioemotional and
cognitive functioning and its cultural variations (Bornstein, 2012; Vygotsky, 1978). In most societies,
as a major socialization agent, parents provide care and monitoring to children and exert a significant
influence on their development, particularly in the early years. Cultural beliefs, norms, and values in
the society determine, to a large extent, parental socialization goals and parenting practices. Moreo-
ver, culture provides a guideline for social judgments of specific parenting behaviors, which in turn
shape their significance for child development.
This chapter is concerned with how culture is involved in parenting. We first discuss some theo-
retical and methodological issues in the study of culture and parenting. Then, we review research on
parental socialization goals and attitudes in cultural contexts. Next, we discuss how cultural values
affect the main aspects or dimensions of parenting, followed by a section on cultural involvement in
determining the developmental significance of parenting in terms of its relations with developmental
outcomes. Finally, we discuss the implications of social and cultural changes for parenting. We con-
clude the chapter with a discussion of future directions in the study of culture and parenting.
The Exploration of Culture and Parenting: A Brief History

The inquiry about culture and parenting has a long and rich history, mostly in cultural anthropology.
The initial interest, appeared in the early 1900s, was in whether children’s experiences, especially
childrearing experiences, in the early years could serve to explain cultural variations in personality.
Combined with the classical psychological theories, particularly Freudian psychoanalytic theory,
the culture and personality school (Benedict, 1934; Mead, 1928) attempted to search for culturally
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patterned early childhood experiences (e.g., mother-infant sleeping arrangement, strict discipline,

feeding, weaning, toilet training) that might lead to distinct adult personality profiles or national
characters in different societies. However, the over-simplistic views of the causal and determinative
force of early socialization practices in the development of modal personalities, and the work using
scientifically inadequate methodologies, became highly controversial and eventually abandoned by
most scholars and researchers in the field.
In the 1950s, a number of researchers, together with mixed training backgrounds of anthropology
and psychology, conducted a series of cross-cultural field studies of childrearing patterns and child
development (Barry, Child, and Bacon, 1959; LeVine, 1960; Whiting and Child, 1953; see Harkness
and Super, 2002, for a review). The most systematic project was the Six Culture Study of Socializa-
tion, led by Whiting and Whiting (1975), conducted in Juxtlahuaca (Mexico), Khalapur (India),
Nyansongo (Kenya), Tarong (Philippines), Taira in Okinawa ( Japan), and Orchard Town (USA).
In this project, the researchers examined typical socialization experiences and children’s behaviors
using ethnographic and observational methods. In societies where extended families lived together
in traditional styles (Nyansongo, Juxtlahuaca, Tarong), a high level of cooperation among family
members was required, and children were assigned various household tasks (e.g., collecting firewood,
fetching water from the river, cooking food, taking care of younger siblings). In contrast, in culturally
and economically more “complex” societies with class structures and occupational division of labor
(Taira, Khalapur, and Orchard Town), children were assigned fewer, and different types of, household
chores (e.g., cleaning their own rooms, picking up toys). Further analyses indicated that children in
the former societies displayed more nurturant-responsible behaviors (e.g., prosocial-cooperative and
compliant behaviors) than did children in the latter societies, and the researchers argued that the
differences were related to the performance of family responsibilities. According to Whiting and
Whiting (1975), the assignment of tasks to children and other socialization arrangements constitutes
a “child learning environment” for the development of behavioral styles, skills, and psychological
attributes.
The anthropological work in the 1950s and 1960s, including its theoretical frameworks and
methodologies, had a pervasive impact on the study of culture, parenting, and child development.
Researchers have continued to explore socialization issues using those data sets. For example, Edwards
(2000) reanalyzed the data from the Six Culture Study to examine peer interactions. The results
showed that children in the relatively “open” societies (Taira, Orchard Town) were more active in
social interaction and engaged in more sophisticated forms of play, such as fantasy or sociodramatic
play, than their counterparts in more “closed” societies (Khalapur, Nyansongo). According to Edwards
(2000), adult encouragement of play and the opportunity to choose playmates without adult supervi-
sion or control were some of the factors that might account for the cross-cultural variations.
Nevertheless, most contemporary developmental researchers, particularly in psychology, have not
followed the direction of the traditional research on culture and parenting. The deviation from the
traditional research may be mainly due to the fact that, although developmental researchers have
sought to understand social and cultural influences on human development, they are less interested
in the broad issues of anthropological paradigms, such as how settlement patterns, land use, and food
accumulation and storage in the society may determine childcare and childrearing practices and
how socialization experiences and children’s and adults’ behaviors are reflected in the “projective-
expressive systems” of societal-level collective beliefs, social-political systems, and general patterns
of psychopathology (e.g., crime, suicide; Worthman, 2010). Developmental psychologists, such as
Bornstein (Bornstein, Tal, and Tamis-LeMonda, 1991), Chao (1994), Cole, Tamang, and Shrestha
(2006), García Coll and colleagues (1996), Kağıtçıbaşı (1996), Keller and colleagues (2004), Hark-
ness and Super (1996), Trommsdorff and Friedlmeier (1993), and Wang and Fivush (2005), are
particularly interested in cultural involvement in specific parenting processes. Their research focuses,
among others, on (1) parenting beliefs, attitudes, and behaviors at the individual level and individual
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Xinyin Chen et al
variations within and across the societies (as reflected in the analysis of variance) and (2) cultur-
ally similar and distinct patterns of relations between parenting and child outcomes (as reflected in
the multi-group regression analysis or the group-invariance test of relations). The data collected
in studies using different types of designs (e.g., longitudinal, mixed-method) and psychometrically
established assessments and sophisticated statistical analyses allow researchers to rigorously examine
the display of specific aspects of parenting (e.g., parental warmth, power assertion) and their devel-
opmental significance in cultural contexts.
Theoretical and Methodological Issues in the Study

of Culture and Parenting
There are a number of theoretical and methodological issues in the study of culture and parent-
ing. For example, the definitions of culture vary remarkably in the literature. As a result, it is largely
unclear what should be included in the construct of culture and what aspects of culture are relevant
to understanding parenting styles and practices. Different theoretical frameworks and approaches,
often incompatible with each other, have been used to study culture and parenting, which is a main
source of controversies and confusions. Moreover, researchers disagree on how to obtain appropriate
and accurate information on culture in empirical studies that would help us interpret the findings in
a straightforward manner. We discuss some of these issues in this section.
Culture: Conceptualizations and Research Approaches

Culture has been defined as a variety of phenomena, ranging from the man-made part of the envi-
ronment (e.g., community settings), the shared lifestyle of a group of people (e.g., sleeping arrange-
ments for infants), the values and beliefs that are endorsed in a society concerning how individuals
should behave (e.g., filial piety in parent-child relationships), to the meaning system that individuals
use to understand the world (Cole and Cagigas, 2010; Oyserman, 2017). These definitions vary in
many aspects, such as levels of human functioning that culture should focus on. The broad defini-
tion of culture as the man-made environment may not be practically useful for research because the
impact of human activities exists in most, if not all, parts of the daily life environment. It is difficult
to conduct studies based on this definition to capture the effects of culture. Thus, many researchers
seem to prefer the narrower definition of culture focusing on values and beliefs that are shared and
commonly endorsed by people within the society (Hofstede, 1994). This definition works well for
comparing individuals in different societies on culture-related attitudes and behaviors; the prevalent
or predominant attitudes and behaviors of the individual in the society (e.g., mean scores) may indi-
cate general cultural orientations or norms. For example, generally greater support reported by Asian
parents than by North American parents for using power-assertive strategies in childrearing may
reflect the higher value that Asian cultures place on parental authority in the family (Chao, 1994).
This type of finding helps us understand the different socialization conditions of Asian and North
American children.
However, the notion of shared culture creates serious challenges for developmental research-
ers who are interested in how culture affects socialization and development at the individual level
because it is difficult to establish logical and empirical connections between the collective endorse-
ment of cultural values by a group and behaviors that are displayed by individuals. For example,
although research shows that Asian parents are more likely than European American parents to
endorse power-assertive and authoritarian parenting and that Asian students may outperform Euro-
pean American students in mathematics and science (Chao, 1994; Steinberg, Lamborn, Dornbusch,
and Darling, 1992; Stevenson and Lee, 1990), the relations among cultural values, authoritarian par-
enting, and children’s academic achievement are not straightforward (Chen, Dong, and Zhou, 1997).
450
An approach that researchers commonly use to address the issue related to the “sharing” aspect
of culture is to treat it as an individual-level construct, such as a personal trait (Triandis, 1995). In
this approach, each individual is viewed as having a unique culture, presumably due to personal
characteristics that affect his or her reactions to the influence of external cultural factors. Using this
approach, researchers may measure “culture” by observing individual responses to culturally relevant
stimuli (e.g., tasks that are used to activate a particular cultural mindset, expectation, or experience)
or by using self-report questionnaires such as those about collectivism versus individualism (views
and attitudes about whether the interests of the group or society should be considered more or less
important than those of individuals) or independent versus interdependent self-construals (under-
standing of the self as unique and separate from others or as intertwined with social context; Chiao
and Ambady, 2007; Oyserman, 2017; Singelis, 1994). The data allow for direct analysis of relations
between cultural variables and parenting or child behaviors.
However, the conceptualization of culture as a personal trait, and the corresponding research
approach, may not be particularly interesting to developmental researchers who are concerned with
culture as a context for socialization and human development (Bronfenbrenner and Morris, 2006;
Hinde, 1987; Whiting and Edwards, 1988). The contextual effect is unlikely to be adequately cap-
tured by an individual cultural trait, as one of possibly many individual traits, and its relations with
other psychological attributes or behaviors. The question is to what extent the contextual aspect of
culture is reflected in a personal trait. Individual reactions to specific tasks that are used to activate a
particular cultural mindset (Chiao and Ambady, 2007; Oyserman, 2017) are inevitably confounded
with other personal characteristics and experiences. Relatedly, many cultural traits, such as collectiv-
ism versus individualism and independence versus interdependence, conceptually overlap with other
personality or behavioral characteristics, such as sociability, communality, social assertiveness, and
autonomy. There are virtually no reasonable standards to categorize some traits as cultural and others
as noncultural. Consequently, maintaining conceptual and methodological clarity is challenging in
research.
Given this background, research using multiple approaches and methods will be important for us
to achieve a relatively comprehensive understanding of culture and parenting. In addition to cross-
cultural comparisons, for example, studies of the acculturation of domestic migrants and international
immigrants may provide valuable information about how cultural experiences (e.g., accommodation
to the new values, maintenance of the original values, integration of diverse values, Berry, 1997;
Bornstein, in press) modify parenting attitudes and behaviors. Lee (1995) assessed the participation
of Chinese immigrant parents and children in Canada in cultural activities (e.g., listening to Western
versus Chinese music, celebrating Western versus Chinese festivals, interactions with Western versus
Chinese friends) as an index of acculturation. The results showed that acculturation was associated
with parents’ and children’s social and psychological adjustment in the new environment.
The macro-level social change that is currently occurring in many traditionally agricultural coun-
tries, due to modernization, globalization, and technological development, also provides opportuni-
ties to explore cultural factors in the modification of parenting and child behaviors. Kağıtçıbaşı and
Ataca (2005), for example, found that, as Turkey became more urbanized over three decades, Turkish
parents changed their socialization goals; parents became more interested in joint leisure activities
with their children and were more likely to appreciate and encourage child autonomy and inde-
pendence. Chen and Li (2012) examined parenting attitudes of rural parents in China who became
urban residents during urbanization. Parents in urbanized families (families that changed the status
from rural to urban residency) were more likely than parents in rural families to encourage initiative-
taking, self-direction, and social participation in childrearing (e.g., “I encourage my child to take the
lead in initiating activities”; “I encourage my child to express his or her opinions in school and other
public places”; “I encourage my child to actively participate in social activities”). Moreover, parental
encouragement of initiative-taking was associated more strongly with children’s sociable-assertive
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behavior and social adjustment outcomes in the urbanized group than in the rural group, suggest-
ing that urban cultural values might strengthen the function of parental support for initiative-taking,
perhaps through enhancing parental specific guidance and assistance and children’s receptiveness to
the parenting effort (Chen and Li, 2012).
Culture, Parenting, and Human Development: Major Perspectives

Cultural influences on socialization and human development have been discussed mostly from
Bronfenbrenner’s socioecological perspective (Bronfenbrenner and Morris, 2006) and Vygotsky’s
cultural-historical perspective (Vygotsky, 1978). The socioecological perspective focuses on culture
as a part of the environment that affects individual behaviors. The early version of the perspective
(Bronfenbrenner, 1979) considered culture a distal influence on the child in the outmost layer of the
environment, along with laws, social class, customs, and broad ideologies. However, current views
have integrated cultural beliefs and values with proximal socialization forces including childcare,
education, and the family (Bronfenbrenner and Morris, 2006; Tietjen, 2006).
Consistent with this perspective, Super and Harkness (1986, 2002) proposed that cultural beliefs
and values are reflected in the “developmental niche” that includes three interacting subsystems:
the physical and social settings, the historically constituted customs and practices of childcare and
childrearing, and the psychology of the caregivers, particularly parental ethnotheories shared within
the community. In this model, culture affects parenting beliefs, attitudes, and practices directly as
well as indirectly through organizing the physical and social setting conditions (e.g., the living space
and daily life schedules for children) and culturally regulated customs of childcare and childrearing
(e.g., sleep arrangements for infants, different roles of fathers and mothers in childcare and educa-
tion). The three subsystems of the developmental niche may contribute to human development in a
manner of dynamic interactions among themselves and between them and child disposition (Super
and Harkness, 2002).
Vygotsky’s (1978) cultural-historical theory asserts that culture is an integral part of human
social and psychological functions. Individual psychological functions, such as thinking, memory,
and problem-solving, are based on, and constrained by, culturally represented mental tools, such as
language and symbols. Moreover, culture provides guidance for socialization processes in the inter-
nalization of language and other symbolic tools from the interpersonal level to the intrapersonal
level and in the organization and reorganization of psychological functions. In a study of the impli-
cations of transformations in sociocultural structures of the former Soviet Union in the early 1930s,
Luria (1976) demonstrated that the participation of young people and adults in rural villages in new
social and cultural activities (e.g., active group discussion and decision making in collectivized labor
and daily life) significantly affected their communication and interaction practices, social relation-
ships, and life goals and experiences. The transformation of socialization conditions and experiences
might lead to changes in individual socio-cognitive functioning (e.g., shift from concrete practical, or
“graphic-functional,” thinking that was constrained by the immediate and physical features of objects
to more mediated and abstract thinking).
The cultural-historical perspective has guided cultural and cross-cultural research in many coun-
tries, particularly those traditionally agricultural countries in Africa, Asia, and South America (Beach,
1995; Rogoff, 2003) Greenfield, Maynard, and Childs (2003), for example, examined mother-
daughter interactions in weaving in two cohorts (1969–1970 and 1991–1993) of Zinacantec Maya
children in Mexico and found that, as the society shifted from a subsistence-based agricultural life-
style to a commercial lifestyle, the pattern of mother-daughter interactions changed from scaffolding
by the mother to more independent trial-and-error learning.
Largely building on Vygotsky’s cultural-historical framework, Chen and colleagues (Chen,
2012; Chen and French, 2008) proposed a contextual-developmental perspective focusing on social
452
interaction as a context for human development. This perspective taps into socialization processes
in parent-child interaction. Specifically, according to Chen and French (2008), parents and children
evaluate and respond to each other’s behaviors during their interaction according to cultural norms
and expectations. Social evaluations and responses in turn serve to regulate parenting and child
behaviors, which jointly affect the development of parent-child relationships and children’s soci-
oemotional and cognitive outcomes. In this perspective, an important mechanism of parental influ-
ence on child development is the evaluation-response process in which parents display attitudes that
reflect their understanding of cultural norms and values. Whereas parental positive attitudes, such as
approval and encouragement, inform children that their behaviors are regarded as appropriate and
should be maintained and enhanced, parental negative evaluations and responses create pressure on
children to modify their behaviors. Children who are “ready” (Maccoby and Martin, 1983) and have
adequate social-cognitive abilities may comply with parents and act as parents expect. However, fail-
ure of children to do so may cause frustration, distress, and other negative emotions in both parents
and children, which may lead to negative parent-child relationships and externalizing problems if
the emotions are directed toward others, internalizing problems if the emotions are directed toward
the self, or both. Parenting, or, broadly, socialization toward culturally valued goals, is likely to be
more effective when parents understand children’s abilities and characteristics and exert influence
accordingly.
With age, children can play an increasingly active role in their socialization and development by
responding to parental demands and behaviors in parent-child interaction. Children’s responses are
also directed, to a large extent, by cultural norms and values (e.g., filial piety in traditional East Asian
cultures, autonomy in Western societies) that they learn over time from family members, peers,
media, and other sources. Whereas parenting affects children’s behavior and development, children’s
attitudes in turn serve to regulate parenting. For example, children’s resistance and defiance in parent-
child interaction press their parents to change their parenting attempts and practices. The failure of
parents to adjust their behaviors or to reach agreement with children may result in coercive family
dynamics (Patterson, 1982) and ultimately maladaptive parent-child relationships and other devel-
opmental problems. The contextual-developmental perspective on bi-directional social evaluation
and regulation processes in parent-child interaction helps us understand the issues related to relations
between culture and parental socialization goals and attitudes, cross-cultural differences in parenting
styles and practices, and culturally distinct outcomes of parenting.
Culture and Parenting Attitudes

As required by the demands of social, historical, and ecological circumstances, societies specify the
outcomes that children are expected to develop, which are indicated by values of children’s behaviors,
such as obedience, cooperation, independence, and emotion expressivity (Bornstein, 2012; Dennis,
Talih, Cole, Zahn-Waxler, and Mizuta, 2007; LeVine, 1988; Ogbu, 1981; Rogoff, 2003). Cultural
values constitute a basis for parental judgments of, and affective reactions to, children’s behaviors.
Chen and French (2008) argued that two basic dimensions of social-behavioral functioning, social
initiative and self-control, are reflected in most of these values and characterize major socialization
goals across cultures. Social initiative refers to the tendency to initiate and maintain social participation,
especially in challenging situations. Children who tend to display anxious and fearful reactions to
social novelty or perceived social evaluation may experience difficulties in spontaneous engagement
in social participation. Self-control refers to the ability to modulate reactivity in interactions to main-
tain social appropriateness, which is often indicated by cooperative or defiant-aggressive behaviors
in social situations.
In most Western cultures, because a major socialization goal is to help children develop auton-
omy and self-expression, social initiative is highly valued and strongly encouraged in childrearing.
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Xinyin Chen et al
Although self-control is viewed as useful for personal achievement, the culture generally empha-
sizes maintaining a balance between the needs of the self and those of others; undercontrol and
overcontrol are both considered maladaptive in social and psychological adjustment and thus not
encouraged in socialization. In many non-Western group-oriented societies, social initiative is not
highly appreciated because it typically does not affect group harmony or interpersonal relationships.
However, controlling personal desires and behaviors in social settings is strongly emphasized because
it is critical to group functioning. Cultural values of social initiative and self-control may have an
impact on parental attitudes toward specific behaviors, such as sociability-cooperation (active social
initiative with effective control), defiance-aggression (high social initiative and low control), and
shyness-inhibition (low social initiative and adequate control to constrain behaviors and emotions
toward self; Chen and French, 2008).
Parenting Attitudes in Socialization of Social Initiative

and Related Behaviors
Parental attitudes toward children’s social initiative may be indicated by how parents view and sup-
port children’s social interaction, particularly play with peers. Edwards (2000) argued that, in cultures
that value social initiative and self-expression, parents tend to encourage their children to engage in
active social interaction, such as fantasy or sociodramatic activities in play with others. Sociodramatic
behavior and pretense in play allow children to express their inner interests and personal styles while
controlling their social-evaluative anxiety in interaction (Chen and French, 2008; Farver, Kim, and
Lee, 1995). Relatively few sociodramatic activities are exhibited by Bedouin Arab children (Ariel
and Sever, 1980), East Asian children (Farver et al., 1995; Parmar, Harkness, and Super, 2004), Mayan
children (Gaskins, 2000), and rural Brazilian children (Gosso, Lima, Morais, and Otta, 2007). The
results from the Six Culture Study showed that children in Nyansongo (Kenya) and Khalapur (India)
displayed significantly lower levels of overall social engagement than children in Taira in Okinawa
( Japan) and Orchard Town (USA; Edwards, 2000). Moreover, children in Nyansongo and Khalapur
were less active, and their activities involved fewer fantastic characters or themes. Whereas socio-
ecological conditions (e.g., the structuring of activities, physical settings) are related to differences in
children’s play, cultural norms and values influence parental attitudes about whether children’s play
is stimulated or directed to develop social initiative skills. As noted by Edwards (2000), mothers in
Orchard Town were more likely than mothers in Nyansongo to encourage their children to enter-
tain themselves by playing and to direct their play to be imaginative, such as playing birthday party,
riding horses, and other sociodramatic activities.
In a cross-cultural study of parental values and attitudes in Israeli and Palestinian samples, Feldman
and Masalha (2010) examined the extent to which parents valued a set of child behavioral attributes.
Israeli parents placed higher value on social engagement and self-expressiveness in childrearing than
did Palestinian parents. Chinese parents traditionally tend to be unsupportive of children to display
initiative-taking (Chao, 1994; Chen et al., 1998), which is incompatible with the requirements of a
contemporary urban, market-oriented society. Chen and Li’s study (2012) revealed that parents in
urban families were more likely than parents in rural families to appreciate and value children’s quali-
ties of social initiative. Urban parents seemed to realize that it would be important for their children
to learn initiative-taking skills to adapt to the competitive environment.
Culturally based parental attitudes are also demonstrated in studies of children’s shyness-inhibition,
which represents a low level of social initiative. Chen and colleagues (1998) examined relations
between maternal acceptance, rejection, and punishment-orientation and shyness-inhibition among
2-year-olds in Canada and China. Children’s inhibition was assessed in free play and stranger-with-
toys (e.g., a toy dump truck, a black, noisy, and “smoking” toy robot) sessions in the laboratory
(García Coll, Kagan, and Reznick, 1984), and mothers completed a measure of positive and negative
454
childrearing attitudes. The results indicated that child inhibition was positively associated with moth-
ers’ rejection, concern, and punishment orientation in Canada. However, the results were the oppo-
site in China: Child inhibition was positively associated with maternal warm and accepting attitudes.
Different parental attitudes toward shyness-inhibition were also found across cultures in middle to
late childhood. Whereas the Western literature consistently indicates positive associations of shyness
with parental disappointment and rejection (see Rubin, Coplan, and Bowker, 2009, for a review),
shyness in elementary and high school children, especially girls, in China was found to be positively
associated with parental approval and support (Chen, Dong, and Zhou, 1997; Chen, Rubin, and Li,
1997). Kim, Rapee, Oh, and Moon (2008) found that shyness was positively associated with posi-
tive parent-child relationships and parental acceptance in Korean adolescents, but not in Australian
adolescents. There is evidence suggesting that Latin American parents may also have more posi-
tive attitudes than European American parents toward children’s shyness and social anxiety (Varela,
Sanchez-Sosa, Biggs, and Luis, 2009).
Parenting Attitudes in Socialization of Self-Control

Self-control has been valued and encouraged in most cultures, perhaps because it is required for
performance on both individual and group tasks (Eisenberg, Zhou, Liew, Champion, and Pidada,
2006; Maccoby and Martin, 1983; Whiting and Edwards, 1988). Parents typically start to make
effort to help children learn self-control in the first or second year of life when children understand
adults’ demands (Kopp, 1982). Nevertheless, research has shown that parents in more group-oriented
cultures tend to place greater emphasis on children’s self-control (Chen and French, 2008). Keller
and colleagues (2004) argued that whereas behavioral control is often viewed as interfering with the
child’s freedom in individualistic cultures, it is viewed as a duty, indicating social maturity and com-
petence, in group-oriented cultures. In many East Asian societies, such as China and South Korea,
training children to exert control is a major socialization goal from early childhood to adolescence
because controlling one’s behavior according to social rules is viewed as a critical component of
cultivating innate virtues (Chao, 1995; Ho, 1986). According to the Confucian principle of filial
piety, children must learn and maintain absolute obedience to parents. The higher expectation of
children’s self-control in Chinese and some other Asian societies is also related to the belief that
human behavior is malleable and controllable (Stevenson and Lee, 1990). Failure to adequately con-
trol one’s own behavior is considered incompetent and unacceptable because it is attributed to a lack
of effort. Consistent with this argument, Chen and colleagues (2003) found that, relative to North
American parents, Chinese parents expected their children to maintain a higher level of behavio-
ral control. Kohnstamm, Halverson, Mervielde, and Havill (1998) found that Chinese parents paid
more attention than Western parents to children’s conscientiousness or diligence and were more
concerned about poor control in their children. Similarly, in a study of parental values and goals,
Jose, Huntsinger, Huntsinger, and Liaw (2000) asked parents “How important do you think it is to
encourage the following personality traits in your child?” and found that both Taiwan Chinese and
Chinese American parents rated persistence, concentration, precision, calmness, and neatness as more
important than European American parents. The results clearly indicated that Chinese parents had
expectations of a high level of self-control in their children.
Greenberger, Chen, Beam, Whang, and Dong (2000) examined parental attitudes toward miscon-
duct and under-controlled behaviors in samples of senior high school students in Los Angeles (the
United States), Seoul (South Korea), and Tianjin (China). The results showed that U.S. adolescents
had higher scores than Korean adolescents, who in turn had higher scores than Chinese adolescents
on problem behaviors, such as risk-taking, substance use (e.g., drank hard liquor), fighting with
others, and school misconduct. The proportion of U.S. adolescents who displayed problem behav-
iors was substantially higher than that of South Korean, Chinese, or both (e.g., 52%, 18%, and 11%
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Xinyin Chen et al
engaged in “hitting or threatening to hit someone” in the United States, South Korea, and China,
respectively). More importantly, according to adolescents’ reports, parents in South Korea and China
had more negative attitudes toward, or greater sanctions against, adolescents’ involvement in mis-
conduct than did U.S. parents. South Korean and Chinese parents were less tolerant of adolescents’
under-controlled behaviors than American parents.
A similar study was conducted by Hackett and Hackett (1993) through interviewing Gujarati
parents, who emigrated from India and East Africa in the 1970s, and English parents of 4- to 7-year-
old children in Manchester, England. When asked about a situation in which a peer hit their child
or grabbed a toy from their child, significantly more Gujarati parents than English parents said that
they would encourage their children to exert self-control and that they disapproved of hitting back
or getting the toy back by force. When their child had a fight with a peer, more Gujarati parents
would stop the fight immediately and punish their child, whereas more English parents would ignore
the fight. The results indicated that Gujarati parents were less tolerant of their children’s under-
controlled behavior than English parents.
Dodge and colleagues (Deater-Deckard and Dodge, 1997; Deater-Deckard, Dodge, Bates, and
Pettit, 1996; Pinderhughes, Dodge, Bates, Pettit, and Zelli, 2000) conducted several studies about
parental attitudes and their relations with externalizing behaviors among African American and
European American children. The results generally indicated that African American parents, particu-
larly in low socioeconomic status, were more likely than European American parents to endorse the
use of harsh discipline. According to the authors, the experiences of heightened social and life stress
among African American parents in low socioeconomic status may affect their cognitive-emotional
processes (e.g., increased sensitivity) in response to socially provocative situations. The parenting atti-
tudes of African American parents and corresponding strategies they use may serve to prepare their
children to function in social settings in which others might treat them harshly. For example, when
presented with hypothetical vignettes describing situations in which a child engaged in misbehavior
(e.g., threatening peers who excluded the child from a game), African American parents were more
likely than European American parents to make hostile attributions about the child (e.g., the child
acted with hostile intent or in a negative-trait-like manner). Moreover, African American parents
were more worried than European American parents about their children growing up to have prob-
lems in the future (Pinderhughes et al., 2000). Thus, African American and European American
parents may differ in their attitudes, such as concern, anxiety, and vigilance, about children’s under-
controlled behaviors.
Cultural values of familism/familismo and respect/respeto in Latin societies are commonly believed to
be associated with parenting attitudes (Halgunseth, Ispa, and Rudy, 2006; White, Zeiders, Gonzales,
Tein, and Roosa, 2013). Familismo is concerned with feelings of loyalty to the family and commit-
ment to the family over individual needs and desires. Latinos place higher value on family respon-
sibility than European Americans, which is reflected in childrearing (Fuligni, Witkow, and Garcia,
2005; Halgunseth et al., 2006). The goal of respeto is to maintain close relationships through respect
for others, particularly elders. In Latino societies, children from the early years are socialized to learn
rules of respect for parents and other adults (Valdés, 1996). The values of familismo and respeto are
associated with the emphasis on parental monitoring, including high expectations of child obedi-
ence, self-control, and harsh parenting, which are believed to be important for familial solidarity and
deference (White et al., 2013).
Parenting Styles and Behaviors Across Cultures

A number of cross-cultural studies have been conducted concerning specific parenting styles and
behaviors. For example, researchers have examined parenting behaviors with infants and toddlers
in different societies in terms of whether parents engage in object-oriented activities or social and
456
emotional communications with the child (Bornstein, Cote, Haynes, Suwalsky, and Bakeman, 2012)
and whether parents display proximal (e.g., body contact, body stimulation) or distal (face-to-face,
object stimulation) behaviors with the child (Keller et al., 2004). Patterns of parenting styles and
behaviors are largely consistent with cultural values and associated socialization expectations in the
society. Moreover, extensive evidence has emerged to indicate that parents may differ across cultures
on the fundamental dimensions of parenting, such as parental warmth/affect and control, that are
emphasized in the developmental literature (Baumrind, 1971; Maccoby and Martin, 1983).
Cultural Variations on Parental Warmth and Affect Expression

Adequate parental warmth and affection in parent-child relationships are necessary for socializa-
tion and human development across cultures (Barnard and Solchany, 2002; Bornstein, Suwalsky, and
Breakstone, 2012). However, the level and phenotype of emotional expression may be moderated
by contextual influence (Bornstein, 1995; Saarni, Campos, Camras, and Witherington, 2006). It has
been argued that the expression of positive emotions in parent-child interactions is more encouraged
in cultures that place higher value on promoting children’s development of sociability, liveliness, and
autonomy (Harkness et al., 2007; Wörmann, Holodynski, Kärtner, and Keller, 2012). In Western
cultures, parental affect and warmth are regarded as important for fostering children’s socioemo-
tional competence, self-confidence, and self-expression (Chao, 1995; Henderlong and Lepper, 2002;
Rothbaum and Trommsdorff, 2007; Wang, Wiley, and Chiu, 2008; Wörmann et al., 2012). In other,
particularly East Asian, cultures, the explicit expression of emotional experiences in social settings is
viewed as inappropriate. Parents’ expression of affection toward children is believed to undermine
parental authority and the hierarchical and authoritarian organization of the family in which chil-
dren are expected to show respect and obedience (Chao, 1995; Cheah and Li, 2010; Chen, 2010;
Ho, 1986).
The results of studies with Asian and Western parents support this argument. Compared with
European American parents, Asian parents are less affectionate and less likely to engage in affective
communication with their children (Camras, Kolmodin, and Chen, 2008; Cheah and Li, 2010; Chen
et al., 1998; Dornbusch, Ritter, Leiderman, Roberts, and Fraleigh, 1987; Huntsinger and Jose, 2009;
Kelley and Tseng, 1992; Porter et al., 2005; Saw and Okazaki, 2010), which is related to their low
motivation to help children develop sociability and social initiative (Chao, 1995; Chen, 2010). For
example, Wu and Chao (2005) found that Chinese mothers were reluctant to express warmth openly
and directly; compared with European American parents, Chinese American parents displayed lower
levels of warmth and responsiveness (e.g., “Smiles at me very often”; “Often praises me”), although
they did not differ on adolescents’ ideals for parental warmth (e.g., “Ideally, how often do you think a
parent should smile at you?”), according to adolescents’ reports. In a study of family emotion sociali-
zation experiences, Saw and Okazaki (2010) noticed that Asian American youth reported consider-
ably fewer experiences of positive emotions with their parents in childhood and adolescence, such
as expression of physical and verbal affection, than European American youth (e.g., “On the phone
my dad’s like, ‘I love you’; ‘Yeah, love you too, Dad’ ”; “We’re affectionate in the sense of hugging, and
kissing on the cheek”).
Cheah, Li, Zhou, Yamamoto, and Leung (2015) illustrated cultural differences in the expression of
parental warmth through interviewing Chinese immigrant and European American mothers. In the
study, the researchers examined cultural variations in not only the amount of warmth expressed, but
also reasons for expressing warmth and the specific practices of expressing warmth. Chinese mothers
reported expressing less warmth than European American mothers. Whereas both groups of moth-
ers believed that maternal warmth was important for children’s development, they provided different
reasons and reported different practices of expressing warmth. Relative to Chinese mothers, Euro-
pean American mothers reported more outward and direct demonstrations and communications of
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Xinyin Chen et al
warmth (“Hugging and kissing”; “Tell her ‘I love her’ all the time”) that served to promote children’s
individuality, emotional expressivity, and self-confidence. They also tended to provide more elabo-
rative explanations of emotions and to encourage children to focus on their own feelings. In con-
trast, Chinese mothers focused on warmth as a foundation for an exertion of parental training and
guidance. Accordingly, Chinese mothers were more likely to indicate expressing warmth through
instrumental support, including satisfying daily routine needs (e.g., “I make him his favorite dump-
lings”; “I help him organize clothes and quilts”) and providing guidance and learning/educational
opportunities (e.g., “I help and guide the child to develop good habits and be polite”; “I try to let
her go to good schools”).
Relatively low levels of parental warmth and affect expression were found in other Asian socie-
ties, such as Indonesia, India (Daga, Raval, and Raj, 2015), Japan (Trommsdorff, and Friedlmeier,
2010), and South Korea (Louie, Oh, and Lau, 2013), as well as non-Asian and non-Western socie-
ties, such as Cameroon (Wörmann et al., 2012), Arab (Rasmi, Chuang, and Safdar, 2012), and Tur-
key (Yaman, Mesman, van IJzendoorn, Bakermans-Kranenburg, and Linting, 2010). For example,
Wörmann and colleagues (2012) found that, compared with German mothers, Nso mothers in
rural communities in Cameroon displayed less positive emotions such as smiles to infants and were
less likely to respond to infants’ smiles. Rasmi et al. (2012) showed that Arab youth in both Canada,
Egypt, and Lebanon perceived lower parental warmth than European Canadian youth. Yaman et al.
(2010) reported that, after controlling for maternal age and education, compared with native Dutch
mothers, second-generation Turkish immigrant mothers in the Netherlands were less supportive, as
indicated by little expression of positive emotions, and were more intrusive, as indicated by the lack
of respect for the child’s autonomy in exploratory or problem-solving situations and by interfering
with the child’s needs and interests. Deater-Deckard and colleagues (2011) found that, based on
parental and child reports, parents of 7- to 10-year-olds in multiple non-Western countries, includ-
ing China, Jordan, Kenya, and Thailand, were significantly lower on warmth (e.g., “I let my child
know I love him/her”) than parents in the United States. European, Asian, and Latino American
parents within the United States did not differ on warmth in this study. Inconsistent with the results
in Deater-Deckard and colleagues (2011)’s study about different ethnic groups within the United
States, Ispa and colleagues (2004) found in an observational study of mother-toddler interactions
at home that Latin and African American mothers expressed a generally lower level of warmth
(e.g., physical and verbal expressions of affect, words of encouragement and praise) than European
American mothers.
Cross-cultural variations in parental warmth and affect have been observed across Western
nations. Harkness and colleagues (2007) found that, compared with mothers in the Netherlands,
Spain, and the United States, Italian mothers had the higher scores on socioemotional closeness,
which indicates parental effort to promote infants’ sociability and parental sensitivity to infants’ need
for social interaction. Other researchers (Bornstein, Putnick et al., 2012; Hsu and Lavelli, 2005) also
found that, relative to North American mothers, Italian mothers were more sensitive and more likely
to display affection in interactions with their children.
Cultural Variations on Parental Control, Directiveness, and Discipline

Parental control is another dimension that has received a great deal of attention in developmental
research (Baumrind, 1971; Maccoby and Martin, 1983). In Western cultures, the views of parental
control and its function in child development have been mixed, depending on the nature of parental
control. For example, psychological control, referring to parenting behaviors that are intrusive of
the child’s feelings and thoughts through invoking guilt, shame, and anxiety, is believed to impede
the development of autonomy and self-direction (Barber and Harmon, 2002). However, behavioral
control, referring to parental monitoring and supervision of the child’s behaviors, is often considered
458
useful and necessary for children to learn social standards and to control their behaviors according to
social standards (Barber and Harmon, 2002; Stattin and Kerr, 2000). Nevertheless, the general cultural
orientation that focuses on promoting individual freedom and autonomy of children in Western
societies constitutes a context that encourages parents to exert control, directive, and discipline with
caution, especially as children grow older (Greenfield, Suzuki, and Rothstein-Fisch, 2006).
Parental control and monitoring are regarded as more acceptable and desirable in many non-
Western cultures. In some Asian cultures that highly value parental involvement in children’s various
activities, parental control is perceived by parents and children as an indication of responsibility and
care (Chao, 1995; Ho, 1986). Moreover, it has been argued that parental assertion of control is one
of the primary means through which parents express their warmth (Nomura, Noguchi, Saito, and
Tezuka, 1995; Rothbaum and Trommsdorff, 2007). Parents who do not exert strict and compre-
hensive control over children’s behaviors and activities are viewed as irresponsible and incompetent,
indicating a lack of affection toward the child or inadequate parenting skills. Wang and Fivush (2005)
found that even during parent-child conversations about emotional events, Chinese parents empha-
sized discipline, focused on understanding social norms, hierarchy, and moral rectitude, and were
direct in correcting children’s responses.
Parents in Asian societies are expected and encouraged to exert control over children’s activities,
such as learning, through a variety of methods. Pomerantz and colleagues (Cheung and Pomerantz,
2011; Ng, Pomerantz, and Lam, 2007; Pomerantz and Wang, 2009) found that, compared with
American parents, Chinese parents used more explicit and systematic control and direction to help
children’s learning. They focused more on children’s errors on tasks, emphasized failure more than
success, and expressed lower satisfaction with children’s performance. Similar results were found by
other researchers (Huntsinger et al., 2000; Miller, Wiley, Fung, and Liang, 1997; Stevenson and Lee,
1990). In the sections that follow, we focus on the discussion of parental use of shaming, as a salient
form of psychological control, and behavioral directiveness across cultures.
Shaming. Fung (2006) found that parents in Taiwan often used shaming practices to help children
develop prosocial and other socially appropriate behaviors. Similarly, Wu and colleagues (2002)
found that shaming was perceived more favorably by Chinese parents than U.S. American parents
and that Chinese parents were more likely than U.S. American parents to use shaming in parenting.
Fung (2006) argued that feelings of shame can activate the processes of self-examination and self-
criticism, which make individuals recognize the discrepancies between their behaviors and social
standards and motivate individuals to improve their performance. Parents in Chinese culture tend to
believe that shaming practices can be used effectively to help children learn social responsibility, self-
control, and other group-oriented behavioral qualities. Thus, Chinese parents use shaming strategies
to teach children right from wrong, including verbal markers (e.g., name-callings, threats of aban-
donment, social comparisons), paralinguistic cues (e.g., emphatic stress, angry intonation), vocal cues
(e.g., sighs, making disapproving sounds), nonverbal techniques (e.g., staring, frowning), and silence
(Fung, 1999; Schoenhals, 1993).
Shaming practices seem to be used commonly in socialization in other non-Western cultures as
well. For example, Cole, Bruschi, and Tamang (2002) found that, in response to situations in which
a child received unjust treatment or made a mistake (e.g., “You’re outside your house, playing with
everyone. Suddenly, everyone is shouting and running to see something. You run after them but then
you fall down and get mud all over you. Your mother and all the others laugh at you.”), whereas
U.S. children were more likely to express anger, Tamang Nepali children were more likely to report
that they would feel ashamed. Keller and Otto (2009) also found that rural Nso mothers expected
to see the expression of shame in their children earlier than German mothers (at approximately 26
versus 34 months). Unlike their Western counterparts, parents in these cultures do not seem to be
concerned that using shaming or similar psychological control may have harmful effects on children’s
development of autonomy and self-worth.
459
Xinyin Chen et al
Behavioral control. Cross-cultural researchers have examined extensively parental behavioral con-
trol, or parental control in general, and found differences between parents in Western and non-
Western societies. For example, Louie and colleagues (2013) found that, based on parental reports,
South Korean and other Asian American parents of preschool-age children were higher on parental
control than their European American counterparts. Chao and Aque (2009) found that Chinese
American adolescents rated their parents as more strict and controlling (e.g., “Insists that I do exactly
as I’m told”) than European American adolescents, which was consistent with findings from other
studies using similar assessments (Lin and Fu, 1990).
Based on observations of family interactions, Jose et al. (2000) found that Chinese American
parents displayed higher directiveness (e.g., parent made the decision and child followed without
question; parents quickly corrected child’s errors) than European American parents. Bornstein, Cote,
and colleagues (2012) observed mother-infant interactions in the home setting and found that,
whereas U.S. American mothers were more likely than their infants to display responses to social
initiations, perhaps in an attempt to foster child independence, Japanese mothers were more likely
than infants to lead and direct interactions, which might reflect a controlling parenting style. Japanese
American immigrant mothers appeared to be in-between, with later generation immigrant mothers
being more similar to European American mothers. Lamborn, Nguyen, and Bocanegra (2013, p. 54)
conducted interviews with Hmong American adolescents regarding their perceptions of mothers’
parenting practices. A salient finding was that adolescents frequently talked about mothers’ assertions
of authority that included supervision (e.g., “She keeps track of us . . . I have a cell phone, so I either
call her and tell her where I’m at, or I’m safe and stuff. Or either she’ll call me and ask me where am
I, or what am I doing, am I safe and all this stuff.”).
Keller and colleagues (2004) observed mother-child interactions in Nso families in rural Cam-
eroon, Costa Rican families, and middle-class Greek families. A major focus of the study was on
proximal parenting (body contact, body stimulation), which was believed to reflect parental control
that facilitates child obedience and regulation. Consistent with the hypotheses, the results indi-
cated that Cameroonian Nso mothers displayed more proximal parenting than did Costa Rican
mothers, who in turn displayed more of such behavior than Greek mothers. Keller and colleagues
(2004) argued that parental behavioral control may be viewed as detrimental to the development of
autonomy in individualistic cultures but as conducive to the child’s development of competence in
group-oriented cultures.
Relatively high levels of parental control and directiveness have been found in multiple stud-
ies with Latin and African American families. Ispa and colleagues (2004) examined, in samples of
Mexican American, African American, and European American families, maternal intrusiveness in
mother-toddler interactions at home, which was defined as the degree to which the mother con-
trolled the child’s play instead of allowing for the child’s preferences. Typical maternal intrusive
behaviors included grabbing toys, taking charge of the activity, and imposing her own agenda with-
out letting the child shape the focus or pace of play. The results showed that Mexican American
mothers, including those who were acculturated toward American values, and African American
mothers scored higher on intrusiveness than European American mothers. Similarly, Carlson and
Harwood (2003) found that Puerto Rican mothers in San Juan, Puerto Rico, were more likely than
European American mothers in the United States to display parenting behaviors that limited and
directed the movements and activities of their infants at 4, 8, and 12 months in mother-child inter-
actions at home. Relatively high parental control has also been found among Latin American and
African American parents of adolescents. For example, Blair, Blair, and Madamba (1999) found that
Hispanic and African American high school students rated their parents as exerting greater control
than European American counterparts (e.g., “How often do your parents check on your home-
work?”“How often do your parents limit you going out with friends?”); the former two groups were
similar to Asian Americans students.
460
According to Ispa and colleagues (2004), Latin and African American mothers might engage in
more intrusive behaviors because they might consider these behaviors less intrusive than European
American mothers. It is also possible that the higher directiveness of Latin and African American
mothers indicates their greater effort to raise well-behaved children with self-control and other
qualities that are crucial for group functioning (Carlson and Harwood, 2003). For example, the values
of familismo and respeto in Latino cultures are likely to promote parental monitoring and strictness in
parenting (Halgunseth et al., 2006; White et al., 2013).
Authoritative and Authoritarian Parenting Styles

Baumrind’s model of parenting (1971), particularly the notion of authoritative and authoritarian par-
enting styles in the model that are derived from different levels of warmth and control, has often been
used as a conceptual framework to interpret findings of cross-cultural studies. Parents in Western and
non-Western cultures have been found to differ on these parenting styles. Authoritative parenting,
based on high warmth and high control, is regarded as ideal in most Western cultures because it is
believed to promote child autonomy and social initiative. Authoritative parents are described as sup-
portive of children’s exploration and independence and respectful of their rights while, at the same
time, monitoring their behaviors and activities. In contrast, authoritarian parenting, based on low
warmth and high control, focuses on child obedience and the use of coercive and punitive strategies in
parent-child interactions. This parenting style is typically viewed as undesirable for child development
in Western cultures because it may undermine children’s independence and initiative-taking abilities.
Relative to parents with a North American or Western European background, Asian, African
American, and Latino parents have been found to use more “adult-centered” and power-assertive
parenting strategies (e.g., “I do not allow my child to question any decisions I make”), which appear
to fit into Baumrind’s authoritarian parenting style. For example, compared with European Ameri-
can parents, Asian parents in Asia or other regions are found to be less likely to use inductive reason-
ing and engage in equalitarian communication with children and more likely to express coercion
and punishment orientation in parenting (Dornbusch et al., 1987; Jose et al., 2000; Lin and Fu, 1990;
Porter et al., 2005; Rudy and Grusec, 2006). However, it has been argued that the coercive and high-
power parenting that Asian parents use is typically associated with parental care and warmth (Chao,
1995), which would be conceptually categorized as authoritativeness, although it may be indeed the
case that Asian parents tend to display relatively few prototypical “authoritative” parenting behaviors,
such as listening to the child, encouraging the child to express opinions, providing explanations, and
engaging in affect communication with the child. For example, Jose et al.’s (2000) observational study
revealed that Chinese American parents and Taiwanese parents were more directive, exerting more
parental control, than European American parents, but they were equally warm with their children.
Therefore, labeling Asian parenting as authoritarian may be incorrect and misleading. There is also
evidence indicating that, although Latino parents tend to be more restrictive and directive than
European American parents in their interactions with children, Latino parents display similar levels
of sensitivity and affection and thus may not necessarily be “authoritarian” (Carlson and Harwood,
2003; Varela et al., 2004). Similarly, it may not be appropriate to characterize African American par-
ents, who tend to use more harsh discipline than European American parents (Deater-Deckard and
Dodge, 1997), as authoritarian if their parenting is based on acceptance of, and concern for, the child.
In short, cross-cultural differences have been found in major parenting behaviors or strategies.
However, broad categories, such as authoritative and authoritarian parenting styles, as described in
the Western literature (Baumrind, 1971) may not be adequate or appropriate for characterizing
parents in Asian, Latin, or African American societies. It may be useful to examine specific parent-
ing behaviors or aspects in different societies, to avoid confusion resulting from the broad parenting
categories, especially in terms of their relations with children’s developmental outcomes.
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Xinyin Chen et al
Relations Between Parenting and Children’s Socioemotional

and Cognitive Functioning in Cultural Context
Cultural norms and values serve to shape the significance, or functional meanings, of parenting
behaviors (Bornstein, 1995), as indicated in their relations with children’s developmental outcomes.
Researchers have investigated the role of culture in affecting the relations between parenting and
children’s adjustment (Chen, Liu, and Li, 2000). In the section that follows, we review and discuss
work on the significance of the major dimensions of parenting in different cultures.
Parental Warmth and Children’s Socioemotional

and Cognitive Functioning
It has been argued that parental warmth may have universal positive implications for children’s
development (Rohner, 1986; Rohner, Khaleque, and Cournoyer, 2005). The findings from different
societies appear to support this argument; despite cross-cultural variations in the level and form of
parental affective expressiveness in childrearing (Cheah et al., 2015; Wörmann et al., 2012), higher
parental warmth generally tends to be associated with children’s competence and positive adjustment
in social, academic, and psychological domains at the within-culture individual level (Chen et al.,
1997; Eisenberg, Liew, and Pidada, 2001). For example, Bronstein (1994) observed family interactions
between parents and children at 7–12 years in a small city in central Mexico and found that parental
warmth (showing affection, praising the child) was positively associated with children’s cooperative
and compliant behaviors and negatively associated with children’s resistance. In a study with 4-year-
old children in China, Chen, Wu, Chen, Wang, and Cen (2001) conducted observations of parenting
behaviors at home and child aggression in peer interactions in a laboratory setting. Maternal warmth
and responsiveness (e.g., loving gestures such as hugging, smiling, physical comfort, praise, support-
ive responses) were negatively associated with child aggression. Kim and Rohner (2002) found in a
sample of grades 6–12 Korean American students that maternal and paternal warmth as reported by
adolescents was positively associated with academic achievement. These results suggest that parental
warmth and emotional support constitute important social and psychological resources for children
and adolescents to learn appropriate social behaviors and achieve in school.
Nevertheless, how parental warmth contributes to the development of specific behaviors or qual-
ities may vary across cultures, depending on the extent to which the behaviors or qualities are valued
in the society. In a 2-year longitudinal study with elementary school children in China, Chen and
colleagues (2000) found that maternal warmth positively predicted perceived self-worth and nega-
tively predicted loneliness and depression, whereas paternal warmth positively predicted social com-
petence and school achievement and negatively predicted aggression. The differential significance of
maternal warmth and paternal warmth might be related to the culturally prescribed distinct roles of
the mother and the father in Chinese families. As indicated by Ho (1987), like their Western coun-
terparts, Chinese mothers are responsible for providing care and affection to the child in dealing with
tasks and problems in daily life. Unlike Western fathers, who typically interact with the child like
a playmate, however, Chinese fathers are expected and required to help children learn social values,
develop appropriate behaviors, and achieve in academic areas because, as the authority figure in the
family, fathers have the most responsibility to maintain and enhance the status and reputation of the
family. This is clearly stated in the famous Three Character Classic—“It is the father’s fault if a child
is not well educated” (Mo, 1996). Therefore, whereas mothers are more sensitive to children’s emo-
tional problems, fathers may provide greater guidance and assistance to children in learning social
skills and achieving school success.
As another example, Chen et al. (2003) examined relations between maternal parenting and
toddlers’ complaint behaviors in a clean-up setting in China and Canada. Maternal warmth was
462
positively associated with toddlers’ committed compliance (working willingly and wholeheartedly
on the clean-up task without adult’s sustained control) in China, but situational compliance (gener-
ally cooperative but requiring repeated maternal control to stay on task) in Canada. Thus, warm
and supportive parenting may be beneficial for the development of self-control in both China and
Canada. However, due to specific cultural expectations, parental effort may be directed to the sociali-
zation of different child behaviors in toddlerhood, such as internally driven self-control in China and
controlled behavior that is maintained by external prompts in Canada.
Parental Control and Children’s Socioemotional

and Cognitive Functioning
Relations between parental control and children’s developmental outcomes vary evidently across
cultures. Whereas parental psychological control is associated with more maladaptive socioemotional
and cognitive functioning than parental behavioral control in general (Stattin and Kerr, 2000), both
types of parental control seem to be associated with fewer child adjustment problems in many non-
Western societies than in Western societies, with a few exceptions (Wang, Pomerantz, and Chen,
2007). The endorsement of parental authority in non-Western cultures and the encouragement
of child autonomy in Western cultures may provide a basis for parents and children to view and
react to the use of control strategies in parenting and eventually affect their significance for child
development.
Several cross-cultural studies with North American and Asian families showed that, whereas
high parental control, particularly psychological control, was associated with children’s externalizing
(e.g., aggression, rule-breaking behaviors) and internalizing (e.g., anxiety/depression, somatic com-
plaints) problems in North America, these associations were not significant in Asian nations such as
Hong Kong and South Korea (Fung and Lau, 2012; Kim and Rohner, 2002; Rudy and Halgunseth,
2005). For example, Louie and colleagues (2013) found that, whereas parental control (e.g., shaming)
reported by European American parents was positively associated with child anger and dysregula-
tion, the associations were not significant for Asian American and South Korean families. Similarly,
Ho, Bluestein, and Jenkins (2008) found that relations between parental ratings of control or harsh
parenting and teacher ratings of child aggression differed in European Canadian and South Asian
Canadian families; the relation was positive in the European Canadian group but negative in the
South Asian Canadian group.
Chao and Aque (2009) examined how Asian (Chinese, South Korean, and Filipino descent) and
European American youth reacted to parental control (e.g., “If my parent insists that I do exactly
as I’m told [or brings up past mistakes when s/he criticizes me], then I would feel [1 = not angry to
4 = very angry].”). The results showed that Asian adolescents felt less angry with parental strictness
and psychological control than European American adolescents. The attitude of gratitude for parents’
sacrifice and a sense of family responsibility are likely to reduce the negative emotional reactions of
Asian adolescents. Adolescents’ feelings of anger served a protective function in buffering against the
effects of parental psychological control on behavioral problems among European American, but
not Asian American, adolescents. Chen et al. (2000) also found a complicated pattern of relations
between parental control and children’s adjustment outcomes in Chinese children. Specifically, this
2-year longitudinal study showed that paternal control positively predicted later social competence
for children who were initially competent, but positively predicted aggression for children were ini-
tially high on aggression. Paternal control appeared to promote adaptive development for children
who were competent, but exacerbate behavioral problems for children who lacked self-regulatory
abilities and displayed negative emotional reactions.
Parental control also seems to have weaker effects on children’s behaviors in African Ameri-
can, Latin American, and other group-oriented societies than in European American societies.
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Xinyin Chen et al
Deater-Deckard and Dodge (1997) reported that parental harsh discipline in early childhood, based
on interviewers’ ratings, was predictive of later teacher ratings of externalizing problems among Euro-
pean American children, but not African American children. Similarly, Ispa and colleagues (2004)
found that parental intrusiveness or directiveness in mother-infant interactions positively predicted
later child negative behavior and decreased engagement with the mother in European American
and more acculturated Mexican American families, but not in African American or less acculturated
Mexican American families. Wood and colleagues (2017) studied the moderating role of accultura-
tion in relations between parental control and toddlers’ behaviors among Puerto Rican families in
the United States. Mothers’ frequent use of control (e.g., assertive commands in a clean-up session
such as “put that in the box,” prohibitions such as “you cannot play anymore,” physically controlling
behaviors such as taking toys from child and physically orienting child toward toy box) was related
to child defiance with negative emotions for families reporting high levels of acculturation, but not
for less acculturated families. Wood et al. (2017) argued that parental control had less harmful effects
on children’s development when it was exerted in the context of Latino cultural values that endorse
familismo and respeto. Finally, Atzaba-Poria (2011) found that parental authoritarian control tended to
be associated with more externalizing and peer problems among Israeli children than Former Soviet
Union (FSU) children. Taken together, these findings indicate that parental control and directiveness
may be associated with children’s social and behavioral functioning in different manners in different
societies. Group-oriented cultural values in Asian, African American, and Latino societies appear to
mitigate the adverse impact of parental control, including harsh discipline, shaming, and other types
of psychological control, on child development that is often seen in Western societies.
Implications of Social Change for Parenting

The implications of social change for socialization and human development have been recognized in
psychology, sociology, and other social sciences (see Elder and Shanahan, 2006; Silbereisen and Chen,
2010). Greenfield (2009) proposed a theory of social change and human development, focusing on
links among changes in sociodemographic conditions, cultural values and learning environments,
and human development. According to Greenfield (2009), the movement between Gemeinschaft
(rural residence, substance-based economy, the use of simple technology, informal education) and
Gesellschaft (urban residence, commercial or industrialized economy, the use of complex technol-
ogy, formal education) conditions corresponds to the shift between collectivistic and individualistic
values, with the former encouraging compliant-cooperative behavior and the latter encouraging
independent behavior. From a different perspective, Kağıtçıbaşı (2012) argued that the global urbani-
zation and modernization may not necessarily lead to cultural change in a linear fashion from an
emphasis on relatedness to an emphasis on autonomy. Instead, as rural societies become increasingly
urbanized, the emergence of combined autonomous-related values may help families and children
adapt to the demands in the new environment.
Largely consistent with Kağıtçıbaşı’s view (2012), Chen (2012, 2015) argued from a pluralist-
constructivist perspective that the migration of populations, advance in information technology, and
interaction among cultural systems across regions have led to an integration of diverse values and
lifestyles, which represents a distinct feature of the context for human development in the socie-
ties today. An important task in socialization is to help children and youth develop competencies
and qualities that allow them to function successfully in the environment with diverse, including
conflictual and complementary, cultural beliefs and values. According to this perspective, although
values from different cultures play a significant role in shaping parenting attitudes and practices, new
values may be integrated with the cultural traditions in the society. The integration of different values
may be illustrated by the experience of Eastern European parents who were exposed to American
464
individualistic cultures as described by Nesteruk and Marks (2011)—although the parents gave their
children more choices and were more willing to listen to their opinions, they still attempted to make
sure that their children learned responsibilities and self-control. In addition, Chen (2015) argued that,
whereas traditionally agricultural societies become more individualistic during urbanization, West-
ern societies may become more inclusive of group-oriented values, even though some aspects of the
Western context may become more individualistic as suggested by Greenfield (2009). Thus, parents
and children in both Western and non-Western societies benefit from their experience of mixed
values because maintaining a balance between pursuing one’s own interests and establishing positive
group relationships is critical to social and individual functioning (Maccoby, 1998).
Research findings have shown the impact of social change on parenting attitudes and behaviors,
although they may not necessarily support a particular perspective or model. For example, Kağıtçıbaşı
and Ataca (2005) found that the parental goals of Turkish families changed over three decades along
with the transformation of Turkish society. As a consequence of urbanization and socioeconomic
development, the need for material dependence within the family was reduced, and accordingly,
parents perceived more positively the child’s autonomy and initiative-taking. Turkish parents in 2003
were also more likely than their counterparts in 1975 to appreciate the “psychological value of the
child” (e.g., pleasure watching children grow, fun to have young children around, to have someone
to love and care for). Chen and Chen (2010) reported that, as China was more urbanized, Chinese
parents became less controlling and less power-assertive. At the same time, they became more sensi-
tive to children’s feelings and needs and more inclined to encourage them to engage in exploration.
Similar changes in parental warmth and control in urban Chinese and South Korean parents have
been noted in other studies (Lu and Chang, 2013; Park, Joo, Quiroz, and Greenfield, 2015; Ren and
Edwards, 2016; Way et al., 2013; Yoshikawa, Way, and Chen, 2012).
Lamm, Keller, Yovsi, and Chaudhary (2008) found cross-generational differences in paren-
tal beliefs in German, Indian, and Cameroonian Nso families, presumably associated with social
changes. Specifically, individuals in the urban Nso community experienced rapid change in their
living conditions and education. Consequently, compared to Nso grandmothers, Nso mothers used
more autonomy-oriented parenting in their care of young infants. In Delhi and Berlin, grand-
mothers and mothers also differed in their discourse style; relative to grandmothers, mothers placed
greater emphasis on autonomy and expressed less concern with the maintenance of their authority
in parenting.
In contrast to Lamm et al.’s study (2008) concerning German parents, however, there is initial
evidence that, parents, particularly fathers, in European countries are becoming more active in moni-
toring children’s activities and education. Collishaw, Gardner, Maughan, Scott, and Pickles (2012),
for example, found that, compared with their counterparts in 1986, English youth in 2006 reported
that their parents had higher expectations concerning going to school, doing homework, and being
polite. Youth in the 2006 cohort also reported that their parents exerted higher levels of supervi-
sion, such as telling parents where they were going and what activities they engaged in out of home.
In addition, youth in 2006 were more likely than youth in 1986 to disclose their activities to their
parents. With the current trend of migration and technological development, it will be interesting
to systematically examine how cross-cultural communications and interactions modify parenting in
different societies.
Conclusions
Findings from a variety of research programs have indicated that cultural beliefs and values, particu-
larly those concerning socialization goals, play important roles in shaping parenting attitudes and
behaviors. Moreover, culture affects the functional significance of parenting practices, as indicated in
465
Xinyin Chen et al
their relations with children’s developmental outcomes. Macro-level social changes that are occur-
ring in many countries due to the movement of populations, technological development, and inter-
action among different political, economic, social, and cultural systems have led to the co-existence
and integration of diverse cultural values, which have substantial implications for socialization and
human development in Western and non-Western societies today.
In this chapter, we have reviewed and discussed theoretical perspectives and research on issues
related to culture and parenting. Our discussion has focused on parental attitudes in the sociali-
zation of social initiative and self-control and related parenting styles and practices, particularly
parental warmth and control. Culture influences broader aspects of parenting, such as parental ideas
about what constitute effective childrearing, parenting strategies (e.g., resources and time parents
allocate to childcare), and parent-child communication styles (Harkness and Super, 2002; Keller,
2012). It will be important to examine the relations between cultural values and these aspects of
parenting.
Several other future directions may be suggested. First, parenting and its impact on child devel-
opment occur in broader social contexts, such as community services, school practices, and peer
relationships. Researchers should investigate how these contexts work together with parenting in
mediating and moderating the influence of culture on children’s behaviors. It will also be interesting
to explore how culture is reflected in macro-level social conditions, such as child- and family-related
policies and the mass media, which in turn affect parenting and child development.
Second, it has been increasingly recognized that children play an active role in socialization
and development (Chen, 2012; Edwards, de Guzman, Brown, and Kumru, 2006). Children may
display their active role through their reactions to parenting. As demonstrated by Chao and Aque
(2009), cultural norms and socialization experiences are likely to affect children’s interpretations
of, and attitudes toward, parenting behaviors and, thus, facilitate or undermine their effects. Chil-
dren’s perceptions of “normativeness” of parenting strategies, such as power assertion, may moderate
the relations between the strategies and developmental outcomes (Davidov and Khoury-Kassabri,
2013; Gershoff et al., 2010; Lansford et al., 2005). There is also extensive research on the moderat-
ing effects of child temperament on relations between parenting and development (see Chen and
Schmidt, 2015). Nevertheless, our understanding of the active role of the child in socialization is
still rather limited, especially in non-Western societies. Researchers need to pay greater attention
to this issue in the future.
Third, cross-cultural studies of parenting have mostly relied on parent or child self-reports because
of relatively low costs for data collection and advantages in data analysis. Self-report methods have
obvious limitations that may threaten the validity of cross-cultural comparisons, including culturally
specific response biases, the “reference group” effect, and differences in the understanding of ques-
tions or items in measures (Schneider, French, and Chen, 2006). Relative to self-reports, observa-
tions, either in the controlled laboratory or naturalistic home settings, can provide more objective
information that allows for more straightforward cross-cultural comparisons. However, conducting
observations of parents and children in equivalent conditions, developing culturally sensitive and
comparable coding systems, training coders to reliably code data from different cultures, and inter-
preting the results beyond observed variables are highly challenging tasks for researchers. Given the
strengths and weaknesses of each of the major methods, a multimethod approach is apparently ideal
to reduce potential biases in cross-cultural research.
Finally, researchers who study culture and parenting are often interested in comparing families
in Western self-oriented societies with those in group-oriented societies. This framework, which
focuses on broad categories, has been criticized for its inadequacy in describing the within-society
heterogeneity and the complex cultural systems (Miller, 2002). Continuous investigation in a variety
of cultures and sub-cultures will be crucial for a more comprehensive understanding of the contex-
tual nature of parenting.
466
Acknowledgments
The writing of this chapter was supported by a grant from the National Science Foundation (#BCS-
1225620).
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15
ENVIRONMENT AND
PARENTING
Robert H. Bradley
Introduction
It’s impossible to know exactly how humans lived in ancient times or exactly how characteristics
of the geography or the dwellings in which they lived and the objects that they used affected what
parents did or the activities of children. For more than two centuries, archeologists have gathered
artifacts from around the globe trying to find clues as to what ancient life was like. Their findings
have sometimes been startling—humans appear to have been engaged in certain types of activities,
such as organized hunting for meat and specialized tool making, centuries or even millennia earlier
than previously thought. Although much remains unclear about the enterprise of child rearing for
most human populations (especially those of long ago), one thing is clear: Life has radically changed
over the millennia, including the activities of parents. Some activities that could hardly have been
imagined even 100 years ago (e.g., watching an infant sleeping in a crib via a baby monitor while one
works in one’s own office; playing a game on cell phones with one’s 9-year-old while the 9-year-old
is spending time with grandma; selecting a video to play in the car for the 4-year-old in the back seat)
are now commonplace, a consequence of new gadgets that have proliferated in the lives of families
in wealthier nations. Changes in the accouterments of life have long led artists to imagine what life
might be like in the future and how human activities would change over time, sometimes amusingly
so (see the TV program, The Jetsons; https://en.wikipedia.org/wiki/The_Jetsons).
Reading academic journals, full of arcane measures and experiments, restricted samples, and statis-
tical findings that show low “effect sizes” can sometimes give the impression that environmental con-
ditions do not have that much to do with how a parent parents. Accounts in the public media offer
a contrast to such an impression. In September of 2015 a picture of a young boy lying face down on
a beach in Turkey was transmitted by the news media. It sent shock waves around the world—and
induced not a few tears. He was an innocent, vulnerable child, drowned along with family mem-
bers trying to escape the horrors in Syria at that time. Over the past quarter century, pictures and
accounts of children caught up in war and ethnic strife have emerged in the popular press (especially
electronic media) and in governmental publications, many containing graphic depictions of parental
efforts to keep children safe and healthy under the challenging conditions present. As an example, the
U.S. press has presented numerous such accounts regarding the tens of thousands of unaccompanied
minors and the tens of thousands of other people traveling as families (mostly mothers and young
children) who have tried to cross the U.S. border in the past 5 years in an effort to escape poverty and
gang violence endemic in Central America. The pictures often show skinny, scantily clad, sometimes
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sick children either alone or with a dazed parent. These pictures and accounts of real lives, like the
graphic depictions of parental efforts to deal with the horrific conditions of Nazi internment camps
found in movies and books make far clearer than most scholarly accounts of parent-child interaction
just how potent environmental affordances can be in shaping what parents do in their role as parents.
Who can forget the tortured agony of Sophie Zawistoska as she faced the primal “choice” regarding
the most basic of parenting acts: staying with one’s child in a time of peril as described in William
Styron’s classic American novel, Sophie’s Choice. And who can forget the frantic efforts of Candido
and America, illegals living in Los Angeles, who tried to save their infant daughter as all three swirled
downhill in the floodwaters that blasted through their makeshift home in the L.A. hills, as depicted
in T. C. Boyle’s novel, Tortilla Flats.
Anthropologists have also published scintillating accounts of how social and physical surroundings
penetrate the actions of caregivers, influencing what they do and how often they do it. Notable is
Scheper-Hughes’s (1989) wrenching portrayal of home life in Alto do Cruzeiro, the largest shanty-
town in Bom Jesus da Mata, located in northeast Brazil. Like many third-world shantytowns, Alto
do Cruzeiro exudes chronic poverty, high mortality, marital instability, and disconnection from kin.
Scheper-Hughes makes clear how the physical surroundings and the social actions that compose the
home environments of Alto children are shaped by the larger economic, cultural, and historical forces
present in urban Brazil. The point of her article is that these conditions produce mothers who do not
care. On average, the women of Alto experience 9.5 pregnancies, 3.5 child deaths, and 1.5 stillbirths.
The women face child disease and death with fatalism and stoicism. They identify infants who have
the strength and will to survive, then nurture them. To others they attribute a “will to die” or the
kind of “child sickness” that cannot be cured—angels awaiting flight to heaven. These infants are
viewed with indifference, often revulsion, and are allowed to die. Most of the infants die at home and
are buried with little ceremony in unmarked graves, a form of passive infanticide to which mothers
have been socialized by other women in the community and even by the church. Scheper-Hughes
(1989, p. 117) concluded that:
Frequent child death remains a powerful shaper of maternal thinking and practice. In the
absence of firm expectation that a child will survive, mother love as we conceptualize
it (whether in popular terms or in the psychobiological notion of maternal bonding) is
attenuated and delayed with consequences for infant survival.
Life in most U.S. American homes and most homes in other wealthy nations tends not to be so
stark. Parents who live in countries ranked high on the Human Development Index (HDI) tend
to have more financial and social capital, resources that give them more control over daily life and
resources that afford their children greater opportunity for optimal development. The HDI was
created by the United Nations based on the idea that people needed more than just money to live
a good life (United Nations Development Programme, 2017). Because people need a diverse set of
capabilities to thrive and to function well as members of society, the experts connected with the UN
identified key markers connected to health and education as well as economic status to rate coun-
tries with respect to the country’s assets for supporting the overall well-being of its citizens. By no
means does every family living in a high-HDI country have access to such capital; and by no means
does every child living in such families receive the “benefits” of such capital. However, the broad
differences in daily life and parenting behaviors observed in high-HDI versus low-HDI countries
makes clear the general impact of environmental affordances on parenting and child development.
To go back to what is frequently revealed by pictures of children in the media, one can contrast those
pictures of middle-class U.S. American parents and children playing together in a scenic city park
with numerous facilities for enjoyable activity and pictures of poor children in a third-world country
roaming with peers in muddy city thoroughfares.
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Robert H. Bradley
The physical and social settings in which families live constitute part of what Super and Harkness
(1986) call the developmental niche. The developmental niche of families regulates the microenvi-
ronment for children. A good example of its impact on parent behavior can be seen when comparing
sleep patterns of Kokwet infants from Kenya and infants living in urban United States (Super and
Harkness, 1986). Kokwet babies generally sleep with their mothers and are rarely left alone. Babies
living in the United States, by contrast, generally sleep in their own beds and often in a room sepa-
rate from where their parents sleep. Similar differences are observed between Mayan infants and U.S.
infants (Morelli, Rogoff, Oppenheim, and Goldsmith, 1992). German babies also tend to sleep sepa-
rately from parents, in a room of their own—what Valentin (2005) describes as a cult of independ-
ence. Such sleeping arrangements increase the likelihood of particular forms of behavior on the part
of both parent and infant. There tends to be other differences in the social and physical affordances
of the developmental niche as well. Homes with lots of toys and material affordances, for example,
increase the likelihood parents will act as companions to the child provide the structure for the child’s
emerging skills (i.e., they set the opportunities and incentives for child action; Sutton-Smith, 1986).
In effect, the setting helps determine what a child can see, hear, and do and what parents are likely
to do in their role as caregivers.
It is important to state at the outset that what parents do in a given moment or across time is not
fully determined by the social and physical features of the settings they inhabit. As Belsky and Jaffee’s
(2006) model of parenting shows, parenting is a complex function of many factors, including the
parent’s own history and personality and the characteristics of the child being parented. Bornstein
(2016) discussed how the characteristics of a parent (e.g., health, beliefs, cognitions, proclivities) can
act in complex ways in response to the characteristics of a particular child and to the multiple aspects
of context that surrounds parent and child to determine the ways that the parent behaves in and
through time as they enact the role of caregiver for the child. It is a process that can be quite mercu-
rial, as both parent and child are active constructors of their environments as well as responders to
what the environment affords (Ford and Lerner, 1992; Wachs, 2000). The actual process of parenting
involves numerous moment-to-moment exchanges with the child and the environment—parenting
in the rain is a bit different than parenting while basking in the sun.
It is not easy to define what constitutes the parenting environment or to specify its boundaries.
In a child’s mind, what home means almost certainly is not confined to the activities and physi-
cal features contained within the four walls of the child’s residence. Moreover, the idea of “home”
evolves throughout the course of development—not even ending when someone becomes an adult.
Lawrence and Low (1990) cited studies done in several cultures that demonstrate that the social
boundaries of households do not necessarily coincide with the physical boundaries of dwellings.
Home experiences may include any number of events that are seen as connected to home and family
but not coterminous with the physical boundaries of a place. For a 4-year-old, home life may include
time spent with dad in the neighborhood park. For the 9-year-old, it may include playing outside
with buddies or the drive with mom to a music lesson. For children of all ages, it may include Christ-
mas dinner with grandma. So, too, it may include arguments between family members—wherever
they occur. Thus, it would seem useful to have a somewhat capacious view on what constitutes the
environment of parenting. Such a view seems particularly relevant today—in a world where connec-
tions between family members include text messages, Facebook posts, and FaceTime encounters. By
the same token, the concept of the parenting environment as fully divorced from any place of resi-
dence is probably not useful either because the idea of home is generally associated with identifiable
places. For both parent and child, many of the ideas and attitudes connected to “home” have their
roots in scenes, episodes, and “scripts” which emanate from particular concrete places where family
activities occur (Abelson, 1981).
The concept of the parenting environment might best be thought of as phenomena emanat-
ing from the family setting broadly construed. Such an expansive definition is consistent with the
476
position of social anthropologists who contend that the social boundaries of household units do not
necessarily coincide with the physical boundaries of their dwellings (Altman, 1977; Lawrence and
Low, 1990). Such an expansive definition seems more appropriate when one considers the actual liv-
ing conditions of many families (e.g., families living out of a car or shelter, parents sharing dual cus-
tody, children who spend large amounts of time at their parents’ sides in places other than home such
as fields and workplaces, and the exchanges that increasingly occur via social media). The concept of
the parenting environment as not fully confined to a place becomes useful because the activities of
parents and the roles they play in children’s lives tend to change as children age—and sometimes as
the parent’s own life changes as a consequence of work, extended family, or community obligations
(Belsky and Jaffee, 2006; Bornstein, 2016; Bronfenbrenner, 1999). Essentially, the parenting environ-
ment encompasses the settings where the activities of parental caregiving take place; it is empirically
defined (Rapoport, 1985).
In this chapter, I describe some of the key tasks of parenting and how they relate to children’s
well-being. I also discuss how aspects of the social and physical environment affect that ways par-
ents carry out these tasks and their success in doing so. I attempt to portray parents as conscious
actors in dealing with the needs of their children, actors who make decisions in light of structural
characteristics of the settings in which they operate and the resources available to accomplish par-
ticular goals. I conclude the chapter with a consideration of the difficulties scholars face in trying to
explicate what are complex connections between environmental affordances and parenting behavior.
Although the behavior and characteristics of a child are an integral part of the environment of par-
enting, my focus on this component of Belsky and Jaffee’s (2006) model will be limited. Most of the
chapter deals with the broader ecology.
Fundamental Tasks of Parenting

For about two decades, we have organized our consideration of parenting around the idea that par-
ents must perform a number of key tasks in their role as caregivers, tasks that help children maintain
health, adapt to the demands imposed by the environment, and exploit opportunities afforded by the
settings they encounter (Saegert and Winkel, 1990). More concretely, we have proposed that parents
must act in ways that assure (1) sustenance, (2) safety, (3) stimulation, (4) socioemotional support, (5)
structure, (6) surveillance, and (7) social integration (Bradley, 2006; Bradley and Caldwell, 1995)—
seven fundamental tasks of parenting. This organizing framework is a simple heuristic; and we make
no assumption that all parenting behaviors can be neatly packaged into these seven categories or that
other ways of organizing would not work as well. The idea behind these seven fundamental tasks of
parenting derives from what is known about human needs and arousal systems (Maslow and Murphy,
1954) and ideas about complex living systems in dynamic environments (Ford and Lerner, 1992;
Wachs, 2000). The framework also recognizes that humans (both parents and children) are conscious,
self-constructing agents who actively engage and structure the environments they encounter (Lerner,
Johnson, and Buckingham, 2015; Lewis, 1997). The framework would appear consistent with the idea
that children must survive and develop the kinds of skills or assets that allow them to function as com-
petent, caring adults who are ready to function well within society (Maccoby and Martin, 1983; Scales
and Leffert, 2004). Based on the idea that positive development in highly advanced organisms requires
a multiplicity of skills that result in self-productivity through time (Cuhna, Heckman, and Schennach,
2010), positive caregiving requires actions that promote skill development in multiple domains.
Sustenance
It is essentially a sine qua non that not being well nourished or having access to decent health
care can lead to poor health and even death (Pollitt, 1988). We use the broad term sustenance to
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Robert H. Bradley
encompass those caregiving actions that are designed to promote basic biological health. In keeping
with the broad idea of sustaining health for children, we have included under the sustenance category
parental efforts protect children from pathogenic conditions, such as pollutants, passive cigarette
smoke, and exposure to heavy metals (Evans, Kliewer, and Martin, 1991; Jacobson, Jacobson, Padgett,
Brummitt, and Billings, 1992; Tong and McMichael, 1992). At present, there is limited informa-
tion on how most conditions in the social and physical environment actually affect what parents
do to provide sustenance to their children. On the face of it, it might seem that when children face
conditions that threaten their health, parents would take whatever actions are necessary to provide
for basic needs, such as food and shelter, or to remove a child from exposure to toxic substances.
However, the few studies that are available suggest that parental actions are not so easy to predict.
Early studies suggested that in times of food shortage, women and children are likely to suffer more
from undernutrition than men because men consume a higher proportion of available food. Later
studies observed that this pattern was more reflective of societies having a high degree of gender
inequality (Safilios-Rotschild, 1980; Schofield, 1974; Zeitlin, 1996). There is also evidence that food
deprivation does not always disproportionately affect children and that the burden of food shortage
does not always fall equally on children of all ages. For example, Leonard (1991) found that caloric
stress was greater on adults than on children among Andean agriculturalists, in which all able-bodied
individuals participated in agriculturally related activities to the greatest extent possible. Wandel and
Holmboe-Ottesen (1992) found that in Tanzania the burden fell more heavily on preschool-age
children than on older children because older children were needed to help with the harvest. By
contrast, McDonald, Sigman, Espinosa, and Neuman (1994) found that in Kenya infants do not suf-
fer as much during food shortages as do older children. In effect, although food availability is known
to affect how much parents feed children, there is no consistency in the response of parents to food
shortage.
Part of what determines what parents do to provide both nutrition and health care for their chil-
dren is where they live. For example, when families live in homes that have serious structural defects
or lack basic amenities, there is greater likelihood of exposure to dust mites, rats, cockroaches, and
other pests that have potential to impair health (Rao and Phipantankul, 2011; Rauh, Chew, and Gar-
finkel, 2002). Infants exposed to indoor wood smoke and cockroaches are also more likely to develop
asthma (Salam, Li, Langholz, and Gilliland, 2004). Indoor exposure to dust mites and smoke appears
to be a higher risk for asthma development than outdoor exposures to allergens (Etzel, 2003). Such
circumstances tend to motivate parents to enact behaviors designed to afford protection to offspring.
Living in such conditions can also cause parents to spend more time dealing with children’s health
problems—perhaps at the cost of spending less time engaging children in activities that promote
competence.
Where the family residence is located can also impact parental actions connected with health. For
example, surveys point to marked differences in the kinds of health care that is accessible to families
both within and across nations (Delamater, Messina, Shortridge, and Grady, 2012; Health and Places
Initiative, 2014; Radley and Shoen, 2012). Consequently, parents who live in areas where access to
health care (basic or more specialized) is limited likely take different actions with respect to meeting
their children’s health needs than parents with ready access to particular forms of health care (Gamm,
Hutchison, Dabney, and Dorsey, 2003). Lack of needed health care facilities can be particularly prob-
lematic if a child has a serious disability or complicated medical condition, as the normal approaches
to keeping a child nourished and healthy may not always work (Floyd and Gallagher, 1997; Gannoni
and Shute, 2010; Hauser-Cram, Warfield, Shonkoff, and Krauss, 2001). Likewise, whether one lives
in a run-down or upscale neighborhood, in a rural or urban area, or in a highly developed or not so
highly developed nation can have a bearing on what foods parents acquire for consumption by family
members, with impacts on the actual food consumed by children (Pearson, Biddle, and Gorely, 2009;
Rasmussen et al., 2006). Some neighborhoods have been characterized as “food deserts” due to lack
478
of access to fresh fruits and vegetables, making it more likely that parents will feed children meals that
contain inadequate amounts of key nutrients and too much sugar and fat (Darmon and Drewnowski,
2008; Gordon-Larsen, 2014). Area of residence is also a factor in how likely it is that parents engage
their children in outdoor physical activity or that they allow children to engage in outdoor physical
activity on their own, leading to better overall fitness for children (Dunton, Almanza, Jerrett, Wolch,
and Pentz, 2014; Kaczynski and Henderson, 2007; Zaltauske and Petrauskiene, 2016).
Socioeconomic status (SES; income, occupation, and educational attainment) is implicated in
many parenting decisions concerning health care and nutrition (Caulfield, Huffman, and Piwoz,
1999; Darmon and Drewnowski, 2008; Ezzati et al., 2002; Janevic, Petrovic, Bjelic, and Kubera,
2010; Nagash, Whiting, Henry, Belachew, and Hailemariam, 2015; Prickett and Augustine, 2016).
Sometimes the components of SES operate independently and sometimes they operate in conjunc-
tion with parental adjustment disorders, such as addiction or depression or geographic location, to
determine patterns of parental behavior ( Janevic et al., 2010; Magura and Laudet, 1996; Kandala,
Madkungu, Emina, Nzita, and Cappuccio, 2011; Miller and Chen, 2013). SES is associated with
differences in parenting behaviors that are connected with nutrition and health promotion begin-
ning very early in a child’s life (e.g., behaviors related to prenatal care and breastfeeding; Hajizadeh,
Tehrani, Simbar, and Farzadfar, 2016; Shahla, Fahy, and Kable, 2010).
Although family income is associated with various parental decisions pertaining to nutritional
intake and health care for children, the association is not a simple one. A study of low-income women
in the United States showed that both initiation and cessation of breastfeeding were connected to
perceived stress of four types: (1) financial, (2) emotional, (3) traumatic, and (4) violent partner behav-
ior (Dozier, Nelson, and Brownell, 2012). Studies of other low-income populations have also shown
that inter-partner violence is associated with undernutrition and illness in young children (Ackerson
and Subramanian, 2008; Hasselmann and Reichenheim, 2006; Rahman et al., 2012).
Cultural beliefs and connections with social networks can also have an impact on parental deci-
sions with respect to feeding and use of health care pertaining to children (Hajizadeh et al., 2016;
Martin, Rogers, Cook, and Joseph, 2004). For example, Latina mothers in the United States are more
likely to breastfeed their infants and follow American Academy of Pediatrics guidelines for feeding
than is the case for African American and European American mothers (Hurley, Knolhff, Dallav-
alle, and Black, 2005). In Nigeria, there is a reticence to give food to children who do not directly
contribute to family survival for fear that children may become spoiled (Zeitlin, 1996). Likewise,
in Nicaragua, where the belief is prevalent that children will eat when they are hungry, mothers
tend not to offer extra food to their young children, even in times of food shortage (Engle, Zeitlin,
Medrano, and Garcia, 1996).
A major contributor to child neglect is not having access to a strong support network and the
skills to know how to use it (Burke, Chandy, Dannerbeck, and Watt, 1998). Lack of social connec-
tions can lead to parental stress, depression, and lack of focus. Accordingly, perhaps nothing impacts
parental behavior pertaining to nutrition and health care than homelessness. According to Hart-
Shegos (1999, p. 2), “Homelessness influences every facet of a child’s life”. It is no wonder that many
children born to homeless mothers are born low-birthweight and experience numerous health
problems so long as they remain homeless.
Safety
Helping children avoid injury or abduction is considered a primary obligation of parents. Almost
every parent can recall specific events or circumstances that left them breathless due to their poten-
tial to cause bodily harm to their child. Injury rates for children, like mortality rates for children,
tend to be higher in rural areas (Cherry, Huggins, and Gilmore, 2007; U.S. Department of Health
and Human Services, Health Resources and Services Administration, Maternal, and Child Health
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Robert H. Bradley
Bureau, 2013). A good example the rural versus urban differences in injury is traumatic head injuries
(Halldorsson, Flekkoy, Gudmundsson, Aarnkelsson, and Arnarson, 2007; Leonhard, Wright, Fu, Lehr-
feld, and Carlson, 2015). Young children living in rural areas are more likely than children in urban
areas to be injured and die from burns, drowning, and motor vehicle accidents (Hwang, Stallones, and
Keefe, 1997). Part of the difference likely reflects generally lower wealth and education among rural
parents and part likely reflects differences in cultural values, the latter leading to different expecta-
tions pertaining to monitoring (see Surveillance) of children’s whereabouts. Many rural households
and adjacent areas contain objects and physical conditions that create risks for injury (Salmon et al.,
2013). To some degree, blighted urban neighborhoods also pose greater risk for injury. As children
get older and are allowed to wander (unsupervised) into the neighborhood, injuries are more likely
(Bradley, Corwyn, McAdoo and Garcia Coll, 2001; Dercon and Krishnan, 2009). Natural physical
hazards can affect what parents do in relation to their children. To reduce some of the risks, Ache
parents living in Paraguay put limits on infant locomotion and exploratory behavior (Kaplan and
Dove, 1987).
Low-SES children are more likely to suffer injuries and to die while still a child (Howe, Huttly,
and Abramsky, 2006; Keall, Baker, Ormandy, and Baker, 2011; Overpeck, Brenner, Trumble, Trifi-
letti, and Berendes, 1998), as their homes often have safety hazards (e.g., water heaters set too high in
temperature) and lack safety protections (e.g., smoke alarms; Bradley, Corwyn, McAdoo, et al., 2001;
Gielen et al., 2012; Evans, 2006; Gauderman et al., 2004). If parents are depressed, whether from
stresses associated with low SES or not, they are less likely to use car seats, safety latches, or covers
for outlets (McLearn, Minkovitz, Strobino, Marks, and Hou, 2005; Prickett and Augustine, 2016).
Residential instability can also make it challenging for parents to address potential safety hazards
(Cohen and Wardrip, 2011).
Another factor that can contribute to decreased effectiveness in assuring child safety is household
instability and chaos, whether from residential moves, job loss or change, the introduction of new
household members, traumatic events, military deployment, loss of child care, or other circumstances
(Wachs, 2000). Such conditions can distract both parents and children and can lead to reduced man-
agement of ordinary activities, miscommunications, and poor monitoring of child activities (Sand-
strom and Huerta, 2013).
Stimulation
To ensure competence and continued effort toward life-enhancing goals, the environment must pro-
vide sensory data that engage attention and provide information (i.e., stimulation; Kagan, 1984). In
their penetrating analysis of how children learn, the Committee on Developments in the Science of
Learning showed that people learn by actively encountering objects, actions, events, and concepts in
their environments (Bransford, Brown, and Cocking, 2000). There is substantial evidence that expo-
sure to key forms of stimulation promotes cognitive, psychomotor, and social development (Bradley
et al., 1994; Horowitz, 1987). Illustrative are studies of parent talk. The research of Hart and Risley
(1995) and Hoff (2003) provide compelling documentation of the value of providing children with
more labels for objects, responding contingently to children’s speech, making efforts to elicit conver-
sation from children, sustaining conversations with children, and just talking to them more often. In
general, to move from novice learner to expert in any particular area of knowledge or skill requires
substantial experience (usually guided experience) in that area (meaning much stimulation). How-
ever, social and physical conditions can sometimes make such actions on the part of parents more
difficult. For example, Evans and his colleagues (1999) found that adults living in crowded conditions
tend to speak in less complex ways to their children than do those who live in more spacious condi-
tions. These researchers suggest that coping with stress may contribute to reduced levels and reduced
quality of stimulation by parents.
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Access to social and material resources at the household, neighborhood, and community level
helps to determine what parents do to provide stimulation for children. Many household amenities
that support health and education require electricity, an obvious one being access to the internet and
use of computers (Hollingsworth, Mansaray, Allen, and Rose, 2011). Recent estimates suggest that
1.3 billion people lack predictable access, with rural areas in developing countries being the most
seriously affected (International Energy Agency, 2017). Crowded conditions can also affect the like-
lihood children will have access to a diversity of play and learning materials (Bradley and Caldwell,
1984), as crowded conditions make it difficult to store such materials.
Having access to a variety of material increases the likelihood children will develop a diverse
array of interests and skills, partly as a consequence of the types of social interactions certain mate-
rials facilitate (Bradley, 2015). For example, a survey done in 43 countries showed that number of
books in the home was positively correlated to reading comprehension in 15-year-olds (Chiu and
McBride-Chang, 2006). Gibson (1988) made the point that man-made and natural objects afford
opportunities for exploration and that such exploration leads to changes in perception, action pat-
terns, and cognition. Access to a diverse array of toys, learning materials, and household objects also
increases the likelihood of diverse types of interactions between parents and children, interactions
that lead to useful scaffolding of learning experiences for children (Tomopoulos et al., 2006). That
is not to say that having more objects and materials available at home only increases the likelihood
that parents will have more socially productive and competence-enhancing encounters with their
children. For example, having a TV and other types of electronic media available to children some-
times allows parents to spend time in activities that do not involve interactions with their children.
Research shows that most of the time even preschool children spend watching TV, using computers,
and playing video games is done independent of their parents (Takeuchi, 2011). Having cell phones
and fire arms in the house also may increase the likelihood parents will have to supervise children’s
behavior more closely and set down rules for their behavior (O’Keeffe, Clarke-Pearson, and Council
on Communications and Media, 2011). However, teens having a cellphone also increases the likeli-
hood that they will communicate with parents, particularly when they are not in the presence of the
parent (Lenhart, Ling, Campbell, and Purcell, 2010).
The factor that most strongly determines whether a child is likely to have access to a diverse array
of materials for learning and enjoyment—and consequently the likelihood that parents will spend
time with children using such materials—is household income (Hamadani et al., 2014). Parental
education and family income help to determine whether children will have access to musical instru-
ments, art supplies, and certain types of sports equipment (Bradley, Corwyn, McAdoo, et al. 2001,
2000; Yeung, Linver, and Brooks-Gunn, 2002). According to data collected by the Pew Research
Center (2015), household income is a major factor in determining whether children will participate
in sports, do volunteer work, take music, art, or dance lessons, and participate in organizations like
the scouts. Numerous studies world-wide show disparities between the “haves” and “have nots” with
respect to the presence of play and learning materials and the likelihood parents will read to children,
speak to them using complex language, and engage them in other types of encounters that promote
learning (Biedinger, 2011; Bradley, Corwyn, McAdoo, et al., 2001; Campbell and Parcel, 2010; Engle
et al., 2011; Hoff, 2003; Rubio-Dodina, Attanasio, and Grantham-McGregor, 2016; Sansour et al.,
2011; UNICEF, 2016). These differences, favoring those with greater resources, is consistent with
the family investment model (Conger and Donnellan, 2007). In some low-SES households direct
stimulation from parents in the form of meaningful conversations, guided interactions with materi-
als, and trips to potentially enriching facilities is diminished as a consequence of parental depression
or addiction (Kiernan and Huerta, 2008; Noll, Zucker, Fitzgerald, and Curtis, 1992). However, it is
difficult to fully assess how household income is implicated in the amount of stimulation children
will be afforded at home because income tends to be confounded with a number of other factors
that also affect the quality of home stimulation, such as single-parent status, maternal education, and
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immigration status (Biedinger, 2011; Campbell and Parcel, 2010; Green et al., 2009; Jackson, Brooks-
Gunn, Huang, and Glassman, 2000; Koury and Vortruba-Drzal, 2014; Sansour et al., 2011). It is also
the case that the availability of household resources is partially determined by the overall level of
social and physical resources available in the larger society (Bradley and Putnick, 2012).
The nature of parental employment also affects the kinds of stimulation parents provide their chil-
dren. Parents who have jobs with greater substantive complexity and self-direction tend to value ide-
ational flexibility and put less stress on conformity for family members (Schooler, 1999). They tend
to engage in more intellectually demanding leisure activities and discourse with their children. They
also take a more open-ended approach to problem solving and promote a self-directed orientation.
It is more common that single parents will have fewer financial resources than households with
two parents present. However, the burdens of being a single parent tend to diminish what the single
parent is able to afford children by way of nurturance and stimulation, even beyond what is attribut-
able to income (Bradley, Elardo, Rosenthal, and Friend, 1984; Crosnoe et al., 2010; Evans, Maxwell,
and Hart, 1999; MacKinnon, Brody, and Stoneman, 1982). In the study by Vortruba-Drzal (2003),
having more children in the household was also related to lower levels of stimulation. Such a finding
speaks to concerns that parents generally provide lower levels of investment for children who are
born later or who have lots of siblings. At present, however, the research is not consistent on how
family size or birth order affects the type or amount of stimulation parents offer children independ-
ent of other factors (Davis-Kean, 2005).
Area of residence is also implicated in what kinds of stimulation parents tend to afford offspring.
Natural landscaping encourages greater use of outdoor areas (Coley, Sullivan, and Evans, 1997; Kuo,
Bacaicoa, and Sullivan, 1998). Access to parks and recreational facilities not only increases the likeli-
hood that children will be afforded more opportunities for physical activity but it also increases the
likelihood they will engage in activities that increase psychomotor skills, including joint activity with
parents (Eime, Charity, Harvey, and Payne, 2015). Children with limited access to outdoor play areas
by their houses tend to be more fearful and more attached to their parent. By contrast, living in dense,
inner-city neighborhoods, particularly with high concentrations of poor families can discourage out-
door activities (Brockman, Jago, and Fox, 2011; Tigges, Browne, and Green, 1998). Residents in such
communities are often socially isolated and report higher levels of incivilities from others (Reisig
and Cancino, 2004). When children living in such areas go outside, they more often go with their
parents and the parents tend to be overprotective due to worries about safety hazards in the nearby
environment (Bartlett, 1997). Where families live can affect other parental efforts to provide other
types of stimulating experiences as well; most specifically, whether parents provide lessons to support
artistic and athletic interests. Although household income and parent education have a significant
role in determining whether children will be involved in music, sports, and art activities, proximity
to facilities and instructors who provide lessons also plays a role in whether a child is given private
lessons (Child Trends, 2015; Corrigall and Schellenberg, 2015). To offer another illustration, Dur-
brow, Jones, Bozoky, Jimerson, and Adam (1996) found that children from the islands of Jamaica and
St. Vincent rarely had access to personal musical instruments nor did they take many trips of greater
than 50 miles.
Sometimes living in a crowded, inner city area means that families have to cope with high levels
of noise. Excessive noise can make it very difficult for parents to maintain focus and offer productive
stimulation or socioemotional support to children on regular basis (Evans, 2006). Noise, and other
forms of household chaos, tend to produce stress in both parents and children, stress that can lead to
reduced engagement in joint activity and exchanges that are not very productive (Evans and Wachs,
2010). Crowded conditions can also lead to conflict between parents, and inter-parental conflict
reduces the quality of stimulation offered children (Cabrera, Hofferth, and Soo, 2011).
Based on his analysis of the parenting literature, Bornstein (2013) concluded that culture helps
shape parental cognitions, which help shape parenting practices. Roopnarine and Davidson (2015)
482
offer several illustrations of how culture influences parenting play behavior with children, noting
differences in the percentage of mothers who act as playmates for young children in Guatemala,
India, and the United States and noting differences in the times mothers versus fathers engaged
in play with young children in various countries. Vandermaas-Peeler (2002) juxtaposed the ideas
that children in all countries tend to play in somewhat similar ways with various types of materials
(e.g., balls, dolls, household objects) and they often do so in the presence of parents; but parents in
different cultures construe their roles as playmates quite differently. Parents living in poor countries,
where survival remains a key issue, spend less time in imaginative play with children or guiding their
skill development in ways that connect to academic learning. Indeed, this role is often turned over to
siblings. Even when parents spend time in joint activity with children, culture often determines the
goals of those activities, with Asian parents often more concerned with teaching proper conduct and
U.S. American and European parents more often concerned with knowledge advancement or just
having fun (Haight, Parke, and Black, 1997; Tamis-LeMonda, Bornstein, Cyphers, Toda, and Ogino,
1992). Even within the same country, cultural differences can be seen in how often parents perform
particular tasks related to their children’s learning (Bradley et al., 2000, Bradley, Corwyn, McAdoo,
et al., 2001); and even within a society where one ethnicity is dominant there can be geographic
differences in how intensively parents are involved in stimulating children’s learning—likely a con-
sequence of movement toward more modern views of life in areas where adult work circumstances
have become more technological (Li and Rao, 2000).
The findings reported by Li and Rao (2000) pertaining to differences between parent behavior
in Beijing, Singapore, and Shanghai point to some of the complexities in trying to determine how
various social and physical factors combine to determine what parents do by way of providing stimu-
lation for their children (Bornstein, 2016). Consider too a cross-cultural comparison where both
cultures would be considered “modern” in their views toward parenting. Specifically, compared with
U.S. American parents, Israeli parents are less likely to include children in their recreational hobbies
and less likely to spend time teaching their children motor skills, the latter partly owing to the fact
that most such skills are learned in the neighborhood from peers. Because of working conditions and
religious practices, Israeli children are also less likely to be taken to the theater or shopping ( Jacobson
et al., 1995). Israeli households tend to be smaller than American households; thus, there tend to be
fewer material possessions aimed at children present in the home.
Support
To be socially competent and emotionally healthy, it is critical that children receive adequate soci-
oemotional support from caregivers and live in environments that afford them protection from
forces that would undermine adaptive functioning (Egger and Angold, 2006). As emotional beings,
we need environments that are both affirming and responsive to our individual needs (Brether-
ton and Waters, 1985; Rohner, Khaleque, and Cournoyer, 2005). Emotions function to prepare
human beings to take action in their own best interest (Cohn and Fredrickson, 2006). Parents must
assist children in enlisting and modulating the motivational properties of emotions to help ensure
optimal fit with environmental demands. There is substantial evidence that development is more
nearly optimal in an environment that is responsive in the sense that a child’s requests for assistance
are dealt with in a timely, predictable, and satisfying way (Ainsworth, 1973; Bowlby, 1969; Rot-
ter, Chance, and Phares, 1972). Emotional health is also fostered when caregivers can accurately
anticipate children’s emotional needs for particular situations and proactively furnish needed social
or cognitive supports and when they engage in behaviors that affirm children’s identity and worth
(Rohner et al., 2005). Such care promotes goal-directed (positively motivated) behavior on the part
of children. By contrast, children’s well-being is undermined when caregivers act in hostile, overly
intrusive, and disrespectful ways, as this undermines a child’s needs for autonomy and connectedness
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( Joussemet, Landry, and Koestner, 2008). These negative forms of parental behavior can lead to poor
social relationships and involvement with antisocial peers (Dishion, Patterson, Stoolmiller, and Skin-
ner, 1991; Power and Chapieski, 1986).
To fully appreciate how social and physical aspects of the environment affect parental efforts to
provide socioemotional support to children, it is important to understand that there are cultural vari-
ations in how adults are likely to express affection, affirm worth, or act in a responsive way toward
their children (Chao and Kanatsu, 2008; Kilbride and Kilbride, 1983; Whiting and Edwards, 1988).
Cultural differences in these parenting behaviors emerge as a consequence of how culture shapes
parents’ interpretation or children’s behavior and needs (Bornstein, 2013). Cultures subscribing to
an interdependence script rely more on continuous caregiving and parent-child contact (Carlson
and Harwood, 2003). Kung infants from Namibia spend many hours a day in direct contact, as do
Longoli children in Kenya (Munroe and Munroe, 1980). Likewise, Indian mothers tend to stay close
to their infants and score high on emotional responsiveness (Agrawal and Gulati, 2005). Between-
country differences in parental affective responses are also seen for older children. For example, Ken-
yan parents rarely encourage school-age offspring to contribute to conversations with visiting adults,
a finding that contrasts with U.S. and European parents (Holding, 2003).
Body contact provides an interesting window through which to understand how culture can
influence parental actions that offer socioemotional support to children. Keller and colleagues (2004)
studied mothers from the Cameroon, Greece, Germany, and Costa Rica. Specifically, they observed
mothers with their infants in a free-play situation. Almost all the mothers (a small number of Greek
mothers being the exception) used some form of body contact during at least part of the observation
period. However, mothers from the Cameroon and Costa Rica (where interdependence in social
relationships is the norm) maintained that contact for at least 75% of the observation period, whereas
Greek and German mothers did so less than 33% of the time. Likewise, almost all mothers had face-
to-face contact with their children during the period; but Greek and German mothers maintained
such contact for about 70% of the time (consistent with a strong focus on education for self-reliance)
compared to less than half the time for the other three groups of mothers. In the three groups (Ger-
man, Greek, Costa Rican) where great value is ascribed to economic independence, mutual eye
contact was maintained throughout almost half the observation period; contrasted to less than 20% of
the time for the groups from Cameroon. When Richman and colleagues (1988) compared the Gusii
mothers from Kenya to mothers from Boston, they found that Gusii mothers rarely talked to their
infants or even looked at them. By contrast, Gusii mothers held infants more than Boston mothers.
In societies where self-reliance and autonomy are highly valued, parents tend to demand less compli-
ance with parental directives and are more tolerant as regards displays of disrespect for elders. Parents
also tend to use physical forms of child control less frequently. For example, in Bahrain, where great
value is placed on parental authority, scores on the Acceptance dimension from the HOME were
substantially lower than those generally obtained in studies done in the United States, Canada, and
Western Europe (Hadeed and Sylva, 1999). The Acceptance dimension includes such items as “par-
ent does not scold, yell at or derogate child more than once during visit”, “parent does not use physi-
cal restraint during visit”, and “parent neither slaps nor spanks child during visit”. These findings, and
others like them, suggest that cultural values help determine the manner and level of socioemotional
support offered to children. That said, there are often socioeconomic differences in the families as
well as cultural differences, leaving unclear just how strongly cultural values are implicated in these
parenting behaviors independent of differences in material and social resources.
At present, there is only a modest amount of information on how often parents offer most types
of support to their children. As a consequence, not much is known with precision regarding how
various aspects of the social and physical environment change the rate at which parents provide
supportive actions and conditions for their children. One interesting exception was a study done
in Spain. In this study, the researchers found that the likelihood a parent would touch a child in a
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positive way depended on the nature of the task they were engaged in. Fathers were more likely to
softly touch children during a storybook reading task and mothers were more likely to softly touch
a child while reminiscing (Anzar and Tenebaum, 2016). The main reason so little is known about
factors that influence particular types of socioemotional behavior on the part of parents is that it is
difficult to perform the kind of exhausting documentation of supportive behaviors such as Hart and
Risley (1995) and Hoff (2003) accomplished with parental language inputs to infants and toddlers.
Some direct observations of parent-child interactions are included in broad home environment
measures such as the HOME Inventory (Caldwell and Bradley, 2016). Unfortunately, researchers
rarely report findings at the item level, so it is difficult to estimate how often parents engage in
particular actions, much less what determines the likelihood they will. One exception was a study
of 10- to 15-year-old children by Bradley, Corwyn, Caldwell, Whiteside-Mansell, Wasserman, and
Mink (2000). During an hour-long home visit, fewer than 6% of parents (representing five different
ethnic groups) made derogatory comments about the adolescent, and almost all (97%) maintained
a positive demeanor when speaking to the adolescent. Over 80% of the parents encouraged the
adolescent to contribute to the conversation and responded appropriately to the adolescent’s com-
ments and questions. In a study done with 16- to 20-year-olds from diverse ethnic backgrounds and
geographic locations, Bradley and colleagues (2017) found that more than 93% of parents asked the
adolescent’s opinion regarding family activities and showed positive feelings toward the adolescent
during the home visit. By contrast more than 30% of parents did not try to work to resolve disagree-
ments with the adolescent.
Not surprisingly, there is abundant documentation of the link between low family income (or
low SES) and less nurturant, more hostile interactions between parents and children (Blair and Raver,
2012; European Child Care and Education Study Group, 1999; Gunning et al., 2004). A number of
scholars point to the stresses connected to poverty as a major mediator of this connection (Conger,
Conger, and Martin, 2010; Mistry, Lowe, Benner, and Chien, 2008). The stresses connected to low
parental education and financial hardship have been linked to maternal distress and low self-efficacy
which in turn leads to neglectful, insensitive, and even hostile behavior on the part of parents (Brody,
Stoneman, Flor, McCrary, Hastings, and Conyers, 1994; Brody, Flor, and Gibson, 1999; Conger et al.,
2010; DeGarmo, Forgatch, and Martinez, 1999). Household instability and overall chaos is more
common in low-SES families too, with research showing that high levels of confusion in the house
decrease maternal responsiveness (Corapci and Wachs, 2002). Ackerman, Kogos, Youngstrom, Schoff,
and Izard (1999) reported that family instability led to increased conflict among family members and
promoted negative emotionality on the part of parents. Research generally shows that household
chaos increases the likelihood parents will display negative behavior toward children and often leads
to inconsistent and ineffective disciplinary practices (Coldwell, Pike, and Dunn, 2006; Dumas, Niss-
ley, Nordstrom, Smith, Prinz, and Levine, 2005). Stress connected with adversity also decreases the
likelihood of positive forms of parenting (warmth, sensitivity, encouragement, stimulation; Bradley,
Corwyn, McAdoo, et al., 2001; Conger and Donnellan, 2007; Grant et al., 2003; Gutman, Sameroff,
and Cole, 2003; Kiernan and Huerta, 2008), which can make it harder for children to control their
emotions and inhibit undesirable behavior (Lansford and Deater-Deckard, 2012; McLoyd, 1998).
It is not always easy to determine how much family income, parental education, and occupa-
tional status feed into parental actions. Low-SES parents are more likely to manifest mental illness or
addiction problems, which also decreases the likelihood they will be emotionally responsive to their
children (Howard, Kumar, and Thornicroft, 2001; Wilson and Durbin, 2010). Eiden, Edwards, and
Leonard (2007) found that paternal alcoholism predicted low warmth and sensitivity for both fathers
and mothers, which in turn predicted poor self-regulation and behavior problems in their offspring.
When depression accompanies divorce or other forms of instability, there is a tendency of mothers
to withdraw from their children, leading to more behavioral maladjustment in the children (Wood,
Repetti, and Roesch, 2004).
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Robert H. Bradley
Low SES and divorce often mean that a parent lacks support from others when enacting the role of
parent. Unfortunately, the construct of social support is a bit “fuzzy” in what it represents. It is not in
conceived or measured in a consistent way in studies of human behavior (Geens and Vandenbroeck,
2012); nonetheless, it is presumed to assist individuals in engaging in productive forms of behavior,
particularly when faced with challenges. Ceballo and McLoyd (2002) found that having support
from others reduced the potentially negative impact of living in a disadvantaged neighborhood on
positive parenting. MacKinnon et al. (1982) and Bradley et al. (1984) found that single-parent house-
holds, family situations where mothers often are faced with handling the job of parenting by them-
selves, scored lower in acceptance and responsiveness on the HOME Inventory than did two-parent
households, even with other family demographic factors controlled. Likewise, Black, Dubowitz, and
Starr (1999) found that the presence of a father in the home was correlated with the total HOME
Inventory score in African American families with 3-year-old toddlers, with maternal education
and age controlled. Mexican American adolescents reported greater parent-child conflict when liv-
ing with a single-parent mother than when living in a two-parent household (Zeiders, Roosa, and
Tein, 2011). When Dodge, Pettit, and Bates (1994) included both low SES and single parenthood as
predictors of children’s externalizing problems and aggression, a socialization composite that included
harsh discipline, parental warmth, cognitive stimulation, and exposure to violence indirectly linked
both household SES and single parenthood to child outcomes. Other studies have shown the paternal
presence is associated with greater maternal responsiveness and less frequent use of harsh punishment
(Bradley et al., 1984, 2000). However, the researchers did not control for other demographic factors.
Moreover, Cain and Combs-Orme (2005) found limited evidence that family structure, by itself, was
a major factor in determining how much warmth mothers showed to children. There have been
relatively few studies, beyond those pertaining to paternal presence or marital status, that examine the
relation between family configuration and the quality of parental nurturance.
Research on three-generation households offers additional evidence of how living arrangements
can influence parenting behavior. At the same time, the findings also make clear how complex
such arrangements can be and how those arrangements might function to determine parenting
actions, especially actions that are supportive of children’s social and emotional well-being (Oyser-
man, Radin, and Salt, 1994; Spieker and Bensley, 1994; Whiteside-Mansell, Pope, and Bradley, 1996).
In general, when young mothers and grandmothers have a high-quality relationship, it bodes well
for the mothers’ parenting and feelings about father involvement if fathers are also present in the
household (Krishnakumar and Black, 2003). However, if mothers do not perceive their parents as
being truly supportive (i.e., grandparents have many expectations for the mother leading to con-
flict between them), then the mothers may find it more difficult to engage in nurturant parenting
(Barnett, Mills-Koonce, Gustafsson, Cox, and the Family Life Project Investigators, 2012 Oyserman
et al., 1994).
There is anecdotal evidence that the presence of siblings can have effects on what parents do, but
only a few studies. In one such study, Shanahan, McHale, Osgood, and Crouter (2007) found that
parents often display greater warmth to later-born children. In another example, Harris and Morgan
(1991) found that girls who had more brothers got more attention from their fathers. The strain of
dealing with sibling conflict can lead to harsher, more restrictive parenting (Crnic and Greenberg,
1990), albeit mothers and fathers tend to respond differently to squabbles between siblings. Mothers
tend to try coaching more than fathers do, whereas fathers tend to directly intervene more often
(McHale, Updegraff, Tucker, and Crouter, 2000). Finally, when a sibling has a severe disability or
chronic illness, it often changes how the parent acts toward the child, including the amount of time
spent in supportive interactions with the child (Rossiter and Sharpe, 2001).
Spousal support is associated with greater involvement and more nurturant care in both moth-
ers (Cummings, 1994; Olsen, Martin, and Halverson, 1999) and fathers (Coley and Chase-Lansdale,
1999; NICHD Early Child Care Research Network, 2000). The findings pertaining to social support
486
are also consistent with findings on coparenting relationships. Specifically, Belsky, Putnam, and Crnic
(1997) found that unsupportive coparenting was associated with less responsive parenting on the part
of both mothers and fathers. By contrast, inter-parental conflict increases the likelihood that parents
will act in harsh and punitive ways towards children, what is sometimes called a “spillover” effect
(Krishnakumar and Buehler, 2000). Low-SES parents more often have a history of trauma during
their own childhood or have been abused by a partner. These traumatic experiences are connected to
the use of severe physical discipline or neglect as regards their own children (Banyard, Williams, and
Siegel, 2003). In households where mothers and grandmothers live together, verbal conflict between
the two is predictive of the mother’s intrusive behavior with her own children (Barnett et al., 2012).
The physical conditions present in a home can have an impact on how supportive parents are to
children (Evans, 2006). Crowded households often lead to tensions between family members (Wachs
and Camli, 1991). Evans and Lepore (1993) posited three major mechanisms that can adversely affect
behavior in crowded situations: (1) behavioral constraint (density interferes with goal attainment and
thereby causes frustration), (2) diminished control (density exposes people to situations over which
they have little perceived control, leading to learned helplessness), and (3) overload or arousal (exces-
sive stimulation from density overloads the sensory system). Although the research to support each
of these mechanisms is limited in terms of precise applicability to parenting behaviors, the broader
literature on stress reactions suggests that high density in the household could lead to both intrusive,
hostile actions and to social withdrawal on the part of parents. Supportive of that hypothesis is a
study by Bradley and Caldwell (1984). They found that household crowding was related to paren-
tal acceptance (lack of restrictiveness and punishment) of infants, even when they controlled for
other family demographic factors. Bradley and Caldwell (1984) also reported a negative correlation
between crowding and parental warmth for preschool-age children, albeit they did not control for
other family demographic variables. Likewise, Bradley et al. (2000) found that household crowding
was related to parental acceptance and responsivity for teenagers, but they did not control for other
demographic factors. In the study of adolescents, there was not a significant relation for African
Americans.
Living in a dangerous or run-down neighborhood can contribute to parenting stress and reduce
parental expressions of warmth and increase punitiveness, but the findings are mixed, with some evi-
dence that the influence is greater for low-income families and immigrant families with low levels
of acculturation (Barnett, Mortensen, Gonzalez, and Gonzalez, 2016; Cueller, Jones, and Sterrett,
2015; Kohen, Leventhal, Dahinten, and McIntosh, 2008). This is particularly the case if families have
been exposed to community violence (Aisenberg and Ell, 2005). In such situations, both parents and
children are likely to become more anxious, sometimes feeding off each other’s insecurities. The pre-
sumed negative impact of living in a bad neighborhood seems to be partially offset if one has social
support (Ceballo and McLoyd, 2002). The negative impact of neighborhood disadvantage also did
not emerge in a study of Latino families (Gonzales et al., 2011) or a study of middle-income families
(Vieno, Nation, Perkins, Pastore, and Santinello, 2010). These latter findings suggest complexities in
how external conditions impact the tendency of parents to engage in positive caregiving.
Among the least controversial are findings are those pertaining to experiences with war and
trauma. As illustration, a study done in Gaza showed that there was less reciprocity between mothers
and infants who were exposed to war conditions (Feldman, Vengrober, Eidelman-Rothman, and
Zagoory-Sharon, 2013). Likewise, a study of mothers who experienced intimate partner violence
showed that such an experience increased the likelihood the mothers would neglect their children
(Lannert et al., 2014).
An emerging area of concern with respect to parenting and parent-child relationships is the rapid
infusion of technology in family life (Villegas, 2013). At this point, research is simply inadequate to
speak to the nuances of how use of cell phones, TV, and various other computer devices changes
interactions between family members. That said, there are worries that TV and other electronic
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devices essentially function as “baby sitters” for young children and that both parent and child are
often distracted while they are using such devices, leaving less opportunity for positive encounters
with other family members (Radesky et al., 2014). Advances in technologies and the rapid uptake
of such devices makes it difficult for scholars to catalog just how access to such devices is impacting
the quality of parental behavior; but this is an area in need of considerable research (Radesky, Schu-
macher, and Zuckerman, 2015).
Structure
To promote optimal development for children is challenging given all the demands, restrictions, and
opportunities afforded by the settings people inhabit through time—not to mention the variability
in children’s behavior. Zolkoski and Bullock (2012) found that children fare better when parents
were able to foster a continuing sense of family cohesion and a general sense that there is still coher-
ence and manageability in life. In effect, parents must not only assure that children remain safe but
also assure that the children get the right amounts and kinds of stimulation, sustenance, and sup-
port. In essence, parents must organize how and when various inputs get to the child and organize
time, space, objects, and events that occur in settings so that there is a “fit” with a child’s needs and
proclivities and what the environment affords—they must structure the child’s encounters with the
environment (Spagnola and Fiese, 2007; Walsh, Craik, and Price, 2000). When families are homeless,
it is almost impossible to provide the kinds of organization needed to assure children’s overall well-
being (Hart-Shegos, 1999). Likewise, family trauma may not just disrupt various ways of organizing
family life but also lead to adjustments in the organization of various tasks and physical arrangements
designed to adapt to the trauma (Gerwitz, Forgatch, and Wieling, 2008). Structuring the activities
of daily life, providing meaningful routines for children—even maintaining a useful organization of
furnishings and materials in the house—can be very difficult if parents themselves are traumatized
and families are dislocated (Banyard, Williams, and Siegel, 2003). Even rather commonplace changes
in family life (moving to a new neighborhood, birth of a sibling, military deployment, change of jobs,
divorce, joining a sport team) can lead to major shifts in family routines or physical arrangements
within the household (Mahoney, Harris, and Eccles, 2006; Mayberry, Shinn, Benton, and Wise, 2014;
Roche and Ghazarian, 2012).
Both parents and children can function more productively within a setting when the conditions
within the setting are orderly and transparent. In an orderly setting, it is easier to get one’s bearings
and make plans for how to meet setting demands and take advantage of the opportunities present
within the setting. To some extent, the tenets of dynamic systems theory seem particularly relevant
when evaluating how various changes or “instabilities” in family circumstances can change the kinds
of structures parents use in their role as parents (Endsley, 1995). According to the theory, the actions
of systems are organized around various “attractors” (physical and social elements) that give direc-
tion to the elements (Thelen and Smith, 2006). When the system is disrupted, those elements may
get re-organized around new attractors. In complex, adaptive systems, like human families, it is not
always easy to predict what new arrangements will be made by parents when disruptions in family
life occur or how stable the new arrangements will be. Part of the challenge in predicting derives
from the fact that changes in one aspect of family life (e.g., changes in family composition) can lead
to changes in other aspects of a child’s life (e.g., day care arrangements; Crosnoe, Prickett, Smith, and
Cavanagh, 2014). That said, Martin, Razza, and Brooks-Gunn (2012) found that household instabil-
ity was associated with fewer productive family routines.
There is evidence that the number of objects or ideas a person can deal with effectively at any
one point in time increases with age (Kuhn, 1992); thus, children (particularly young or otherwise
less competent children) can benefit when a parent (or teacher) organizes a child’s encounters with
objects and ideas so that they are easier to assimilate (Bjorklund, 1990). Learning is also easier in
488
the absence of distracting stimuli (Wohlwill and Heft, 1977). There is an emerging literature on
how best to guide or “scaffold” children’s learning so that learning is more efficient and productive
(Blewitt, Rkump, Shealy, and Cook, 2009; Pea, 2004). The emergence of new forms of media and
electronic devices has motivated parents of young children to actively mediate children’s engagement
with the devices, to enhance both enjoyment and learning (Nikken and Schols, 2015). A number
of environmental factors function to affect maternal effectiveness in scaffolding children’s learning
and problem solving skills, including maternal education, household income, single-parent status,
and depression (Hoffman, Crnic, and Baker, 2006; Lowe, Erickson, MacLean, Schrader, and Fuller,
2013; Stright, Herr, and Neitzel, 2009). In addition, children’s characteristics influence the type and
amount of scaffolding offered (Evans, Moretti, Shaw, and Fox, 2003; Lowe et al., 2013; Robinson,
Burns, and Davis, 2009).
The rapid infusion of computer and media devices is also having an effect on family routines.
They are changing much about family dynamics and how parents manage common daily activities
(Villegas, 2013). Most critically, perhaps, they are changing the structure of interpersonal interactions
and how family members communicate (Misra, Cheng, Genevie, and Yuan, 2014; Przybylski and
Weinstein, 2012).
Crowded conditions in a residence often make it difficult for parents’ children to organize things
for themselves and to make maximum use of persons and objects in the environment (Reynolds,
2005). Unfortunately, parents tend to provide less supervision in crowded homes and may involve
themselves less intensively with their children (Bradley and Caldwell, 1984; Wachs, 2000). In general,
when conditions like crowding and stressful life events produce chaotic conditions in a household,
parents are less able to instill productive routines for their children, a circumstance that leads to stress
reactions and poor achievement (Chen, Cohen, and Miller, 2010; Muniz, Silver, and Stein, 2014).
However, Bradley et al. (2000) found only a small correlation between household crowding and fam-
ily efforts to regulate adolescent behavior. Somewhat by contrast, parental supervision is made con-
siderably more difficult in the case of “latchkey” children. It is more difficult to monitor or supervise
youth if the parent is not physically present (Belle, 1999; Mulhall, Stone, and Stone, 1996).
Parents use different approaches to supervision and impose different rules, depending on what
their neighborhood affords. In dilapidated neighborhoods, there were greater efforts on the part of
parents to make their own homes safe for a child (McDonnell, 2007). As mentioned earlier, because
of physical hazards present in their surroundings, Ache parents living in Eastern Paraguay tend to
restrict children’s locomotion and freedom to explore (Kaplan and Dove, 1987). Parental efforts to
organize their children’s activities in the neighborhood depend on the distinctiveness of features
in the area and the level of safety present. In an effort to protect their children from the physi-
cal danger and lure of gangs and drugs present in the inner city, parents use a variety of strategies,
including keeping children inside the home, walking them to the school bus or school, restricting
their children’s associations within the neighborhood, and enrolling them in after-school programs
(Halpern, 1992; Jarrett, 1997). Having distinctive, well-situated landmarks can enable children to ori-
ent themselves and to move throughout the environment with greater facility (Head, Bradley, and
Rock, 1990; Weinstein and David, 1987). The quality and resources available in the neighborhood
and community affect the kinds of regulatory strategies parents use and, ultimately, children’s social
competence (O’ Neil,Parke, and McDowell, 2001).
Structure is important for children’s socioemotional functioning and adaptive behavior. For exam-
ple, parents often serve as facilitators of interactions with peers for children, especially young children
(Bhavnagari and Parke, 1991). Likewise, parents often try to address the anxieties that can arise when
children encounter new circumstances, such as moving to a new house or day care arrangements
(Coddington, 1972; Sandstrom and Huerta, 2013). Even when changes are more normative, such
as when children are not in school, parents often have to re-organize how children will spend time
in daily activity. For example, parents often arrange for children to be more physically active on
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weekends and during the summer (Atkin, Sharp, Harrison, Brage, and van Sluijs, 2016). Data from
the Survey of Income and Program Population show that families often make major adjustments
in child care arrangements, sports participation, and lessons during the summer, depending on child
age, family composition, availability of relatives, ethnicity, and family income (Laughlin, 2010). As
intimated above, the likelihood parents will engage in actions that help children manage situations
and take advantage of the opportunities situations afford depends on the characteristics of the child.
Unfortunately, some child characteristics result in parents backing off their role as supervisor. This
appears to be especially the case as children get older and begin involvement in delinquent behavior
( Jang and Smith, 1997).
SES, family composition, and ethnicity are implicated in certain forms of structuring for children.
For example, high-SES parents tend to provide more effective scaffolding for their children’s learn-
ing (Sigel, 1982), and they are more likely to eat meals together as a family and impose limits on
TV viewing (Anderson and Whitaker, 2010; Bradley, Corwyn, McAdoo, et al., 2001; Prickett and
Augustine, 2016). Single-parent and low-income households are more often characterized by house-
hold disorganization, and children in such homes are less likely to be fully supervised (Anderson and
Whitaker, 2010; Cookston, 1999; Goldstein, 1984).
Surveillance
To keep children safe and healthy requires that parents must keep track of the child’s whereabouts
and activities. It also means being aware of conditions present in whatever setting the child is in.
A significant proportion of accidents and injuries occur because parents fail to adequately moni-
tor children, (Greaves, Glik, Kronenfeld, and Jackson, 1994; Morrongiello, Kane, and Zdzieborski,
2011). Clearly, the type and amount of monitoring and supervision children need to protect them
from harm change with age (Peterson, Ewigman, and Kivlahan, 1993). As children get older, there
is increased concern about sexual predators and exposure to violence on TV and other forms of
media (Lee, Bartolic, and Vandewater, 2009). Access to social media has only worsened the situation
for adolescents (Pew Research Center, 2016; Shin, 2015). Moreover, children’s engagement in risky
behavior is more likely if parents do not adequately monitor children’s activities and social networks
(Bradley and Corwyn, 2013; Dishion and McMahon, 1998; Dittus et al., 2015; Han et al., 2012;
Ryan, Roman, and Okwany, 2015). Research makes unclear just how monitoring fits into the equa-
tion of harm avoidance (including risky behavior) in older children (Kerr, Stattin, and Burk, 2010),
with evidence indicating that the tendency to monitor and monitoring effects may vary by culture
(Chao and Kanatsu, 2008; Ryan et al., 2015).
The social and physical affordances in a setting have a bearing on parental monitoring behavior,
together with the child’s personal characteristics. Each setting, because of the particular persons,
objects, and activities it contains, exerts different pressures on a parent to maintain visual and/or
physical contact with a child, pressures that vary according to the child’s age, competence, and pro-
clivities (Barker, 1978). Closer proximity is required at the state fair or in the parking lot at McDon-
alds than in a park in an up-scale neighborhood or in church. In chaotic resource-poor settings,
such as those in the Brazilian shantytowns described by Scheper-Hughes, monitoring a child can
be extraordinarily difficult. There is very little information on how particular environmental condi-
tions influence parental monitoring behavior, but there are clues from research on parental beliefs
on how long it is appropriate to leave children alone in different settings (Peterson et al., 1993). Not
surprisingly, parents recommended that children be supervised most closely in those situations that
pose the most imminent dangers, and they advocated less supervision as children get older. The study
did not assess how closely parents actually monitor children in these various situations. Furthermore,
there is concern that parental beliefs and intentions based on their appraisal of what settings afford
by way of potential harm only partially determines actual practice. For example, the National Survey
490
of American Attitudes on Substance Abuse included 12 questions on surveillance by parents for

adolescents to answer. These included questions about monitoring TV and internet use, restrictions
on what music they buy, knowing their whereabouts after school, expectations about academic per-
formance and being told the truth about where the adolescent planned to go, imposition of curfews,
assignment of chores, and eating dinner with the teen 6 or more times a week. Only 27% of teens
indicated that their parents consistently took 10 or more of the 12 actions. By contrast, 18% indicated
that their parents took five or fewer of the actions. Finally, part of the appraisal process derives from
parents’ beliefs regarding the ability of a child to deal with the affordances present in a given situation,
and parents often overestimate children’s capacity to understand the dangers present in a situation
or to exert control over their own behavior in situations that induce engagement (Karstad, Kvello,
Wichstrom, and Berg-Nielsen, 2014).
As stated earlier, the pervasiveness of media devices has made it more difficult for parents to effec-
tively monitor adolescent activities. The majority of parents report checking the websites their chil-
dren have visited and their social media profiles, and they report looking through phone call records
(Pew Research Center, 2016). Most have some restrictions on children’s use of computers, and about
40% of used parental controls on cellphone use (Shin, 2015). This said, many parents are somewhat
ambivalent as regards how much to monitor and control children’s use with media, and their patterns
vary depending on child age (Nikken and Schols, 2015; Pew Research Center, 2016). As it happens,
some electronic devices have also given parents additional means of engaging in monitoring. Not
only are there monitoring tools to track where children are if they have a phone; but, there are also
video devices that can help parents be aware of what is happening to children when the parent is not
physically present with them. A good example is baby monitors and other cameras that allow parents
to observe children when parents are not present (e.g., when baby is asleep in a crib or when a pre-
schooler is at home with a nanny). In effect, if parents and offspring both have smartphones, parents
can connect more easily with the child during times when they are not together.
Just as it is harder to supervise children who live in certain circumstances, so it is harder to moni-
tor them as well. If a child is at home alone, for example, monitoring is quite difficult (Belle, 1999;
Mulhall, Stone, and Stone, 1996). Somewhat by contrast, when families live in dangerous or dilapi-
dated neighborhoods, there seems to be a tendency for parents to monitor children more closely and
perhaps be more restrictive ( Jarrett, 1997; Jones, Forehand, O’Connell, Armistead, and Brody, 2005;
Letiecq and Koblinsky, 2004; O’Neil, Parke, and McDowell, 2001; Vieno, Nation, Perkins, Pastore,
and Santinello, 2010). However, there is not yet much evidence for strong neighborhood effects
(Cueller et al., 2015).
Part of what determines how much and what kind of monitoring parents do relates to whether
there are others available to keep track of the children. In families in which there are other kith and
kin to help a parent monitor children, the burden of monitoring particular children is distributed
across networks of adults and older children. In situations where there is only one parent in the
household and no other adults are present, research points a reduction in monitoring and a higher
risk of injury (Chao and Kanatsu, 2008; Rivara and Mueller, 1987).
As the process model of parenting promulgated by Belsky and Jaffee (2006) would suggest, a
major factor in determining whether a parent will monitor a child behavior in a given situation is
the child’s behavior and characteristics. As the text above implies, parents are to some extent more
inclined to monitor a child’s behavior if the child has previously engaged in some form of risky
behavior; but socially deviant behavior also motivates some parents to withdraw (Bradley and Cor-
wyn, 2013). What seems more clear is that parents are want to be more attentive to children who have
health problems. There is a quite remarkable literature on parental efforts to keep track of and attend
to technology-dependent children (Elias, Murphy, and the Council on Children with Disabilities,
2012; Meltzer, Sanchez-Orhuno, Edinger, and Avis, 2015; Wang and Barnard, 2004) or children with
other complex medical conditions (Rempel and Harrison, 2007; Uzark and Jones, 2003; Woodgate,
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Robert H. Bradley
Edwards, Rpiat, Borton, and Rempel, 2015). Such efforts to “be there” for fragile children induces
significant stress and often leads to sleep deprivation on the part of parents, especially if they have
no one else to help them in providing critical caregiving tasks. The impact of various forms of dis-
ability is less clear. In some cases, parents “up the ante” to make sure circumstances are favorable for a
disabled child; but some disabilities (e.g., attention deficit hyperactivity disorder) can wear on parents
and actually decrease their level of monitoring (Algood, Harris, and Hong, 2013; Hassall, Rose, and
McDonald, 2005; Ullsperger, Nigg, and Nikolas, 2016). It is also the case that some disabilities actu-
ally decrease the likelihood children will encounter some types of harm (e.g., engagement in risky
behavior with peers or solicitation on social media from potential predators); thus, parents may feel
they have less need to engage in certain types of monitoring (Hogan, Shandra, and Msall, 2007).
It is also likely that parents will become more attentive if they are aware of circumstances that lead
to physical or emotional harm for their children, such as bullying, suicidal ideation, anorexia, and
self-inflicted wounds. However, research on such actions is quite limited and it suggests that parents
sometimes become quite cautious when dealing with such issues (Buus and Steager, 2013; Honey
and Halse, 2005; Sweet and Whitlock, 2009).
Not surprisingly, most of the research on parental monitoring pertains to efforts designed to
reduce harm. However, parents also pay attention to the affordances of settings and their children’s
engagement with objects and people in them to determine how well suited the conditions are for
promoting productive and enjoyable encounters as well. There is a literature that advises parents on
how to make situations more productive and enjoyable for children, but there is little research on
what family members actually do for children in most situations or what motivates them to do so.
Social Integration
For children to succeed in any society, it is important that parents connect them with the social
fabric of the society as it is through such connections that children receive the opportunities and
supports they need to live productively (Weisner, 2002). Connecting to extended social networks
can be particularly important for children living in adverse circumstances, such as poverty or family
instability (Horvat, Weininger, and Lareau, 2003). It is through connections with social networks
(e.g., extended family, other adults in the community, work associates, community agencies) that
parents can access potential resources for children (Runyan et al., 1998). Single-parent homes gener-
ally have less access to extended family and the resources they can provide; and, as a consequence,
children’s well-being tends to be compromised (Amato, 1995). As it happens, parents with mental
illness or problems with addiction are often in difficult positions to connect their children to use-
ful social networks because they themselves are socially isolated (Howard, Kumar, and Thornicroft,
2001; Semple, Strathdee, Zians, and Patterson, 2011).
The larger the parent’s own friendship network, the more likely parents are to put their children
in contact with potential playmates and friendship networks (Homel, Burns, and Goodnow, 1987;
Oliveri and Reiss, 1987). Strong social networks impact parents’ motivational beliefs, including those
about involvement in institutions and activities that can strengthen the quality of their children’s
social connections (Curry, Jean-Marie, and Adams, 2016). Involvement in a number of different types
of community groups and organizations can assist parents in forming and maintaining the kind of
supportive networks that facilitate good social connections for their children. These include mem-
bership in a religious community (Assari, 2013). Based on their review, Horwath and Lees (2010)
contended that constant involvement in a church can improve a parent’s overall capacity to act in
behalf of their children and motivates the parent to involve the child in the religious community as
well. It is not unusual for churches to have special classes for young children where they engage other
children of their age and get to know other adults from the church. Although religious affiliation
has been examined more often than affiliation with other social institutions, there are organizations
492
that provide support to certain classes of families in ways that provide parents access to key social
networks and services for children. For example, there are organizations designed to support military
families (Gerwitz, Erbes, Polusny, Forgatch, and DeGarmo, 2011) and high-risk families (Trivette
and Dunst, 2014). These organizations offer connections for the parents and assist families in dealing
with stress and making connections for their children as well (Saltzman, Lester, Milburn, Woodward,
and Stein, 2016).
Access to certain community groups and agencies, including day care centers, playgroups, schools,
churches, and libraries, can help parents strengthen parental ties and help parents increase children’s
social connections and access to social supports (American Library Association, 2017; Hancock,
Cunningham, Lawrence, Zarb, and Zubrick, 2015; Juvonen, 2007; Monahan, Oesterle, and Haskins,
2010). When mothers and their children become involved in playgroups, it enhances the mother’s
sense of social support and promotes children’s social connections (Hancock et al., 2015). When
schools and agencies such as Head Start reach out to families, parents are encouraged to be involved
with teachers and school personnel, involvement that increases their children’s connection to the
school (Bohan-Baker and Little, 2002; Hancock et al., 2015; Juvonen, 2007; Monahan et al., 2010;
Trivette and Dunst, 2014). A meta-analysis of efforts made by schools to engage families revealed
positive impacts of those programs that focused on parent-child joint reading, teacher-parent part-
nership, and teacher-parent communication ( Jeynes, 2012). Such efforts increased student engage-
ment in learning and feeling of connectedness to school. However, studies show that lower class
parents are generally less able to connect with teachers in behalf of their children (Horvat et al., 2003;
Lee and Bowen, 2006).
Poverty at the household, neighborhood, and country levels make it difficult for parents to foster
or maintain good social connections (DeSilva, Huttly, Harpham, and Kenward, 2007; Runyan et al.,
1998). Low SES leads to stress and makes self-regulation and positive interactions in behalf of others
(including children) more difficult (Evans and Kim, 2013). Life in poor families can lead to inter-
parental conflict and household instability, both of which make it difficult for parents to promote
good social connections for their children (Fomby and Cherlin, 2007; Krishnakumar and Buehler,
2000; Mayberry et al., 2014). In addition, low-SES parents are likely to have employment situations
that make it more difficult to maintain good family routines and robust social connections (Sheely,
2010). Poor parents are also more likely to be rearing a child alone; and that often reduces the size of
the social network the parent and the children have to draw on (Barajas, 2011).
By contrast, high-SES parents are better able to form connections with others and to work with
others toward common purpose (Horvat et al., 2003). Social class becomes a cultural frame that
motivates parents to do many things that increase their children’s connections with individuals and
institutions that provide benefits to the children (Kraus, Piff, and Keltner, 2011). Part of the parenting
motif that comes from being middle or upper class derives from engaging in higher-order occupa-
tions. Such occupations lead to different approaches to problem solving and different values (Luster,
Rhoades, and Haas, 1989). Those values, together with work-connected relationships, motivate mid-
dle- and upper-class parents to spend time in a variety of ways that promote social connections for
their offspring.
As with most of the parenting tasks discussed, the area in which families live has a bearing on
the likelihood the parent can or will engage in actions that foster high-level social connections for
their children. Living in poor countries and poor neighborhoods decrease the odds of parents or
children will participate in organizations that offer specialized programs or enriching activities and
materials (Cueller et al., 2015; Kakwani and Silber, 2008). Chung and Steinberg (2006) demonstrated
that parents living in dangerous neighborhoods had fears about the dangers present and, therefore,
would be hesitant to engage others in the neighborhood or let children go outside to play with
other children. In general, when there is low neighborhood cohesion, parents do not trust what may
happen in the neighborhood; thus, they back off efforts to engage others (Kohen et al., 2008). In
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neighborhoods that are orderly and are high in social cohesion, parents have more opportunities to
form social ties with community members, ties that bring social capital to children (Parcel and Bixby,
2016). In such neighborhoods, one sees greater monitoring of children—people know one another,
including children (Rivara and Mueller, 1987; Runyan et al., 1998). However, it is not just the social
attributes of neighborhoods that matter as regards parental behavior that foster social connections
for children; physical attributes matter as well. In some communities, streets are easier to traverse
and more pleasant to walk down. The physical affordances in some neighborhoods (e.g., availability
of parks, limited debris, sidewalks that encourage walking) present fewer threats to children’s activi-
ties and may even motivate child play and productive behavior—they are “child friendly” (Islam,
Moore, and Cosco, 2016). In such neighborhoods, children spend more time outdoors, sometimes
in the company of parents. In such situations, both parent and child are likely to engage others and
the parent is provided opportunities to foster the child’s social connections with other children and
adults. Rural areas can present something of a contrast to more densely populated urban areas with
respect to opportunities for building social capital (Bradley, 2012). Mothers in rural areas more often
perceive themselves as having a strong social network and less often voice concerns about safety in
the immediate area. Thus, they allow or encourage outdoor activities (Salmon et al., 2013). Rural
areas typically do not present as many immediate social dangers as are often found in poor urban
neighborhoods. However, rural areas typically do not afford many high-quality settings that increase
the likelihood of engaging others (day care centers, high-quality schools, libraries, museums, facilities
for lessons; Provasnik et al., 2007; Smith, 2010; Swan, Grimes, and Owens, 2013).
Not everything parents do in behalf of their children can be neatly organized in accordance with
the seven broad parenting tasks described in this section. Moreover, a single act on the part of a par-
ent may fulfill more than a single goal for a child. That said, the vast literature on parenting indicates
that, when parents accomplish tasks in a sustained and consistent manner, it bodes well for children—
perhaps particularly when the children have biological or environmental risks that otherwise make
it difficult for them to grow and develop well (Bradley et al., 1994).
The Contexts of Parenting

Scholars from the social and biological sciences generally agree that human behavior derives from
ongoing transactions between person and environment, with each transforming the other in and
through time (Ford and Lerner, 1992; Lerner et al., 2015; Sameroff, 2010; Wachs, 2000). Individu-
als are conceived as being active and goal-oriented in responding to and shaping the affordances
present in the settings where they spend time (Lewis, 1997). Parents experience, arrange, and use
themselves and other elements within the environment for purposes of achieving different parent-
ing goals (Wapner, 1987). Sometimes their use is proactive (such as when they introduce a toy to a
child for the purpose of helping the child learn a new skill), sometimes reactive (such as when they
remove a toy from a child when the toy poses a danger to the child). For example, Holden (1985)
found that some parents attempt to reduce accidents through verbally structuring the environment
proactively to divert attention from potentially dangerous conditions and to direct children toward
activities that match their capacities. But parents are not always fully in command of situations, so
sometimes the affordances of the environment disturb efforts to act in ways that are fully consonant
with parental goals. Findings pertaining to household chaos are pertinent. In the study by Johnson
and colleagues (2008), household disorder was more influential in situations where mothers had
greater reading skills themselves; in effect, household chaos seems to disrupt the generally productive
involvement of otherwise competent mothers. This likelihood also manifests itself in the findings by
Vernon-Feagans, Garrett-Peters, Willoughby, Mills-Koonce, and the Family Life Project Key Inves-
tigators (2012) in that household chaos had a stronger impact in terms of reducing positive aspects
of parenting than increasing negative ones. From the standpoint of children’s development, parental
494
actions (or nonactions) then become part of the child’s environment, part of the larger environment
in which the child is a transactor.
In an article on complex adaptive systems, Holland (1992) argued that humans operate by no single
governing equation, or rule, that controls the entire system. He further argued that humans, as com-
plex adaptive systems, are constantly engaged in “a kaleidoscopic array of simultaneous interactions”
(p. 19) with other systems. The majority of parents appear to be adapted such that they can execute
most basic parenting functions in a manner that supports offspring well-being; and they can do so
in more or less the same way within a broad range of environmental conditions. Within these broad
ranges, parents can make small modifications in particular action sequences to suit conditions, but the
impetus to act, and the character of the action pattern directed toward a particular goal, remain largely
unaffected. However, there are some aspects of the environment that, if encountered in an extreme
range, cause marked changes in attitude and behavior. Moreover, productive pursuit of personal goals
(including goals for children) requires living in settings where daily events are predictable and at least
somewhat controllable (Ursin and Eriksen, 2010). Otherwise, parents will become overwhelmed and
resort to inconsistent and/or negative patterns of behavior (Repetti, Robles, and Reynolds, 2011).
This section of the chapter is devoted to an examination of how climate and the ambient, built, and
community environments broadly construed affect parenting. There is also a consideration of how the
materials present in the home (particularly toys and learning materials) affect parenting and how the
child affects parenting as both can affect parenting beliefs and cognitions (Bornstein, 2016).
Climate
In their review of geography and its relation to human well-being, Sachs and his colleagues (2001)
observed that the majority of poor countries are found in the tropics. They go on to state “a more
precise divide of this geographical divide can be obtained by defining tropical regions by climate
rather than by latitude” (p. 72). Although the effect of climate on parenting practices has been inves-
tigated to only a limited degree, climatic conditions place broad constraints on human behavior.
For example, when precipitation and temperature fall outside moderate ranges, conflict goes up
(Hsiang, Burke, and Miguel, 2013). Such would not generally bode well for most of the parenting
tasks described above, albeit cultural practices can sometimes offset general human propensities for
behavior. Seasonal affective disorder is more common in latitudes outside the tropics (Mersch, Mid-
dendorp, Bouhuys, Beersma, and van den Hoofdakker, 1999). Seasonal affective disorder can affect
both parents and children, leading to sadness and lethargy.
Whiting (1981) demonstrated that climate, apart from cultural history, constrains infant care prac-
tices in Europe, Africa, and Asia. The temperature of the coldest month of the year was the most
important factor in determining whether infants were carried in cradles versus a sling or shawl.
These carrying practices also affected mother-infant contact and parenting behaviors. For example,
Tronick, Thomas, and Daltabuit (1994) observed that the practice of tightly swaddling infants in
high-altitude cold climates, while acting to maintain body warmth and to conserve calories, also
resulted in restricting the infant’s motor activity and the mother’s provision of stimulation. Not
surprisingly, there is also evidence for seasonal variations in the onset of motor milestones in those
regions where there are distinct seasonal changes in temperature. Hayashi (1990) found a consistent
periodic relation between mean monthly temperature in Osaka and the onset of rolling, creeping,
grasping a foot, and grasping a toy. Relatedly, Benson (1993) found that, if the window of locomotor
onset occurs during winter, experiential factors associated with the winter may limit opportunities
for mobility, including heavier clothing that restricts movement, fewer daylight hours for outdoor
play, and restriction to a smaller play area inside the house.
As children get older, they are often allowed greater autonomy in keeping with their increased
competence. Accordingly, out-of-home activities increase. Climatic conditions directly affect the
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amount and the type of parental monitoring and involvement with children (seasonal variations,
mean annual rainfall, mean temperature). Climate also affects parental willingness to allow outdoor
play (Brockman, Jago, and Fox, 2011). Parents’ employment can also be affected by seasonal and
climatic conditions. Crouter and McHale (1993) found that the quality and types of parent-child
involvement, parental monitoring, and children’s involvement in activities all reflected seasonal vari-
ations in parental employment patterns.
The Ambient Environment

Worries about obesity have increased interest in how ambient surrounding conditions affect human
behavior, particularly physical activity. Ambient conditions can affect parent motivational states for
better and for worse (Evans, 2006). Importantly, they can affect a parent’s sense of personal control in
situations; and external conditions that put pressure on parents can lead to more controlling behavior
(Grolnick, 2003). As a consequence, ambient environmental conditions could affect parental efforts
pertaining to most of the parenting tasks discussed earlier in this chapter. Some ambient stressors
(like noise from traffic or nearby airports) tend to be chronic and nearly intractable. Living in high-
density urban spaces for extended periods of one’s life, as is the case for a high percentage of the
world population at present, can affect neural social processing in humans (Lederbogen et al., 2011).
Living in deteriorating, dangerous urban neighborhoods has been linked to fear of crime on the
part of adults (Taylor and Brower, 1987). This fear leads to restricted outdoor activity, avoidance of
certain areas, and more intense efforts to control children’s behavior (Gates and Rohe, 1987; Henson
and Reyns, 2015; Kail and Kleinman, 1985). Long-term residence in crowded urban areas can lead
to mood disorders, anxiety, and poor mental health. Such conditions may induce parents to provide a
kind of “stimulus shelter” for themselves and their children. However, if parental efforts to cope with
these conditions fail, the parent may develop more pervasive feelings of helplessness and a generalized
tendency to give up in the face of obstacles (Cohen, Evans, Stokols, and Krantz, 1986). Unfortunately,
not being able to escape crowded conditions can also lead to child maltreatment (Garbarino and
Kostelny, 1992). Living in crowded, deteriorating conditions can have negative consequences for
children’s behavior as well; and, if it does, it can result in a negative cascade of caregiving on the part
of the parent. In effect, living in crowded conditions can lead to less and less cooperation between
parent and child and more and more negativity (Brock and Kochanska, 2016; Dix, 1991).
People have a preference for environments that contain natural features, such as lakes and oceans
(Hartmann and Apaolaza, 2010; Ogunsteitan, 2005; White et al., 2010). Studies of place attachment
and neighborhood satisfaction appear to corroborate the idea that people feel a stronger and more
positive connection to their surrounding community when it affords them encounters with veg-
etation and other natural elements, including opportunities for gardening (Comstock et al., 2010;
Hur, Nasar, and Chun, 2010; Scannel and Gifford, 2010), a major contributor to the “liveability” of
a geographic area (Badland et al., 2014; Kaplan, 1983). As a case in point, Kaplan (2001) found that
being able to see trees from one’s home, whether in the suburbs or inner city, was associated with
a heightened sense of ease and relaxation. Such findings are consonant with the idea that aesthetic
conditions create a perception of how supportive an environment is. In a study of families from 64
urban public housing units, Taylor, Wiley, Kuo, and Sullivan (1998) observed more play in high
vegetation spaces, more children engaged in creative play when in high vegetation spaces, and more
children with access to adults when in high vegetation spaces.
Unfortunately, much remains unknown about the impact of ambient surrounding conditions on
parenting practices. Part of the problem derives from a poorly specified framework regarding how
the affordances of settings influence parental behavior (Stokols, 1995). Notions about area liveabil-
ity would seem somewhat helpful in this regard, as those notions incorporate perceptions that lead
to both positive and negative motivational states (Badland et al., 2014). On the negative side, the
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cognitive activation theory of stress stipulates the people make evaluations pertaining to the control-
lability of a situation and the potential consequences of their own efforts to exercise control (Ursin
and Eriksen, 2010). A positive expectancy (“coping”) arises when a person concludes that he or she
can handle the situation with positive results. By contrast, when an individual concludes that most or
all responses lead to a negative outcome, the acquired expectancy is “hopelessness”. A neighborhood
or nearby geographic area that is dilapidated or poses threats to safety, may move parents toward a
sense of hopelessness. To some extent an area which simply has few facilities for recreation and fam-
ily support can be perceived so as well. By contrast, ambient conditions have the potential to impact
a parent’s sense of subjective well-being and sense of purpose as well (Lamis, Wilson, Trantino,
Lansford, and Kaslow, 2014). A sense of well-being enables one to adapt to daily challenges and take
advantage of existing opportunities by appraising them somewhat more positively and objectively
and taking time to plan one’s actions (Deiner, Suh, Lucas, and Smith, 1999). Areas where there are
social and physical amenities that improve chances for employment, education, and health care, and
places that afford opportunities for recreational pursuits, likely enhance parents’ sense of well-being
and motivate them to take productive actions in behalf of their children. However, at present there is
little direct research on how ambient conditions impact parents’ subjective well-being or the adapta-
tions parents make as a consequence.
For the better part of a half century, environmental psychologists have promulgated the idea that
the amenities or affordances of one’s place of residence have an impact on how one feels about the
place and one’s dispositions regarding activities and other people in the place (what is called “place
attachment”). It has never been easy to define or measure exactly what place attachment is, but there
is some evidence in support of the idea that the affordances present in a setting or place influences
how much one identifies with the place and how much one feels bonded to the place (Lewicka,
2011). There is also some evidence that one’s attachment to a place connects to how one feels, thinks,
and behaves while in the place. The happier and more comfortable one feels in a place, the more one
is inclined to act in productive ways with others who share that place (Scannel and Gifford, 2010).
Perhaps unfortunately, there is very little research on how place attachment affects parental cogni-
tions or behavior; thus, one can only speculate at present on what that means for parents and children.
The Built Environment

The components of physical settings tend to facilitate certain activities while obstructing others
(Weinstein and David, 1987). What parents tend to do with their children (from play arrangements
to sleeping arrangements to the use of control techniques) is partly determined by the physical fea-
tures of their residence (i.e., size, number of rooms, proximity of certain spaces to others, acoustical
properties, amenities). For example, in Kenya, among the Embu community, children spend very
little time inside their homes, which are situated such that they open onto communal areas. When
children become mobile, they join other children from the community outside to play or to per-
form chores. As a result, children do not tend to be in close proximity to their parents; thus, they
are not very likely to engage in verbal exchanges with adults. By contrast, Egyptian homes tend to
be quite small; and they are not arranged so that they open onto communal areas. Toddlers spend
most of their time within the residence or immediately outside it. Because of their close proximity
to adult family members, Egyptian toddlers tend to have their verbalizations responded to with high
frequency (Wachs et al., 1992).
When a residence or nearby areas cease to accommodate family needs, parents often try to correct
the problem: by moving or making adjustments to the property. Otherwise, they change their own
behavior so that it fits with the conditions present. Adjustments of many types are made in family
residences to accommodate children’s needs or interests (e.g., buying new furniture, “baby proof-
ing” the house, changing the decorations in a child’s room to satisfy the child’s emerging interests,
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establishing a place for a child to study, reworking the outside play area and installing more advanced
equipment, moving a child to a separate room as the child gets more independent, installing door
and cabinet locks to keep a child safe). When homes are crowded parents are more apt to send their
children out to play (Gove, Hughes, and Galle, 1979). In Japan, because most families live in close
quarters, there is typically very little space contiguous to the house for children to engage in outdoor
play. Thus, parents typically have to arrange for children (or permit, in the case of older children) to
go to parks, playgrounds, and game halls for recreation (Weinstein and David, 1987). As regards mak-
ing changes to the nearby environment, parents sometimes engage homeowner’s associations, neigh-
borhood organizations, or governmental bodies to make adjustments in nearby areas, sometimes in
the company of their children. Such actions are more likely when there are active organized groups
in the community (Center for the Study of Social Policy, 2011). That said, most of what is known
about the impact of the accouterments of physical settings on making such changes is known by
anecdote and not as a consequence of household surveys or more focused research.
The Community Environment

Neighborhoods and communities vary considerably with respect to the number of places and
organizations that are available for recreation and learning. A major avenue of potential stimulation
for children is the library. Americans have a favorable view of libraries, but actual use has declined
(Meyer, 2016). The decline in utilization of libraries and museums is associated with a decline in
expenditures by state and local governments on such facilities (Institute of Museum and Library
Services, 2016). The National Association of State Park Directors (2015) estimated that 739 mil-
lion people visited a state park or recreational area in 2014. The National Park Service (2016)
recorded over 307 million visits to national parks in 2015 as well. In effect, relatively little is known
about parents’ actual use of community cultural and recreational services such as libraries, museums,
playgrounds, youth clubs, and parks. Likely, the motivation to use them involves a diversity of con-
siderations (e.g., other family obligations, family composition, income, and ease of access). Culture
and parental work status also play roles (Hutchinson, 1987, 1988; Outdoor Foundation, 2013). As
an example, Asian Americans tend to use certain recreational facilities (e.g., ball parks) less often
than other ethnic groups even when such facilities are available, partly owing to a belief that there
are more long-term opportunities in other domains of life, such as formal education (Bradley et al.,
2000; Leong, Chao, and Hardin, 2000).
Changing technologies, changing patterns of employment, changes in residential location, changes
in family configuration, and governmental efforts to increase leisure and cultural opportunities have
led to major changes in the ways family members spend time that is not committed to work, to
routine family maintenance functions, or to education (Veal, 1987). According to the 2015 American
Time Use Survey, people over the age of 15 spend almost 5 hours each day in various leisure pursuits
and sports activities (based on all seven days of the week). Married women with children under the
age of 6 only spend about three hours in leisure and sports, however, The time use survey provides
breakdowns for both mothers and fathers who have various levels of employment, breakdowns into
major categories of leisure activities, travel, and other activities (e.g., shopping) that occur outside
the home. Although these figures provide a glimpse of how parents spend their time, some of which
might include involvement with children, there are no actual details on how often children are
involved or the nature of their involvement.
Some sense of what parents do with respect to children’s activities comes from data on how much
children are involved in certain types of activities. There is information on children’s participation
in sports, for example (The Aspen Institute, 2016). The National Household Education Survey also
offers information on children’s involvement in various forms of arts and crafts activities, board
games and information on how often they go to places like libraries art galleries, museums, zoos, and
498
church (U.S. Department of Education, 2016). The NHES also includes information on how often
a 3- to 5-year-old child was read to, was taught basic concepts, or told a story by a family member
in the past week. Having access to extracurricular programs at local schools seems to motivate chil-
dren’s involvement in arts, drama, music, and dance. The Physical Activity Council (2016) reported
that most school-age children are physically active and that almost half of parents whose children
participate in sports paid an extra fee for them to do so. According to a report by the Pew Research
Center (2016), more than 70% of school-age children are involved in some type of sport or athletic
activity, more than 60% in a religious activity, and more than 50% in some type of music, art, or dance
lesson. Such activities not only require parental permission, but often entail costs and parental time
(e.g., going to performances).
Toys and Materials

Another factor that helps determine what parents do as caregivers is their access to objects and
materials that can serve as resources for children’s recreation and development. The U.S. Bureau of
Labor Statistics (2016) conducts a consumer expenditure survey each year. Respondents are asked
to estimate expenditures for many items and activities, including expenditures on foods, education,
various types of equipment, toys, entertainment, and other items that may reflect parental decisions
linked to having a child. There are breakdowns in the reports indicating considerable difference
in the allocation of expenditures in these categories by family composition. As examples, married
couples with children spend a greater share of their income on housing, children’s clothing, toys and
playground equipment, and education. A Gallup Poll done in 2013 found that U.S. Americans who
have at least one child spend an average of $29 a day more than those without children. Moreover,
the differential goes up as the number of children increases (Wilkie, 2014). There remains limited
research on how access to particular types of toys and materials influences parental interactions with
children, but the existing research indicates that both the type and amount of materials available in
the home help frame those interactions (Bradley, 1985).
As stated earlier, the deep infusion of various types of media and communication devices have
altered much about family life (O’Keefe, Clarke-Pearson, and Council on Communications and
Media, 2011; Villegas, 2013). It is influencing how parents structure children’s time and they ways
they monitor children’s behavior (Nikken and Schols, 2015; Pew Research Center, 2016). Impor-
tantly, there is evidence that when children are engaged with media, it decreases the quality of
parent-child interactions (Kirkorian, Pempek, Murphy, Schmidt, and Anderson, 2009). Although
devices, such as cell phones, have the potential to increase connectivity between parents and children,
there is increasing evidence the having access to smartphones and other devices that allow access to
the internet interferes with ongoing parent-child interactions (Pew Research Center, 2012).
Child Effects
According to the Belsky and Jaffee (2006) model of parenting, a major social influence on parental
behavior is the characteristics and behavior of a given child. Family decisions regarding where to
go on vacation, whom to visit, and what to buy are influenced by the characteristics of children
within the family (Crompton, 1981; Howard and Madrigal, 1990; McNeal, 1992). Families with
children are more likely to engage in outdoor recreation, for example (Outdoor Foundation, 2013).
Corrigall and Schellenberg (2015) found that parents were more likely to enroll children in music
lessons and continue those lessons for a longer time when the child was high in agreeableness and
showed and openness to experience. In support of the idea of child effects is a history of research
on consumer spending that shows child influences on parental purchases (Hall, Shaw, Johnson, and
Oppenheim, 1995; Martensen and Gronholdt, 2008). According to a YouGov Omnibus Parents
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Survey, children are often successful in persuading parents to purchase certain types of toys, clothes,
and equipment; and they are also persuasive in determining the places families eat and where they go
for entertainment (Harmeling and Gammon, 2015). As children get older, they are more inclined to
solicit information from peers and media. As the field of marketing has come to better understand
children’s influences on parental decisions, and how children get information about products and
services, those in marketing have done more to target ads to key subgroups of children (Martensen
and Gronholdt, 2008). Details about exactly what parents do vis-à-vis their children during particular
activities or at particular venues remain fuzzy, but there are clear indications that parents adjust their
own time use, pursuits, and purchases in response to their children (Harmeling and Gammon, 2015;
Horna, 1989; Howard and Madrigal, 1990).
Just as a child’s characteristics and behavior impact what parents do, so too does a parent’s own
history of being parented (Belsky and Jaffee, 2006). Wells and Lekies (2006) found this to be the case
with respect to children’s spending time out-of-doors. If a parent had a history of spending time in
nature as a child, it was more likely that the parent would spend time with his/her own child doing
things in natural surroundings.
Complexities and Limitations in Studying

Environment-Parenting Relations
Human beings are complex creatures, and most of us live in complex, dynamic environments (Born-
stein, 2016; Lerner et al., 2015). Accordingly, it is not easy to pin point how a particular experience
or aspect of the environment affects a particular type of parental behavior (a parenting task to use
the nomenclature above). To begin with, parents do not experience their environments as isolated
elements. Any one event or circumstance is experienced in the context of other objects, situations,
and happenings both in time and across time (Bronfenbrenner and Morris, 2006). The “effect” of one
element on parent or child behavior is likely to be conditioned by the presence of the other elements
in the contemporaneous surround; that is, a parent’s take on what an element means is simply one
element of a much larger composite—it is a constructed meaning. Unfortunately, very few studies
of parental behavior have simultaneously considered multiple aspects of the environment, much
less how one aspect of the environment might moderate the impact of others. As a simple example,
having lots of toys in a room would generally increase the likelihood a parent would provide useful
stimulation to a child and perhaps structure the child’s experiences in such a manner that the child
would enjoy and learn more. But, if one adds an element of outside noise, a text on a smartphone,
an older rambunctious sibling, and work stress to the mix, the odds of taking such actions might be
reduced or their exact nature altered. Taking a slightly more positive view, having a second, interested
parent might increase the likelihood a parent would provide nicely structured, joyous stimulation.
An important corollary to the idea that humans do not react to experiences as isolated bits is the
idea that the co-occurrence of particular environmental elements tends not to be random, either in
or through time. Some sets of co-occurrences reflect more general ecological circumstances, such as
poverty (high crime, more single-parent families, more household crowding, greater likelihood of
exposure to environmental toxins) or area of residence (isolation from neighbors, lack of access to
culturally enriching events and facilities, more frequent encounters with “nature”). Likewise, some
reflect the ongoing interplay of skill sets and personality dispositions (intelligence and positive moti-
vations versus mental illness and addiction problems). To add one more dimension to the problem of
co-occurring elements, there is also person-environment covariation; that is, certain environmental
conditions tend to increase the likelihood of certain personal characteristics and vice versa (Wachs,
2000). Without sufficient inclusion of variables that represent these connected elements, findings can
be misleading, sometimes spurious. Unfortunately, most research on factors that contribute to the
likelihood parents will engage in particular behaviors has not examined environmental elements that
500
often co-occur as part of a compound (e.g., the factors that contribute to low levels of employment
also increase the likelihood of residential and marital instability and household crowding).
A third factor that makes it difficult to ascertain how environmental circumstances affect parent-
ing behavior is that theories of human behavior do not always provide adequate guidance as to the
nature of the presumed “effect”. In some cases, there can be multiple paths to the same outcome. In
some cases, the effects can be indirect rather than direct. In some cases, the effects only emerge after
a long accumulation of exposure. And, in some cases, the relation may be curvilinear rather than
linear. Extreme events or conditions (trauma, catastrophe) are more likely to produce curvilinear
relations (Little, 1999). One of the principles of dynamic systems theory is that minor adjustments
in one component of a system can result in major shifts in a second (Prigogine and Stengers, 1984).
For example, a single incident of harassment at a playground or ball field may induce a formerly
active mother to quit involving her children in organized sports activities altogether. In real life
there are both thresholds above which more of some condition (presumed cause) does not produce
a concomitant degree of additional “effect” (Bornstein and Manian, 2013) and points beyond which
matters start to quickly cascade to a lower level (Bornstein, Hahn, and Haynes, 2010).
Finally, and perhaps most fundamentally, it can be difficult to accurately estimate environmental
effects because human beings are conscious actors in their own lives, and they do not all react in the
same way to identical environmental conditions (Ellis, Boyce, Belsky, Bakermans-Kranenburg, and
van IJzendoorn, 2011; Lewis, 1997). As Wallenius (1999, p. 132) has argued, “No goals or environ-
mental settings are isolated units in a person’s life. The psychological significance of situations and
settings depends on their relationships to the individual’s overall life situation and plans”. Parenting
goals are enormously diverse and so are relationships between family members. Therefore, what
is of minor interest to one parent may be quite consequential to another. Within the confines of
intimate family relationships, a particular event can lead to a spiraling escalation of behavior of the
kind described by dynamic systems theorists as turbulence (Prigogine and Stengers, 1984). Some-
times small events induce rapid, unpredictable changes in behavior. Turbulence in family systems is
probably rather common, particularly at times when highly significant goals and issues emerge in
interactions between parents and children (e.g., when a child has not spent enough time preparing
for a test, when an adolescent comes home later than expected from a date or activity with peers,
when a child talks back to a parent or refuses to comply with a parental directive, when a child reveals
she has been victimized or reveals suicidal thoughts, when the parent catches a child playing with a
loaded gun or using drugs, when a 15-year-old daughter says she is pregnant by her boyfriend). Such
moments are fraught with significance and can radically alter parental thinking and behavior; but
these are moments about which there is little authoritative research.
Conclusions
Both parents and children face an extraordinary array of circumstances during the march through
life. For the vast majority of these circumstances, research is silent or speaks with limited authority
and precision. Moreover, the literature is replete with disagreements between parents and children
on exactly what parents do as regards the tasks of parenting discussed. For a few dire circumstances
(war, trauma, deep poverty, divorce, family conflict, parental death), there is reasonable coverage of
how parents tend act to accomplish some of the tasks of parenting; but even for those circumstances
critical details are missing (Davies and Cummings, 1998; Garbarino, 1999; Lieberman, Chu, Horn,
and Harris, 2011; Shaw, 2000)—too much is lacking to strongly inform educational or intervention
efforts. Unfortunately, when circumstances put the parents themselves under stress and erode their
own support, there is a tendency for the parent to become disorganized and to withdraw from the
child and become less aware of what the child is actually experiencing. If so, it can make matters even
more difficult for those who wish to intervene in productive ways (Banyard, Rozelle, and Englund,
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2001; Cerel, Fristad, Weller, and Weller, 2000; Coates and Gaensbauer, 2009; Knott and Fabre, 2014;
Saler and Skolnick, 1992). These limitations in the research canon acknowledged, the broader litera-
ture on what children need in times of crises provides at least some direction on the kinds of research
required to determine what social and physical aspects of the environment might influence parental
efforts to engage in key parenting tasks that might assist child well-being (e.g., socioemotional sup-
port, stimulation, structure, surveillance, social connections). Some of those conditions might be
more amenable to change than others, and, thus, might be more useful targets for the next generation
of research on how to assist parents in building useful skills and supports. Toward this end, ideas from
broaden and build theory and selective investment theory (Cohn and Frederickson, 2006) and the
technology of skills formation (Cuhna et al., 2010) seem potentially applicable.
Even though many of the details connected to the complex interplay between parents and their
environmental circumstances remain to be determined, it seems accurate to state that the objects and
the materials available to parents in their place of residence, the conditions present in the area sur-
rounding the place of residence, and other facilities in the nearby community play an integral role
in how parents parent. One could make such a statement with more confidence if there was con-
sensus on just how the environment should be conceptualized (Garling, 1998). But that, like many
of the other issues discussed, is an evolving matter. The research needs to move in directions that
make clearer how parents can adapt to whatever constraints their environments impose and can take
advantage of the opportunities afforded so that parents can more productively engage in the critical
tasks of parenting. In pursuing the agenda of more fully explicating relations between the parenting
environment and parenting, it is critical that scholars bear in mind that parent-child relationships are
themselves an ever evolving dialectic, one that begins before a child is born and continues after the
parent’s death (Kuczynski, Pitman, and Mitchell, 2009). The notion of homophily in social networks
is also worth bearing in mind. Parents are likely to connect to people and entities that are in some
ways similar to themselves; and, therefore, their actions as parents will in various ways reflect these
“birds of a feather” connections (McPherson, Smith-Lovin, and Cook, 2001).
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518
INDEX
Note: Page numbers for figures are in italics, and page numbers for tables are in bold.
1000 Genomes Project 128 adrenocortical hormone 41

adrenocortical synchrony 233 – 235
abandonment 21, 289 adrenocorticotropic hormone (ACTH) 41, 180 – 181,
abortion 303 196, 230
abuse: depression and 237; environment and 479, 487; Adverse Childhood Experiences (ACE) study 197,
evolutionary perspectives and 4, 20 – 22; modern 359 – 360
history and 333 – 334, 340; primate 85, 109 advice literature 334 – 335
acceleration 13 Aeschylus 288; The Libation Bearers 303, 306
accidents 327, 480, 490, 494 Aetius 316
acculturation 378, 451, 464, 487 affect 44 – 45, 49, 273, 457 – 458
accuracy vs. precision 356 – 357 affective prediction hypothesis 265
ACE (Adverse Childhood Experiences) study 197, affiliative hormones 222, 223, 236 – 238
359 – 360 Afonso,V. 50
Ache culture 480, 489 Africa 291, 324 – 326; culture in 448, 452;
Ackerman, B. 485 environment and 495; evolutionary adaptedness
ACTH (adrenocorticotropic hormone) 41, 180 – 181, and 9; grandmother hypothesis and 17;
196, 230 individualism in 342; labor in 328; modern history
Adams, E. 482 and 337 – 338
adaptation 6, 9 – 10, 177, 196 – 199, 495 African American families: childcare and 328;
Add Health (Longitudinal Study of Adolescent culture and 456 – 464; environment and 479,
Health) 382 – 384 486 – 487; neighborhoods and 376 – 378, 382 – 390;
ADHD (attention deficit hyperactivity disorder) 139, parental educational attainment and 406 – 407;
152, 183, 187 – 189, 344 socioeconomic status (SES) and 422, 426 – 428
adiponectin 190 aggression: in adoptees 127; culture and 455 – 456,
adolescence: developmental studies of 145 – 152; in 463; evolution and 20; hormones and 228, 232;
nonhuman primates 79 – 80, 95 – 96, 100, 105 – 107; hostile parenting and 140, 143 – 144; maternal
parental negativity and 139; in Sparta 309; 52 – 53, 99, 104; neighborhoods and 379, 384;
temperament in 127 – 128; as Western idea 321, in nonhuman primates 95, 97 – 98, 108; social
330 cohesion and 11 – 12; socioeconomic status (SES)
Adolescent/Adult Family Development Project 149 and 422
adoption: ancient history and 290, 297; hormones Agora 306
and 221; infant cues and 270; studies 125 – 133, Agrati, D. 56
131, 136 – 144, 152, 365 agriculture 9, 452, 465, 478
adrenal gland 36, 180 AHN (anterior hypothalamic nucleus) 48 – 49
adrenal steroids 41 Ainsworth, M. 19
519
Index
Akhenaten 292 arousal 262, 487

Akkad 291, 295 – 296 ASD (autism spectrum disorders) 233 – 234, 237 – 238
Alenina, N. 52 Asia 291; 19th century in 336; culture in 448 – 452,
alert states 83 457 – 458, 461 – 465; environment and 483, 495;
alleles 57, 134, 140 – 142, 153, 225 illegitimacy in 338; individualism in 341 – 342
alloparents 17, 79, 98 Asian American families 457, 460 – 461, 498
alpha males 100 assisted reproductive technologies (ART) 194 – 195
ALSPAC (Avon Longitudinal Study of Parents and Assyria 291, 295 – 296
Children) 135 – 138, 183, 190, 351, 357 – 358 Ataca, B. 451, 465
altriciality 13 Ateles 104
amargi 295 Athens 288 – 290, 300 – 309, 317
ambient environment 496 – 497 attachment 6, 251 – 252; bonding and 191 – 192;
American Academy of Pediatrics 479 Bowlby and 254; evolutionary perspectives on
American Time Use Survey 426 – 427, 498 17 – 23; hormones and 220 – 221, 225 – 226; infant
amniotic fluid 38 – 40 crying and 262; influences on 136; in nonhuman
amphidromia 305 primates 104, 107 – 108; in rats and mice 39;
amplification model 383 – 388, 386 secure 135
amygdala 44 – 49, 52 – 53, 224, 227, 235, 271 – 272 attachment Q-Sort 136
anaptic animals 47 attention deficit hyperactivity disorder (ADHD) 139,
Anatolia 291 152, 183, 187 – 189, 344
ancient Egypt 288 – 297, 300, 316 – 317 attitudes 425 – 426, 450, 453 – 456
ancient Greece 288 – 292, 300 – 310, 317 Atzaba-Poria, N. 464
ancient history 287 – 319 Au, E. 404
ancient Israel 288, 291, 296 – 300, 317 auditory cortex 40, 49
ancient Mesopotamia 288 – 292, 295 – 296, 300, 317 auditory cues 40, 253, 267
Andean society 478 auditory cuteness 262 – 263
Anderson, K. 22 Augustine 313, 314
androcentrism 290 – 291, 296 aunts 16, 132
andron 301 Australia 22, 191, 333, 336, 455
anemia 184 – 185 Australian Twin Registry 148
anhedonia 51 Australopithecus afarensis 12
Annales school 321 Australopithecus genus 9
anogenital regions 33, 38 – 40, 189 authoritarianism 151, 461
anorexia nervosa 323 authoritative parenting 426, 461
anosmia 38, 47 autism spectrum disorders (ASD) 233 – 234, 237 – 238
antagonistic neural systems 46 – 49 autistic behaviors 189
antenatal care 192 autoradiographic method 48
anterior hypothalamic nucleus (AHN) 48 – 49 avoidance 109
anthropology 291, 448 – 449 Avon Longitudinal Study of Parents and Children
anthropometry 190 (ALSPAC) 135 – 138, 183, 190, 351, 357 – 358
antiestrogens 47 AVP (vasopressin) 33 – 36, 46, 220 – 224, 227 – 228
antipsychotic drugs 52 – 53 avuncular relationship 132
antisocial behavior 139 – 141
antisocial personality disorder 127 baboons 17, 97 – 100
anxiety 34 – 36, 44, 88, 129, 141, 182 – 183, 343 – 344 Babylonia 291, 295 – 297
apes 9, 80, 82 – 85, 94 – 96 baby-schema 18, 253, 257, 266
Aphorisms (Hippocrates) 308 baby talk 306
Apollo 288 Bachofen, J.: Myth, Religion, and Mother Right 290
apprenticeships 90, 301 Bahrain 484
Aque, C. 460, 463, 466 Barbary macaques 97
Arab societies 454, 458 Bard, K. 80, 92, 108
Archaic Homo sapiens 9 Baring, A. 290 – 291
Ardipithecus genus 9 Barker, D. 179, 185
Ariès, F.: Centuries of Childhood 289, 322 – 323 barrenness 295 – 297
aristocrats 292, 296, 300 – 306, 316 Barros, A. 351
aristoi 300 Bates, J. 486
Aristophanes: The Clouds 306 – 307 Bauer, J. 39
Aristotle 302 – 305, 304, 308 – 310, 313; Politics 304 Baumrind, D. 426, 461
520
Index
Beach, F. 37, 46 Bogin, B. 14

Beam, M. 455 – 456 Bohlin, G. 183
Beaver, K. 151 Bollen, K. 401
bed nucleus of stria terminalis (BNST) 48, 52 Bolten, M. 236
Bedouin Arab culture 454 bonding: pair 11, 85, 100, 226 – 228; prenatal
behavior: cues 18, 267 – 269; culture and 450, 191 – 192, 225; selective 39
456 – 461; evolutionary perspectives and 5 – 6; home bonnet macaques 99
environment and 405 – 407, 487; neighborhoods bonobos 9 – 11, 82, 85
and 377 – 381; in nonhuman mammals 30 – 77; in books 350 – 351, 429, 434, 481
nonhuman primates 78 – 122; parenting negativity boredom 332
and 138 – 140, 148; stress and 109, 182 – 184 Bornstein, M. 449 – 450, 460, 476, 482
behavioral biology 170 Bosch, H. 289
behavioral genetics 123 – 165, 124 Bosch, O. 36
Behavior Genetics (Fuller and Thompson) 124 – 125 Bosnia 350
Behavior Genetics journal 135 Bossou 82
Beijing 483 Bowlby, J. 6, 17 – 19, 254
Belarus 351 Boyle, T.: Tortilla Flats 475
beliefs 404 – 405, 425 – 426, 430, 450, 479 boys 14, 140, 183 – 184, 189, 382 – 386, 389
Belsky, J. 19, 151, 476 – 477, 487, 491, 499 Bozoky, I. 482
Benediktsson, R 181 BPA (Bisphenol A) 187 – 188
Benner, A. 408 Bradley, R. 485 – 487
Benson, J. 495 brain development 182
benzodiazepines 36 brains: neurobiology and 252 – 258, 255, 260 – 261,
benzyl butyl phthalate (BzBP) 188 268 – 271, 274; in nonhuman primates 107 – 108;
Bergeman, C. 126 size and structure of 10 – 15, 270, 423; social
Berkson’s bias 362 neuroendocrinology and 235 – 236
Bes 293 brainstem 264 – 265, 268
beta-hydroxysteroid dehydrogenase type 2 168, brain wave patterns 83
180 – 182, 185 – 186 Brazil 351, 454, 475, 490
Beveridge, M. 182 breastfeeding 39, 237, 294, 305, 309, 479
biallelic expression 57 Brett, Z. 56
bias 356 – 366, 366 Bridges, R. 39, 42, 107
Bible 296 – 300 Britain 322, 325, 330, 337, 350, 380 – 381
bioassays 17, 221 British Scientific Advisory Committee of the Royal
biobehavioral synchrony model 222, 232 – 235 College of Obstetricians and Gynecologists 195
bioecological interaction model 126 broaden and build theory 502
biological mechanisms 173 – 184 Bronfenbrenner, U. 374, 403, 426, 452
biological stressors 178 – 180, 195 – 196 Bronstein, P. 462
biomonitoring studies 187 – 189 Brooks-Gunn, J. 377, 402, 488
bipedality 9, 13 Brouette-Lahlou, I. 40
birds 18, 228 Brown, L. 194
birth 41, 293, 302 brown lemurs 105
Birth, Death, and Motherhood in Classical Greece Bruschi, C. 459
(Demand) 304 – 305 built environment 497 – 498
birth control 303, 320, 325 – 326 Bullock, L. 488
birth rates 321, 324 – 327, 335 burial 302, 341
birth weight and length 21, 170, 175 – 187, 194, bushbabies 107
197 – 198 Byzantine medicine 316
Bisphenol A (BPA) 187 – 188 BzBP (benzyl butyl phthalate) 188
Black, M. 486
Blair, M. 460 cadmium 187
Blair, S. 460 Cadoret, R. 126 – 127
blue monkeys 96 – 98, 98 Cain, D. 486
Bluestein, D. 463 Caldwell, B. 485, 487
BNST (bed nucleus of stria terminalis) 48, 52 California mice 229
Bocanegra, J. 460 calling 96, 101
body mass index (BMI) 187, 190 Callithrix jacchus 103
Boesch, C. 90 callous-unemotional behaviors 138, 141
521
Index
Cambodia 337 – 338 Child Development Project 151

Cameron, A. 180 child effects 434 – 435, 499 – 500
Cameroon 458 – 460, 465, 484 child homicide 22
Canada 17, 336, 380, 451, 454 – 455, 458, 462 – 463, 484 childhood: brain size and extended 15; early
canalization 169 – 170 135 – 142; history of 288 – 290; middle 142 – 145,
cancer 194 154; psychosocial stress in 197 – 198; social
candidate gene-by-environment interaction (cGxE) construction of 303 – 308, 311 – 313
128, 134, 141 – 142 childhood mortality rates 16, 297, 302
candidate genes 124, 134 – 137, 140 – 142 childlessness 292, 302, 345 – 346
cannibalism 37 – 38 child outcomes: culture and 450; evolutionary
CAP (Colorado Adoption Project) 125, 142 – 144 ecology and 166; influence of parental brain
capuchin monkeys 100 – 105, 102 and hormones on 235 – 236; neighborhoods
CAR (cortisol awakening response) 230 and 372 – 375, 379 – 391, 386; parent education
caregiving 78, 87, 254, 270, 304, 314, 426 – 427 attainment and 400 – 404, 408 – 410; socioeconomic
Carlson,V. 460 status (SES) and 421
carrying 87, 94, 101 – 106 childrearing: advice literature on 327; ancient history
Casciano, R. 377 and 288 – 290, 293 – 295, 299 – 302, 309 – 313;
Cashford, J. 290 – 291 beliefs about 404 – 405; culture and 448 – 450, 462;
catarrhines 81 evolutionary perspectives and 3 – 29; parenting
catechol-O-methyltransferase (COMT) 136 model and 22 – 23; permissive and restrictive
Cato the Elder 312 98 – 99, 383; socioeconomic status (SES) in
caudate activation 235 – 236 425 – 426
causality 351 – 366, 431 – 432 children: evolutionary perspectives and 5 – 6;
CD38 225 – 226 extended immaturity and dependency of 3 – 5,
CeA (central amygdala) 53 330; historical narrative and 287; hormonal
Ceballo, R. 486 synchrony and 232 – 235; in hunter-gatherer
cebus 101 societies 15; neighborhoods and 379 – 381; parent
Cebus apella 104 educational attainment and 408 – 413; parenting
Cebus capucinus 102 and characteristics of 137 – 140, 143 – 144, 499 – 500;
Celsus: Historia Naturalis 315 – 316 social construction of 303 – 308, 311 – 313; as
Cen, G. 462 workers 325, 329 – 330
Central America 337 – 338 Children of More Caring, Less Controlling Parents Live
central amygdala (CeA) 53 Happier Lives 359
central nervous system (CNS) 182 Children-of-Twins (COT) studies 132 – 133,
Centuries of Childhood (Ariès) 289, 322 – 323 148 – 149
Cercopithecus mitis stuhlmanni 98 Childs, C. 452
cGxE (candidate gene-by-environment interaction) child sacrifice 297
128, 134, 141 – 142 child-twin based designs 129, 130, 133, 136,
Champagne, F. 50, 57 140 – 147
chance 366 chimpanzees 9 – 15, 78 – 91, 86, 94 – 96, 107 – 111
Chao, R. 449 – 450, 460, 463, 466 Chin, E. 184
Charles, A. 175 China 324 – 328, 337, 341 – 342, 404, 448, 451 – 459,
Chaudhary, N. 465 462 – 465
Cheah, C. 457 – 458 Chinese American families 451, 455 – 461
chemosensory functions 43 – 45 choes 307
chemosignaling 227 Christenson, S. 408
Chen, C. 455 – 456 Christianity 297, 310, 313, 340
Chen, H. 462, 465 CHRM2 151
Chen, W. 53 Chung, H. 493
Chen, X. 451 – 455, 462 – 465 Churchill, W. 287
Chernobyl disaster 183 Cicero 315
Child Behavior Check List 183 Cilissa 303, 306
childbirth: ancient history and 297 – 298, 304 – 306, 311; circumcision 298
macaques and 96; orientation in 82; rates 10, 14 Civil Partnership Act 2004 (United Kingdom) 273
childcare: ancient history and 298, 313 – 316; Clark, C. 90 – 91
environment of evolutionary adaptedness and 9; in cleft lip and palate 260 – 261, 261, 266
neighborhoods 389; parental investment and 7 – 8, climate 495 – 496
15 – 17 Clouds,The (Aristophanes) 306 – 307
522
Index
clozapine (CLZ) 52 – 53 Cote, L. 460

CNS (central nervous system) 182 Côte d’Ivoire 110
Coall, D. 175 COT (Children-of-Twins) studies 132 – 133,
Coan, J. 268 148 – 149
cognitive abilities 10, 137, 251 – 253, 400, 403, cotton-top tamarins 103 – 104
462 – 464 counterfactual models 351 – 352, 352 – 353, 355
Cole, P. 449 – 450, 459 cousins 132
Coleman, J. 423 CPP (conditioned place preference) tests 44 – 45
collective efficacy theory 376 – 377, 382 – 383, 387 CRC (Convention on the Rights of the Child) 350
collective socialization 377 – 378, 384, 464 Creation of Patriarchy,The (Lerner) 290
collider bias 364 CRF (corticotrophin-releasing factor) 36
Collishaw, S. 465 CRH (corticotropin-releasing hormone) 44,
colobus monkeys 96, 99 – 100 180 – 181, 186, 196, 230
Colorado Adoption Project (CAP) 125, 142 – 144 cribs 341
Colorado Sibling Study (CSS) 142 crime 153, 333, 376 – 386
Combs-Orme, T. 486 Crnic, K. 487
Committee on Developments in the Science of Cronin, P. 39
Learning 480 cross-cultural research 449, 452 – 453, 456 – 466
Common Sense Book of Baby and Child Care,The cross-fostering approach 56, 235
(Spock) 334 cross-lagged approaches 139 – 140, 143, 147 – 148
community groups 492 – 493 cross-sectional studies 270, 361, 365
compensatory model 382 – 383, 386, 388 Crouter, A. 486, 496
competence 78 – 80, 110, 141, 429, 481 crowding 487 – 489, 496
competition 7 – 11, 105 crying 83, 94, 97, 261 – 262, 306
Comstock, A. 331 CSS (Colorado Sibling Study) 142
COMT (catechol-O-methyltransferase) 136 CT (cortisol) 168, 179 – 183, 186, 194 – 196, 221 – 224,
concerted cultivation 425, 429 227 – 238
conditioned place preference (CPP) tests 44 – 45 C-tactile afferents 263
conduct disorder 127, 384 cultural anthropology 448 – 449
conflict 104 – 105, 145 – 151 cultural-historical perspective 452 – 453
confounding 354, 362 – 365, 363 cultural history 287
Confucianism 341 – 342, 455 culture 448 – 473; environment and 474 – 518;
confundere 362 evolutionary perspectives and 5 – 8; Roman family
conspecifics 10, 47, 108 – 109 life and 310 – 311; socioeconomic status (SES) in
Consumer Expenditure Survey 431 438 – 439
contextual-developmental perspective 452 – 453 culture and personality school 448
control, parental 142 – 143, 146, 322, 458 – 465 culture differences 254
Convention on the Rights of the Child (CRC) 350 cumulative risk 356, 424
cooperative breeding 14, 222 current-future trade-off 172 – 173
cooperative care 78 – 79 curvilinear models 431, 501
cooperative-compliant behavior 448 cuteness 18, 251, 257 – 263, 258, 268, 272
coparents 235 – 236, 487 cytosine 170
coping 177 – 178 Czilli, T. 264
Cornelia 313 – 314
corporal punishment 139, 350, 385, 427 DA (dopamine) 34, 49 – 56, 185, 235, 274
correlated risk 356 dACC (dorsal anterior cingulate cortex) 235 – 236
correlational studies 32, 125, 377 – 378, 424 Daltabuit, M. 495
cortex 46 Daly, M. 22
corticosterone 41, 184, 220 Darwin, C. 4, 7 – 8, 20, 23, 221; The Expression of the
corticotrophin-releasing factor (CRF) 36 Emotions in Man and Animals 252 – 253; On the
corticotropin-releasing hormone (CRH) 44, Origin of the Species by Natural Selection 169
180 – 181, 186, 196, 230 Davidson, K. 482 – 483
cortisol (CT) 168, 179 – 183, 186, 194 – 196, 221 – 224, Davis, A. 422
227 – 238 Davis, E. 183
cortisol awakening response (CAR) 230 Davis-Kean, P. 402, 405 – 410
cortisone 180 day care 328 – 329
Corwyn, R. 485 DBDs (disruptive behavior disorders) 183
Costa Rican families 460, 484 DBP (dibutyl phthalate) 188 – 189
523
Index
De Almeida, R. 52 directiveness 458 – 461, 464

Dearing, E. 384 discipline: ancient history and 303, 309; culture and
Deater-Deckard, K. 458, 464 456 – 461, 464; epidemiology of 350; genetics and
Deborah 299 136 – 139, 146 – 150; modern history and 321 – 323,
deconstruction 287 – 288 328; neighborhoods and 377 – 378, 385 – 387;
DEHP (di(2-ethylhexyl) phthalate) 188 – 189 parental educational attainment and 407
dehumanization 272 disgust 272
Deir el-Medina 292 – 293 Disney 332
Del Giudice, M. 18 disruptive behavior disorders (DBDs) 183
delinquency 148 distal behaviors 457
Demand, N.: Birth, Death, and Motherhood in Classical distress calls 40, 104, 262 – 263
Greece 304 – 305 diversity 388 – 391
deMause, L.: The History of Childhood Quarterly 289 divorce 311, 316, 338
democratic culture 343, 426 Dixon, S. 314 – 315
demographic transition 324 – 327, 372 dizygotic twins (DZ) 129 – 132, 145 – 146
Demos, J. 321 – 322 DNA 56, 123, 134, 169 – 170, 184 – 188
Denmark 190, 195, 198 Doane, H. 39
Denney, D. 125 dodecyl propionate 38
dentate gyrus (DG) 53 Dodge, K. 456, 464, 486
DEP (diethyl phthalate) 188 DOHaD (Developmental Origins of Health and
dependency: of children 3 – 5, 250 Disease) 175 – 179, 183, 192 – 194
depression: hormones and 233 – 235; parental dominance status 99, 110
negativity and 143; postnatal (PND) 196, 271 – 272; domus 310 – 317
postpartum 236 – 237; stress and 128, 183; weaning Dong, Q. 455 – 456
and 91 D’Onofrio, B. 153
descriptive epidemiology 349 – 351 dopamine (DA) 34, 49 – 56, 185, 235, 274
determinism 5 – 6, 351 dopamine D4 receptor (DRD4) 136
developing countries: antenatal care in 192 dopamine receptor D2 (DRD2) 137, 140 – 142
development: consequences of delaying 13 – 15; dorsal anterior cingulate cortex (dACC) 235 – 236
culture and 452 – 453; influences on 55 – 56, dorsal raphe (DR) 53
235 – 236; maternal effects and 168 – 173; maternal dorsal riding 87 – 88, 105
stress and 177 – 184; neighborhoods and 383; Dotterer, A. 407
outcomes 173 – 184; phthalates and 189; Plato DRD2 (dopamine receptor D2) 137, 140 – 142
and Aristotle on 303 – 304, 304; prenatal toxins, DRD4 (dopamine receptor D4) 136
teratogens, and 187 – 191; Quintilian and Augustine Drug and Alcohol Dependence 135
on 313, 314; socioeconomic status (SES) and Dubowitz, H. 486
421 – 423 Duncan, G. 402
developmental events 124 Durbrow, E. 482
developmentalists 5 – 6, 142 Dutch Hunger Winter 170, 177, 185 – 186
Developmental Origins of Health and Disease DZ (dizygotic twins) 129 – 132, 145 – 146
(DOHaD) 175 – 179, 183, 192 – 194
developmental plasticity 174 – 177, 199 early childhood 135 – 142
developmental psychology 449 – 450 Early Childhood Longitudinal Study, Kindergarten
developmental systems approach 5 – 6 Cohort (ECLS-K) 431
developmental timing: individual differences and Early Childhood Longitudinal Study–Birth cohort
388 – 390 (ECLS-B) 136 – 138
development studies 123 – 165 Early Growth and Development Study (EGDS) 137 – 144
de Waal, F. 104 East Asia 321, 330 – 332, 340 – 342, 454 – 457
DG (dentate gyrus) 53 Eastern European parents 464 – 465
dibutyl phthalate (DBP) 188 – 189 ecometric standards 373
Dick, D. 151 ECOT (Extended Children-of-Twins) studies
diet 15, 39, 95, 176 – 177, 294, 299 132 – 133, 133, 148 – 149, 152
diethyl phthalate (DEP) 188 Ecuador 439
differential parental investment 20 – 21 education: ancient history and 294, 299 – 300,
differential selection 357 311 – 313; environment and 481 – 483; modern
differential susceptibility theory 151 history and 320 – 322, 325, 329 – 331, 335, 344;
direct control practices 427 neighborhoods and 371; parent attainment of
direct effects models 430 – 432 400 – 420; socioeconomic status (SES) and 436 – 437
524
Index
Edwards, C. 181, 449, 454 Evans, G. 480, 487

EEG (electroencephalography) 258, 265 – 267, evaporation model 383 – 388, 386
271 – 273 Eve 297
EGDS (Early Growth and Development Study) event-related potential (ERP) 267, 270
137 – 144 evolutionary-developmental framework (evo-devo)
eggs, production of 7 168 – 171
Egypt: ancient 288 – 297, 300, 316 – 317; modern 342, evolutionary perspectives 3 – 29, 168 – 173
458, 497 ewes 38 – 39, 43, 176
Ehrenberg, M. 290 – 291 excitatory neural system 46 – 49
Eisler, R. 290 – 291 exercising behaviors 108
Elam, K. 139 Exodus 297 – 299
electroencephalography (EEG) 258, 265 – 267, expectations 405 – 407, 425
271 – 273 experiential effects, maternal behavior and 40 – 45,
electronic media 481, 487 – 491 53 – 55
elevated plus mazes 33, 44 experts, authority of 334 – 335
Elias, N. 322 – 323 exploitation 336 – 339
Eller, C. 291 exposure, of newborns 302 – 303
Elo, I. 401 Expression of the Emotions in Man and Animals,The
Embu 497 (Darwin) 252 – 253
emotions: culture and 457 – 458; Darwin on 253; Extended Children-of-Twins (ECOT) studies
dysregulation of 272; expression of 448; of new 132 – 133, 133, 148 – 149, 152
nonhuman mothers 35 – 36; parental educational extended investment model 430
attainment and 407; prenatal stress and 183 – 184 externalizing 138 – 139, 147 – 151, 463 – 464
empathy 235 – 236, 272 extirpation and replacement strategies 32 – 34
encephalization quotient (EQ) 12, 12 eye contact 251 – 252, 484
endocrine fit 222, 232 – 235 Ezekiel 298
endocrine system 34, 41
endorphins 34 faces/facial expressions 18, 84 – 86, 253, 257 – 262,
England 138, 193, 465 260 – 261
Enlightenment values 340 – 342 factor analytic techniques 358
environment 4 – 5, 474 – 518; ambient 496 – 497; built familia 310 – 312
497 – 498; community 498 – 499; of evolutionary families: ancient history of 287 – 319; evolutionary
adaptedness 9 – 10; genetics and 123 – 136, 139, perspectives and 3 – 4, 10 – 12, 15; modern history
139 – 145; parental educational attainment and of 320 – 348; neighborhoods and 371 – 399;
404 – 408; parental sensitivity and 135 – 137, 141 – 142 neurobiology and 273
environmental chaos model 431 familism/familismo 378, 456, 461, 464
Environmental Risk Longitudinal Twin Study Family and Communities Health Study (FACHS)
(E-Risk) 138 – 139, 146, 380 – 381 382 – 385, 390
environmental stressors 167 – 168 family design methods 364 – 365
epidemiology 349 – 370 family investment model 379 – 381, 385 – 387, 481
epigenetics 4 – 6, 57 – 58, 169 – 171, 186 – 187 Family Life Project 424
EQ (encephalization quotient) 12, 12 Family Life Project Key Investigators 494
ERP (event-related potential) 267, 270 family planning 302 – 303
Espinosa, M. 478 family process studies 152 – 153, 404
Esther 297 family stress models 379 – 381, 385 – 387, 431
estradiol 34 – 39, 47 – 49, 194 fast thinking 256
estrogen 44, 50 fathers: ancient history and 290 – 293, 296, 299 – 301,
estrous cycles 42 304 – 306, 310 – 311, 315 – 316; culture and 462, 465;
ethnicity 154, 272, 386, 389 – 390, 422, 428 educational attainment of 400 – 401; environment
ethnographic methods 422, 449 and 9, 485 – 486; hormones and 221 – 235; labor
ethnotheories 411 and 327 – 329; modern history and 339 – 340;
Etienne, R. 289 neighborhoods and 377, 383; parental investment
eugenics 309 and 7 – 8, 16; see also males
Europe 345 – 346, 458, 483 – 484, 495 Fearon, R. 136
European American families 378, 384 – 390, 406 – 407, Feinberg, M. 150
422, 455 – 463 Feldman, R. 454
Eutherian mammals 173 females: adolescence and 95 – 96, 100; cooperative
evaluation-response process 453 care and 78 – 79; historical narrative and 287;
525
Index
infant cues and 269; parental investment and 3 – 8; GCs (glucocorticoids) 36, 39, 175, 179 – 186, 198
patriarchal state and 290 – 291 Gemeinschaft 464
feminism 288, 291, 346 gender 269, 287 – 293, 328 – 329, 389, 478
fertility rates 16 – 17 gene-by-intervention research 154
fetal alcohol syndrome 261 gene-environment correlations (rGE) 124 – 156,
fetal development 166 – 168, 175 – 177, 181 – 187, 129, 365
197 – 198 gene-environment interplay (GE) 124 – 130, 138, 153
fetuses 166, 171 – 173, 174, 192 – 193 gene expression 57, 170 – 171, 186
FFA (fusiform face area) 266 generalizability 131 – 132, 391, 438 – 439
fight or flight response 179, 231 generic parents 314 – 316
filial piety 455 Genesis 297
financial capital 423 – 424, 436 – 437 genetically informed designs 123 – 126, 129,
Finland 17, 183 133 – 137, 438
FinnTwin study 145 – 150 genetics 4 – 8, 56 – 58, 123 – 165
Fisk, N. 180 genetic variants/genetic markers 134 – 137
fitness, reproduction and 4 – 7, 169, 172 gene x environment interactions (GxE) 124 – 128,
Fivush, R. 449 – 450, 459 134 – 138, 141, 144 – 145, 149 – 154
Fleming, A. 36, 43 – 50 Gennetian, L. 413
fMRI (functional magnetic resonance imaging) 252, genomes 5, 24, 128
257 – 258, 260 – 261, 263, 273 genome-wide association studies (GWAS) 124, 134,
folate (folic acid) 185 149, 153
food deserts 478 – 479 genotypes 32, 55 – 58, 125 – 126, 134, 149 – 151,
food/feeding: availability 478 – 479; aversion 193; 167 – 170
environment and 479; nonhuman primates and geographic information systems (GIS) 374
83, 89 – 92, 104; parental investment and 8; see also Georgia 338
hunter-gatherer societies Germany 145, 343, 458 – 459, 465, 476, 484
foraging 17, 92. see also hunter-gatherer societies germ cells 56 – 57
Former Soviet Union 464 germ theory 325
Forsman, M. 147 – 148 Gesellschaft 464
fossil records 9, 14 gesturing 87, 92
foster caregivers and parents 33 – 34, 99, 290 gibbons 85, 93 – 96, 94
France 321 – 323, 329 Gibson, E. 481
French, D. 452 – 453 Gillis, J. 322
Freudian psychoanalytic theory 448 Gimbutas, M. 290 – 291
Friedlmeier, W. 449 – 450 girls 14, 183 – 184, 384 – 386, 389
frontal lobes 252, 257 GIS (geographic information systems) 374
Fuller, J.: Behavior Genetics 124 – 125 Gitau, R. 180
functional magnetic resonance imaging (fMRI) 252, globalization 451
257 – 258, 260 – 261, 263, 273 Glover,V. 180, 182
Fung, H. 459 glucocorticoid receptors (GRs) 180 – 182, 186, 223
Fun with Dick and Jane 289 glucocorticoids (GCs) 36, 39, 175, 179 – 186, 198
furies, the 288 glutamatergic neurotransmission 53
fusiform face area (FFA) 266 Glynn, L. 183
goal emulation 90
G1219 cohort 146 – 147, 150 goals, socioeconomic status (SES) and 425
GABA 34 – 36, 51 – 53 God 296 – 300, 317
GABRA2 151 goddess-hypothesis 291
Gall, J. 264 Godfrey-Smith, P. 169
Gallup Poll 499 Godinot, F. 40
games 293 – 294, 307 – 308, 315, 434 golden lion tamarins 104
Ganiban, J. 150 Golding, J. 182
Gao, J. 53 gonadal steroids 229
García Coll, C. 449 – 450 Goodall, J. 85
Gardner, F. 465 Goodison, L. 291
Garrett-Peters, P. 494 Goodnow, J. 403
gathering. see hunter-gatherer societies Google Street View 373
Gaza 487 Gopnik, A. 23
gaze 84 – 87, 86, 97, 101, 104, 225 – 227, 233, 261 Gorilla gorilla 93
526
Index
gorillas 84 – 85, 90 – 95, 93, 111 Havighurst, R. 422

Gottesman, I. 153 Havill,V. 455
government, parenting and 324, 333 – 334 Hawkes, K. 17
Gracchi 313 Hayashi, K. 495
grades, responses to 409 – 410 HCC (hair cortisol concentrations) 231, 234
grandmother hypothesis 17 hCG (human chorionic gonadotropin) 194
grandparental support 16 – 17 HDI (Human Development Index) 475
grasping 83, 89, 96 Head Start 391, 493
gray matter 107 – 108, 270 health, environment and 477 – 479
gray mouse lemurs 106, 106 Heath, A. 127
great apes 80, 80 – 96, 81, 93, 111 Heath, S. 422, 426
Great Recession 372 Hebrew Bible 296 – 300
Greek families 460, 484 Hellhammer, D. 236
Greenberger, E. 455 – 456 helping mode of parenting 289
Greenfield, P. 452, 464 – 465 hemoglobin 184
greeting 88, 106 heritability 4, 123 – 130, 135 – 154, 170, 438
Greven, P. 322 Herman, G. 268
grief 338 Heron, J. 182
grooming 39, 85, 88 – 89, 92, 95 – 98, 105 Herzegovina 350
group-orientation 454 – 455, 464 – 465 Heston, L. 125
GRs (glucocorticoid receptors) 180 – 182, 186, 223 heterochrony 13
Guatemala 483 HG (hyperemesis gravidarum) 193 – 194
Gunung Palung 84 higher-order capacities 272
Gusii 484 high-risk populations 138, 236 – 238
GWAS (genome-wide association studies) 124, 134, Hill, N. 409
149, 153 hippocampus 46, 52, 109, 182
GxE (gene x environment interactions) 124 – 128, Hippocrates: Aphorisms 308; On Dentition 308
134 – 138, 141, 144 – 145, 149 – 154 Hippocratic writings 305, 308, 315
histone modification 170 – 171
Hackett, L. 456 Historia Naturalis (Celsus) 315 – 316
Hackett, R. 456 history: ancient 287 – 319; of behavioral genetics
Hadza 17, 22 124 – 128; of culture and parenting 448 – 450;
haemochorial placenta 166 modern 320 – 348; of neurobiology 252 – 254
Haidt, J. 268 History of Childhood Quarterly,The (deMause) 289
Haig, D. 173, 176 Hmong American families 460
hair cortisol concentrations (HCC) 231, 234 Ho, C. 463
Haley, A. 153 Ho, D. 462
half-siblings 132 Hoekzema, E. 270
Halle, T. 406 Hoff, E. 406, 480, 485
Hallett, J. 315 Hoffman, L. 403
Halpern, R. 351 Holden, G. 494
Halverson, C. 455 Holland, J. 495
hamadryas baboons 100 Hollingshead Four-Factor Index 402, 423 – 434
Hammurabi 295 Holmboe-Ottesen, G. 478
Hannah 297 – 298 home-based parent involvement 409 – 410
Hannibal 313 home environment 474 – 518
Hannigan, L. 149 HOME inventory 423, 428, 484 – 486
Hansen, S. 49 homelessness 488
Hanuman langurs 99 homeostasis 104, 179 – 180, 230
Harding, K. 56 Homer 303
harems 90, 111 homework 409
Harkness, S. 449 – 452, 458, 476 hominins 9 – 14
Harlow, H. 30 Homo genus 9 – 10, 14 – 15
Harold, G. 365 Homo Ludens (Huizinga) 323
Harris, K. 486 homophily 502
Hart, B. 406, 428, 480, 485 Homo sapiens 3 – 14, 17, 23
Hart-Shegos, E. 479 Honduras 351
Harwood, R. 460 Hong Kong 404, 463
527
Index
Hoover-Dempsey, K. 411 inconsistency 144, 149

hoo vocalizations 87 indenture 321, 337
hopelessness 497 India 329 – 330, 342, 449, 454, 458, 465, 483 – 384
Hopkins, W. 108 indirect effects models 430 – 434
hormones: changes and 23, 32; dopamine (DA) and individualism 340 – 342, 455, 464 – 465
50; of human parenting 223; morning sickness Indonesia 458
and 193 – 194; neuroendocrinology and 220 – 249; industrialization 320, 327 – 328, 331, 338
nonhuman mammals and 30 – 53; priming 31, 35, inequality 421, 424 – 425
42, 58 – 59; stress 167 – 168 infancy: ancient history and 304 – 306; communicative
Horwath, J. 492 cues and 250 – 251; dependency of 250;
hostility 135, 138 – 140, 143 – 144, 365, 456 developmental studies of 135 – 142; differential
howlers 100 – 101, 104 parental investment and 20 – 21; in nonhuman
HPA (hypothalamic-pituitary-adrenal) hormones 36, primates 79 – 111, 91; parental instinct and
167, 179 – 183, 186, 194, 230, 233 – 237 250, 253, 268, 272; perceptual biases of 18 – 19;
HPG (hypothalamic-pituitary-gonadal) axis 228 separation anxiety in 88; temperament in 110
Hrdy, S. 21, 291 Infant Care 325, 334 – 335
HSD enzymes 180 – 182 infant cues 259, 268, 269 – 275
Huizinga, J.: Homo Ludens 323 infant faces 257 – 261
human capital 423, 436 – 437 infanticide 4, 20 – 22, 82, 98, 289, 293, 302 – 303
human chorionic gonadotropin (hCG) 194 infant mortality 10, 17, 304, 324
Human Development Index (HDI) 475 infant schema (Kindchenschema) 18, 253, 257, 266
Human Genome Project 128 inhibitory neural system 46 – 49
humans: childbirth in 82; evolutionary perspectives initiative-taking 451 – 454
and 9 – 15; hormones and 31; maternal competence injuries 479 – 480, 490
in 80, 110; neurobiology and 250 – 284; newborn innate releasing mechanisms 253, 266
period of 82 – 84; separation anxiety in 88; social insecure attachment 19 – 20, 104; see attachment
neuroendocrinology and 220 – 249 instability 480, 485, 488
hunter-gatherer societies 9 – 10, 15 – 16, 22 – 23, 290 instincts 250, 253, 268, 272, 291
Huntsinger, C. 455 institutional resources 391
Huntsinger, P. 455 insulin 190, 198
hygiene 325 – 327 integrated models 386, 386 – 388
Hylobates lar 94 interactive models 381, 381 – 386
hyperactivity 144, 183 intergenerational phenotypic inertia 170 – 171
hyperemesis gravidarum (HG) 193 – 194 internalizing 144, 148 – 149, 384 – 385, 463
hyperhomocysteinemia 185 International HapMap Project 128
hyperscanning 273 International Journal of Epidemiology 354
hyperthermia 41 internet access 481
hypothalamic-pituitary-adrenal (HPA) hormones 36, interoception 235
167, 179 – 183, 186, 194, 230, 233 – 237 intersubjectivity 251 – 252
hypothalamic-pituitary-gonadal (HPG) axis 228 intranasal OT (oxytocin) 226 – 229
hypothalamus 36, 44 – 46, 57, 179 – 180, 224, 235 intrauterine environment 132, 170, 174 – 177,
186 – 187
Ichabod 298 intrauterine growth restriction (IUGR) 175
iconography 292 intrusiveness 141, 230, 460, 464
illegitimacy 338 – 339 intuitive parenting 107 – 110, 251 – 254, 266 – 267
illness, ancient history and 294 – 296, 308, 315 – 316 in vitro fertilization (IVF) 194 – 195, 365
imitation 14, 84, 90 IQ 189
immaturity 3 – 5, 15, 23 iron deficiency 184 – 185
immigrants 327 – 329, 332 – 333, 337, 388 – 389 Isaac 297
immune biomarkers 221 – 223 Isaiah 297
immunohistochemistry 47 – 48, 53 Islamic world 324, 340 – 342
immunological responses 108 isolation rearing 109
imperialism 336 – 337 Ispa, J. 458 – 461, 464
imprinted genes 57 Israelis 454, 464
impulsivity 139, 149 Italy 458
inattention/hyperactivity 144, 183 IUGR (intrauterine growth restriction) 175
inclusive fitness 4 – 7 IVF (in vitro fertilization) 194 – 195, 365
income effects 436 – 437 Izard, C. 485
528
Index
Jacka, F. 186 Kohnstamm, G. 455

Jaffee, S. 476 – 477, 491, 499 Kokwet 476
Jamaica 350, 482 Komisaruk, B. 48
Jansson, T. 186 Korean American students 462
Japan 138, 151, 191, 320, 325; advice literature in Korja, R. 183
334; childlessness in 345 – 346; consumerism in Korsmit, M. 47
332; culture in 257, 449, 454, 458 – 460; education Kramer, M. 175
in 330; environment and 498; individualism in Krasnegor, N. 107
341; labor in 327 – 328, 337; parental educational Kringelbach, M. 265 – 266, 272
attainment in 404; state authority in 333 Kummer, H. 97
Japanese American families 460 Kung 484
Japanese macaques 99 Kuo, F. 496
Jarrett, R. 383 kurios 300 – 302
Jason 302 Kurtz-Costes, B. 406
Jaynes, J. 37 Kuzawa, C. 170 – 171
Jenkins, J. 463
Jeremiah 297 – 299 Labartu 296
Jerusalem 296 – 299 labor 297 – 301, 321, 324 – 329, 337 – 339
Jewish tradition 297 – 298. see also ancient Israel lactation 34 – 36, 41, 106
Jimerson, S. 482 L.A. FANS (Los Angeles Family and Neighborhoods
Joffe, T. 15 Study) 376, 384
Johnson, A. 494 Lamborn, S. 460
Jones, E. 482 lambs 38 – 40, 43
Jordan 458 Lamentations 298 – 299
Jose, P. 455, 460 – 461 Lamm, B. 465
Journal for the History of Childhood and Youth 324 Land of Punt 291 – 292
Journal of Abnormal Child Psychology 135 language 405 – 406, 428 – 429, 485
Jungian psychology 290 langurs 97 – 99
juvenile period 10, 79 – 80, 94 – 107 Lansford, J. 350
Lareau, A. 422, 425, 429
Kağıtçıbaşı, C. 449 – 451, 464 – 465 Larsson, H. 147 – 148
Kahneman, D.: Thinking Fast and Slow 256 Lasch, C. 334
Kalil, A. 432 latchkey children 489
kangaroo care (KC) 238 lateral habenula (LH) 49
Kanyawara community 87 lateral OFC (lOFC) 266
Kaplan, H. 15 lateral septal nucleus (LSv) 53
Kaplan, R. 496 lateral septum (LS) 47
Kasson, J. 323 Latin America 321, 324 – 326, 334, 338 – 339
kawaii 257 Latin American families 379, 382, 385 – 390, 434,
Keeble, S. 39 455 – 464, 479
Keio Twin Project 151 laughter 88, 262 – 263
Keller, H. 449 – 450, 455, 459 – 460, 465, 484 Lawrence, D. 476
Kendler, K. 153 Laws (Plato) 304
Kenya 449, 454, 458, 476 – 478, 484, 497 Leah 297
Kenyon, P. 39 learning 41 – 45, 53 – 55, 270 – 271, 483. see also
Kerr, M. 387 education
Ketambe 84 Lebanon 342, 458
Kibale National Park 82, 87 Lee, A. 45
Kidd, C. 13 – 14 Lee, B. 451
kidnapping 98 – 99 Lee, R. 360
Kim, K. 462 Lees, J. 492
Kim, P. 270 Leiden and London Twin Studies 135 – 136
Kindchenschema (infant schema) 18, 253, 257, 266 Lekies, K. 500
Kinsley, C. 39 lemurs 105 – 106
Knafo, A. 139 Leon, M. 39 – 41
knockout approaches 56 Leonard, W. 478
Kogos, J. 485 Lepore, S. 487
Kohn, M. 424, 437 – 438 leptin-adiponectin ratio 190
529
Index
Lerner, G.: The Creation of Patriarchy 290 MacKinnon, C. 486

letdown reflex 39 MacLean, P. 261
Leung, C. 457 – 458 Madagascar 106
Leve, L. 141, 154 Madamba, A. 460
Leventhal, T. 377 magnetic resonance imaging 33
lex talionis 295 magnetoencephalography (MEG) 252, 258, 265 – 268
Leydig cells 188 Magnuson, K. 407, 413
LFPs (local field potentials) 264 – 265 Magnusson, J. 45
LH (lateral habenula) 49 Mahale 82, 89
Li, D. 451 – 454 Mahoney, J. 406
Li, H. 483 maladaptive behaviors 140, 143, 453, 463
Li, J. 457 – 458 male-elite interests 288 – 290
Liaw, F. 455 males: adolescence and 95 – 96, 100; cooperative care
Libation Bearers,The (Aeschylus) 303, 306 and 78 – 79; historical narrative and 287 – 288;
Lichtenstein, P. 147 – 148 infant cues and 269; nonhuman primate 92 – 106;
Lichtman, C. 264 parental investment and 3 – 8, 16; patriarchal state
life course epidemiology models 354 – 356, 355 and 290 – 291; uncertainty of paternity and 8,
life history theory 167, 167 – 172, 175 – 177, 197 11, 90
limbic systems 44 – 49, 59, 109 Mali 351
Lindsay, R. 181 malnutrition 170
linear direct effects models 430 mammals 9, 18 – 20, 30 – 77, 222
lions 17 managerial control 427
Lipscomb, S. 139 mangabeys 97
literacy 137, 300, 428 – 429 Mann, J. 21
liveability 496 MAOA (monoamine oxidase A) 128
local field potentials (LFPs) 264 – 265 mapping tools 373
locomotor behavior 85, 90 – 93, 97 – 99, 104 marital satisfaction models 151
lOFC (lateral OFC) 266 marking 86, 105
Lomako 82 Marks, L. 465
Lomanowska, A. 56 marmosets 101 – 105, 103, 228 – 229
longevity 17 Marriage Act 2013 (United Kingdom) 273
longitudinal studies 126, 138 – 140, 147, 273, 361, Martin, A. 488
365, 382 – 384 Martin, J. 426
Longitudinal Study of Adolescent Health (Add Masalha, S. 454
Health) 382 – 384 Mason, J. 181
Longoli 484 Massachusetts 321 – 322
Lönn, S. 153 Massey, D. 377
Lonsdorf, E. 90 master-apprenticeship education 90
Lonstein, J. 56 mate guarding 11
Lorenz, K. 18, 253, 266 mater familias 310
Los Angeles Family and Neighborhoods Study (L.A. Mater Matuta 312 – 315
FANS) 376, 384 maternal alleles 57
Louie, J. 460, 463 Maternal Antenatal Attachment Scale 191
Louisville Twin Study 136 maternal behavior: aggression 52 – 53, 99, 104;
Love, O. 184 competence in 78 – 80, 110; deficits in 46, 51 – 53,
Low, S. 476 56 – 57; development of 55; experiential effects
lowland gorillas 92 on 41 – 45, 53 – 55; genetics of 56 – 58; hormonal
LS (lateral septum) 47 effects on 34 – 35; neuroanatomy and 35, 45 – 49;
LSv (lateral septal nucleus) 53 neurochemistry of 49 – 53; psychobiology of
Lucion, A. 52 nonhuman mothers and 30 – 77
Lumley, J. 177 Maternal Behavior in the Rat (Wiesner and Sheard) 33
Luria, A. 452 maternal birth weight 175
Lynd, H.: Middletown 422 maternal core 46 – 52
Lynd, R.: Middletown 422 maternal educational attainment 400 – 420
maternal effects 166 – 181, 167
macaques 13, 96 – 100, 108 maternal environment 171 – 173
Maccoby, E. 426 maternal experience effect 42
Machiavelli, N. 287 maternal grandparents 16 – 17
530
Index
maternal instinct 291 Mexico 326, 448 – 449, 452, 462

maternal investment 174 MFTS (Minnesota Family Twin Study) 145 – 150
maternalism 328 mice 38 – 40, 52, 56, 183 – 184
maternal mortality 192, 304 Michael, R. 403
maternal phenotypes 166, 171 Mickey Mouse 257
maternal stress 168, 177 – 184 Microcebus murinus 106
maternal styles 98 – 99, 109 – 110 microdialysis 32, 50
maternal values 288 – 291 middle childhood 142 – 145, 154
Matijasevich, A. 351 Middle East 324 – 326, 337 – 338
mating 3 – 4, 21 Middle Kingdom Egypt 292
matriarchy 290 – 291 Middletown (Lynd and Lynd) 422
Matsuzawa, T. 89 – 90 midwives 293, 297 – 298, 305
Maughan, B. 465 migration 337
maximin reproductive strategies 171 militarism 291, 296, 309 – 311
Mayan culture 454, 476 Millennium cohort 351
Maynard, A. 452 Mills-Koonce, R. 494
McDonald, M. 478 mindset, defined 429
McGillicuddy-De Lisi, A. 403 minimax reproductive strategies 171
McHale, S. 486, 496 Mink, I. 485
McLoyd,V. 486 Minnesota Family Twin Study (MFTS) 145 – 150
Meaney, M. 57 Mintz, S. 344
measurement bias 358 – 360, 365 mirroring 235
Medea 302 miscarriages 297
medial amygdala (ME) 48 – 49 misclassified twins 130
medial orbitofrontal cortex (mOFC) 265 – 267 Moberg, T. 147 – 148
medial prefrontal cortex (mPFC) 46, 51 – 53, moderators 127, 354, 434
271 – 272 modern history 320 – 348
medial preoptic area (MPOA) 46 – 55, 58 modernity 322 – 323
mediated models 354, 379, 379 – 381, 385, 386, modernization 322, 451, 464
431, 435 mOFC (medial orbitofrontal cortex) 265 – 267
mediation vs. confounding 362 – 364, 363 molecular genetic research 124, 127 – 128, 133 – 135,
medical care: ancient history and 294 – 296, 303 – 304, 140 – 142
308, 315 – 316; modern history and 326 – 327 monitoring behaviors: culture and 456, 456 – 461;
Medical Research Council National Survey of environment and 480, 490 – 494; genetics and
Health and Development 359 145 – 146, 149 – 151; neighborhoods and 377,
medieval history 287 – 289, 323 382 – 383, 387 – 389; in nonhuman primates 85,
Mediterranean civilizations 288 – 300 88 – 89, 93, 97 – 99; parent education attainment
MEG (magnetoencephalography) 252, 258, 265 – 268 and 411
Meinlschmidt, G. 236 monkeys 11, 30, 56
Melo, A. 56 monoallelic expression 57
memory 42 – 45 monoamine oxidase A (MAOA) 128
men. see fathers; males monogamy 11, 78, 85, 94, 101, 111
Menaghan, E. 437 monotheisms 291
menarche 14, 95, 99 monotropy 269
Mendel, G. 124 – 125 monozygotic twins (MZ) 129 – 132, 138, 143 – 148
Mendelian principles 57, 124 – 125 morbidity 17, 175, 194
mental health 182 – 184, 432 Mordecai 297
mentalization 272 Morgan, S. 486
Mervielde, I. 455 morning sickness 193 – 194
meshing 87 Morrell, J. 45
mesocortical dopamine system 49 – 52 Morris, C. 291
mesolimbic system 49 – 52 Morris, N. 153
metabolic syndrome 198 mortality rates: adverse childhood experiences and
methylation 170 – 171 197; ancient 297, 302 – 304, 311 – 313; maternal
methyl-binding proteins 184 192; modern 324 – 327; of nonhuman primates
methyl transferases 184 101; paternal investment and 16; rank of mothers
Mexican American families 378, 383, 412, 460, and 110; reproduction and 172; in rural areas 479
464, 486 Moses 297
531
Index
motherese 92, 252 National Park Service 498

Mother Goddess fertility figurines 290 – 291 National Survey of American Attitudes on Substance
mother-litter interactions 31 – 36, 45, 50 – 51 Abuse 490 – 491
mothers: ancient history and 290 – 293, 298 – 301, natural order 288 – 291
305 – 306, 309 – 310, 313 – 316; attachment and natural selection 3 – 4, 7 – 10, 169 – 172
19; differential parental investment and 20 – 21; nature/nurture dichotomy 3
educational attainment of 400 – 420; hormones and Naunakhte 293
221 – 235; infant cues and 270; labor and 327 – 329; nausea and vomiting in pregnancy (NVP) 193 – 194
maternal effects and 166; modern history and NCES (National Center for Educational Statistics)
339 – 340; neighborhoods and 375 – 386; parental 402 – 403
investment and 6 – 8, 15 – 17; psychobiology of NEAD (Nonshared Environment in Adolescent
nonhuman mammal 30 – 77; as social partners 88; Development) 146 – 151
see also females; maternal behavior Neanderthals 14
motivation, maternal 49 – 53 Nefertiti 292
motivational entity model 253, 256, 266 negative amplification model 383 – 385
motivation-based mechanisms 410 – 411 negative controls 364
motor areas 257 negative parenting 135 – 151, 360
motor control 52, 84 – 87 negative/risky environments 126 – 128
motor skills 107 neglect 4, 20 – 22, 479
mountain gorillas 84 neighborhoods 371 – 399, 487 – 497
moving studies 375 – 377 neocortex 11 – 15, 46
Moving to Opportunity for Fair Housing Neolithic Europe 290
Demonstration (MTO) 375 – 377 neophobia 35 – 36
mPFC (medial prefrontal cortex) 46, 51 – 53, neoteny 257
271 – 272 Nepal 185, 459
MPOA (medial preoptic area) 46 – 55, 58 nepios 303
mRNA 44 Nesteruk, O. 465
multimodal cues 87 nests 37 – 39, 51 – 53, 105
multiparous mothers 40 – 43 Netherlands 135, 189, 382, 458
multiple births 194 Nettle, D. 199
Murray, A. 253, 256, 266 Neuman, C. 478
music 434 neural activity 268
mutual gaze 84 – 87, 86, 97, 101, 104 neural plasticity 221
mutual shaping 89 neural responses 270
myelination 185 neuroanatomy 35, 45 – 49
Myth, Religion, and Mother Right (Bachofen) 290 neurobehavioral integrity 84
MZ (monozygotic twins) 129 – 132, 138, 143 – 148 neurobiology 222, 250 – 284
neurochemicals 11, 32 – 34
N1 responses 270 neurochemistry 49 – 53
N2CAP (Northeast-Northwest Collaborative neurocognitive outcomes 185
Adoption Project) 137 – 139 neuroimaging 273 – 274
N2 responses 267 neuropeptide oxytocin 40
N100 component 265 neuropeptides 33, 36, 227
N170 267, 270 neurotransmitters 34, 52 – 53, 134
NA (nucleus accumbens) 46, 49 – 53, 235 newborn period 79 – 85, 96 – 97, 100 – 101, 105, 298
Nakamura, K. 404 New Kingdom 292 – 294
Namibia 484 new world monkeys 80, 81, 100 – 105, 111
naming practices 293, 298, 302, 305, 341 New Zealand 194
Narusyte, J. 148 Nguyen, J. 460
Nast, I. 236 NHES (National Household Education Survey)
National Association of State Park Directors 498 498 – 499
National Center for Educational Statistics (NCES) nicotine receptors 194
402 – 403 Nielsen, M. 14
national characters 449 Nigeria 479
National Household Education Survey (NHES) night monkeys 103 – 104
498 – 499 Nile River valley 292 – 294
National Longitudinal Study of Adolescent nipples 40, 47, 83, 90, 96 – 97
Health 151 Nishida, T. 88 – 90
532
Index
No Child Left Behind 408 ōmēn 297

nomadic groups 290 On Dentition (Hippocrates) 308
nondifferential selection 357 On the Origin of the Species by Natural Selection
nonhuman mammals 20, 30 – 77 (Darwin) 169
nonhuman primates 78 – 122 ontogeny 6, 14, 78
nonlinear direct effects models 430 oppositional behavior 139
nonpatriarchal states 290 – 291 optogenetics 33, 57
Nonshared Environment in Adolescent Development orangutans 84 – 85, 89 – 90, 91, 95 – 96
(NEAD) 146 – 151 orbitofrontal cortex (OFC) 252 – 258, 262 – 272
nonsystematic measurement error 358 Orestes 288, 303, 306
nonverbal techniques 459 organs, marketing of 337
nonvisual cues 251 Oribasius 316
nonvisual neocortex 15 orientation 83 – 84
norepinephrine 34, 185 orienting system 251 – 257, 266
normativeness 466 Orpen, B. 43 – 44
North America 421 – 422, 448 – 450, 455, 458, 463 orphans 297, 333, 351 – 352, 356
Northeast-Northwest Collaborative Adoption Osaka 495
Project (N2CAP) 137 – 139 Osgood, D. 486
Norway 127, 186, 194 other-race effect 272
Nso 458 – 460, 465 Otto, H. 459
Nubia 291 out-groups 272
nuclear families 301, 310 – 311 overinvolvement 148 – 149
nucleus accumbens (NA) 46, 49 – 53, 235 overload, crowding and 487
nulliparous rats 36 – 38 overnutrition 184
Numan, M. 46 – 47 overprotectiveness 145
nursing: ancient history and 298; diet and 39; overreactive parenting 139
dopamine (DA) and 51 – 52; grandparental support ovulation 294
and 17; infancy and 14, 89, 94; newborns and 83; own vs. other infant studies 270 – 271
parturition and 33, 41; Pleistocene and 10; touch oxytocin (OT) 33 – 36, 39 – 41, 46 – 49, 52, 56 – 57,
and 39 – 40 220 – 229, 232 – 237, 262, 274
nurturant behaviors 23, 80, 108 – 110, 449 oxytocin receptor (OXTR) 136 – 137, 140,
nutrition 168 – 171, 184 – 186, 192 – 194. see also 224 – 226, 236
food/feeding
NVP (nausea and vomiting in pregnancy) 193 – 194 P1 responses 269 – 270
P3a responses 270
obesity 184, 198, 496 P300 267, 271
objective stress 195 – 196 PAG (periaqueductal gray) 47, 235, 264 – 265, 268
object manipulations 92 paidagogues 308
object salience 228 pair bonding 11, 85, 100, 226 – 228
observation, performance bias and 360 Paleolithic Europe 290
observational measures 136, 142, 146 Palestinians 454
observational studies 449, 458 Panesar, S. 36
obstetrics 305 Pan troglodytes 86
occupational status 423 – 424 Papousek, H. 266
occupation effects 436 – 437 Paraguay 480, 489
O’Conner, T. 182 paralinguistic cues 459
odor cues 37 – 40, 44, 264, 273 parasites 294
OFC (orbitofrontal cortex) 252 – 258, 262 – 272 paraventricular nucleus (PVN) 36, 46
Ohlsson, H. 153 Parcel, T. 437
oikos 300 – 302, 310 parental authority 138, 332 – 333, 383 – 386, 426,
Okazaki, S. 457 450, 461
old age 10, 293, 299 parental bias, in gene expression 57
Old Kingdom 292 Parental Bonding Instrument 359
old world monkeys 80, 81, 96 – 100 parental brain 256 – 257, 260 – 261
olfactory bulbs 37 – 38, 48, 52 parental care 3 – 8, 13 – 17
olfactory cues 37 – 40, 44, 264, 273 parental evaluations 453
olfactory-mediated responses 36 – 40 parental instinct 250, 253, 268, 272, 291
olfactory systems 46 – 49, 59 parental investment 3 – 8, 15 – 17, 20 – 21, 173 – 175
533
Index
parent-based twin designs 129, 130, 133, 142, PFHxS (perfluorohexane sulfonate) 189 – 190
145 – 147 PFNA (perfluorononanoic acid) 189
parent-child adrenocortical synchrony 233 – 235 PFOA (perfluorooctanoate) 189 – 190
parent-child oxytocin synchrony 233 PFOS (perfluorooctanesulfonate) 189 – 190
parenting: ancient history of 287 – 319; culture pharaohs 292, 297
and 448 – 473; environment and 474 – 518; PHDCN (Project on Human Development in
epidemiology of 349 – 370; evolution of Chicago Neighborhoods) 376 – 379, 384 – 385
3 – 29; genetics and 123 – 165; modern history phenotypes 124 – 134, 150 – 154, 166 – 171, 167,
of 320 – 348; neighborhoods and 371 – 399; 183 – 186
neurobiology of human 250 – 284; in nonhuman phenotypic plasticity 174 – 175
primates 78 – 122; parent education attainment Philippines 229, 449
and 400 – 420; prenatal 166 – 219; psychobiology Philistines 297
and 30 – 77; social neuroendocrinology of human Phillips, K. 264
220 – 249; socioeconomic status (SES) and Phinehas 297 – 298
421 – 447 phthalates 188 – 189
parenting variables 354 Physical Activity Council 499
parent-offspring adoption studies 126 – 127, 132 – 133, physical cues 18
137 – 138, 152 – 153 physical health 432
parent-offspring conflict theory 168, 171 – 173, 196 physiological stressors 195 – 196
Parents’ Magazine 334 Piantadosi, S. 13 – 14
parent talk 480 Pickles, A. 465
parity effects 42 – 45 pigtail macaques 99, 108 – 109
PARs (predictive adaptive responses) 170, 198 – 199 pituitary glands 34, 46
parturition 33 – 34, 37 – 43 place-based initiatives 371
parturitional hormones 45 – 46, 52 placental CRH (pCRH) 196
patas 98 placental efficiency 175, 186 – 187
paternal alleles 57 placental ratio 175 – 177
paternal care 220 – 221 placental weight 175 – 177
paternal effects 166 – 168 placentas 166 – 168, 173 – 177, 180
paternal grandparents 16 plasticity 151, 170, 174 – 177, 182, 199, 221, 267 – 269;
paternal investment 3 – 8, 16 in children 303 – 305, 308, 312
paternal phenotypes 166 Plato 302 – 310, 304, 313; Laws 304
paternity 8, 11, 90 – 92 play: ancient history and 298, 307 – 308, 315; culture
patria potestas 310 – 311, 315 – 316 and 449, 454; environment and 483 – 484, 496;
patriarchy 288 – 299, 316 epidemiology of 350 – 351; evolution of human
Paul of Aegina 316 intelligence and 14; modern history and 323,
Pawluski, J. 56 332; in nonhuman primates 86 – 88, 92, 97,
pCRH (placental CRH) 196 101, 105; social neuroendocrinology and 226;
peasants 292 – 294, 329 socioeconomic status (SES) and 428
pediatrics 327 pleasure cycle 254 – 256, 255, 270 – 271
peer relations 94, 109, 138, 371, 449, 464 pleasure network 255
Pener-Tessler, R. 140 Pleistocene 9, 168
peptides 41, 44, 228 Plomin, R. 126, 139
perceived stress 195 – 196 Plooij, F.X. 87
Pereira, M. 50 pluralist-constructivist perspective 464 – 465
perfluoroalkyl substances (PFAS) 189 – 190 Plutarch 312 – 315
perfluorohexane sulfonate (PFHxS) 189 – 190 PND (postnatal depression) 196, 271 – 272
perfluorononanoic acid (PFNA) 189 poleis 310
perfluorooctanesulfonate (PFOS) 189 – 190 policing 333
perfluorooctanoate (PFOA) 189 – 190 polis 300 – 302
performance bias 360 Politics (Aristotle) 304
periaqueductal gray (PAG) 47, 235, 264 – 265, 268 Pollock, L. 323
permissiveness 98 – 99, 426 pollutants 168, 187 – 191
Persia 291, 309 polygenic risk scores (PRS) 124, 134 – 135, 142, 149
person-centered approaches 437 – 438 polymorphisms 134
Pettit, G. 486 polyvinyl chloride 188
Pew Research Center 481, 499 Pomerantz, E. 409, 459
PFAS (perfluoroalkyl substances) 189 – 190 Pomeroy, S. 302
534
Index
Pongo pygmaeus 91 protein 34, 186

Porter, R. 39 proto-oncogenes 47
positive parenting 135 – 142, 145 – 147, 150 – 151 Proverbs 299
positive/protective environments 126 proximal behaviors 457
positive reinforcement 138, 141 PRS (polygenic risk scores) 124, 134 – 135, 142, 149
post-modernist perspectives 288 Psalm 131 299
postnatal depression (PND) 196, 271 – 272 psychoactive drugs 32
postnatal parental investment 8 psychobiology 30 – 77
postpartum depression 236 – 237 psychological control 458, 463
postpartum period 35 – 36, 41 – 45, 56 psychological processes 35
poverty 321, 337, 372, 375 – 391, 433, 436, 475, 493 psychological theories 448 – 449
Powell, T. 186 psychomotor development 189
power-assertive strategies 450, 465 psychopathology 141, 153, 449
precision vs. accuracy 356 – 357 psychosocial acceleration theory 19
predictive adaptive responses (PARs) 170, 198 – 199 psychosocial stress 178 – 181, 195 – 198
predictive relations 431 – 432 PTSD 236 – 238
pregnancy: ancient history and 293, 297, 304 – 305, Puah 297
313; evolutionary ecology and 166; gray matter puberty 10, 19 – 20, 95, 104, 187
and 270; infant skull size and 13; maternal stress Puerto Rican families 439, 460, 464
and 168 – 171, 179, 184 – 186, 192; modern history punishment 99, 310 – 313, 350, 385 – 388, 409 – 410,
and 345; Pleistocene and 10; social support and 427, 454 – 455
178; unintended 345 Punt, Land of 291 – 292
prehistory 288 – 291 pup-avoidance 36, 40
prematurity 21, 238, 261 pup-induction/pup-sensitization 34 – 36
premodern/modern dichotomy 322 – 323 pup reinforcement 44 – 45
prenatal bonding 191 – 192, 225 pup-seeking behavior 50
prenatal brain growth 13 Purcell, S. 127
prenatal influences 130 – 131 purring 106
prenatal parenting 166 – 219 Putnam, S. 487
prenatal stress 173 – 184 PVN (paraventricular nucleus) 36, 46
preparatory motor responses 267 – 268 Pythagoras 169
preputial glands 38 – 40
prereproductive period 5, 14 quadrupedal steps 87
preschool children 18, 137 – 141, 306 – 308 Quakers 325
Preston, S. 401 qualitative research 376 – 378, 383
preterm birth 181, 184, 194 quantitative research 372
preweaning 56 quantitative trait loci 56
Primate Research Institute, Kyoto University (PRI) Quebec Newborn Twin Study (QNTS) 135,
83 – 87 138, 148
primates 9 – 13, 31, 78 – 122, 80, 81, 186 Quintilian 312 – 313, 314
primatology 291
primiparous mothers 40 – 42, 105 Rachel 297
primogeniture 311 Ragan, C. 56
proactive/supportive parenting 387 – 388, 494 randomized controlled trials (RCTs) 365 – 366
problem solving abilities 400 Rao, N. 483
proestrus phase 44 Rasmi, S. 458
progesterone 34 – 37, 41, 49 – 50 rats: attachment and 20; diets of 186 – 187; dopamine
projective-expressive systems 449 (DA) and 50 – 53; folate and 185; hormones and
Project on Human Development in Chicago 228; maternal behavior in 31 – 39, 42 – 44, 47,
Neighborhoods (PHDCN) 376 – 379, 384 – 385 50 – 59; olfaction and 39, 264; phthalates and 188;
prolactin (PRL) 34, 39 – 41, 47, 220 – 222, 228, 231 touch cues and 39 – 40
property rights 290, 315 Rattus norvegicus 31
proportionate birth weight 175 Raudenbush, S. 373
Propper, C. 141 Raviv, T. 406
prosimians 80, 81, 105 – 107, 110 – 111 Razza, R. 488
prosocial behavior 137 – 139, 168 reaching 87
prostitution 338 reaction norms 167
protectiveness 151 reactive parenting 387 – 388, 494
535
Index
reactivity 99, 110, 183 role construction 411

reading 137, 429, 434, 481 Romans 310 – 317
recall bias 359 – 360 Romantic concepts 340
reciprocity 234 Rome 288 – 292
recognition systems 251 – 256 Romulus 310 – 311
refugees 272, 321, 337 – 338, 474 – 475 Roopnarine, J. 482 – 483
regional case studies 335 – 344 Rose, G.: Sick Individuals and Sick Populations 349
registry data 127, 153 – 154 Rosenwein, B. 323
rejection 99, 103, 109 – 110 Rossen, L. 191
relatedness, modeling of 123 routines 434
relational developmental systems theories 374 RSA (respiratory sinus arrhythmia) 141
religion 296, 299, 320, 492 – 493 ruffed lemurs 105
remarriage 311, 316 rule-based strategies 387
removals, of children 336 runaway selection 13
REM sleep 83 rural environments 390, 451 – 452, 464, 479 – 480, 494
repetition priming paradigms 271 Rusak, B. 47
replicability 135 Russia 324 – 328, 343
reporting bias 358 – 359 Ruth 298
reproductive fitness 4 – 7, 169, 172 Rutter, M. 126
reproductive strategies 19 – 20 Rwanda 338
Republic of Ireland 192
resilience 109, 382, 387 Sachs, J. 495
resource allocation 184, 192 – 193 safety 479 – 480, 489, 494
resource availability 19, 23, 172, 178, 391 St.Vincent 482
resource flow 175 – 176 Salafia, C. 175
respect/respeto 456, 461, 464 salience detection 263 – 269
respiratory sinus arrhythmia (RSA) 141 salivary alpha amylase (sAA) 231 – 232, 238
responsibility training 422 Salvatore, J. 149
responsive care intervention 107 – 108 SAM (sympathetic-adrenal-medullary) 231 – 232
responsiveness: culture and 457 – 458, 462; maternal sampling, of neighborhoods 390
30, 34 – 47, 104, 110, 141, 485; of newborn Sampson, R. 373
chimpanzees 83 – 84 Samuel 298
responsivity 262, 266 – 267, 428 Sandler, H. 411
restrictive childrearing 98 – 99, 383 Sandman, C. 183
retardation 13 Santos, I. 351
retrieval behavior 34, 37 – 41, 49 – 53 Sarah 297
reverse causality 361 – 362 Saul 297
reward-related cues and processes 55, 252 – 257, 266, Saw, A. 457
269 – 274 scaffolding 89, 92, 434, 452, 481, 489
rGE (gene-environment correlations) 124 – 156, Scandinavia 154
129, 365 Scarr, S. 23
rhesus macaques 99, 109 scent marking 105
rhesus monkeys 80, 97 – 100, 108 – 110 Scheper-Hughes, N. 475, 490
Rhoades, B. 437 – 438 Scheumann, M. 106
Richman, A. 484 schizophrenia 125
Riddle, J. 303 Schoff, K. 485
Rijt-Plooij, H. 87 – 88 school-based parent involvement 408 – 411
ring- tailed lemurs 105 – 106 Scipio Africanus 313
risk 232, 236 – 238, 351, 354 – 356, 424 Scott, J. 465
risk and resilience perspectives 382, 387 scouting 328
Risley, T. 406, 428, 480, 485 scrounging 92
RNA molecules 187 seasonal affective disorder (SAD) 495
Robinson, D. 50 – 51 Seckl, J. 181
rodents 31 – 33, 220, 223 – 228, 234 – 236, 264. see also secure attachment 19, 104; see attachment
rats sedentary lifestyles 9
Rodriguez, A. 183 segregating genes 129, 132
Rogerson, F. 181 SEI (Socioeconomic Index of Occupations) 423 – 434
Rohner, R. 462 selection bias 357 – 358
536
Index
selection pressures 4 – 6, 10 – 12 signaling 88 – 89

selective bonding 39 silence 459
selective investment theory 502 silverbacks 84, 92
selective placement 131 – 132 simians 81
selectivity, maternal 40, 43 Simons, R. 383 – 385
self-construals 451 Simpson’s paradox 362
self-control 138 – 140, 143, 453 – 456 Singapore 483
self-efficacy 135, 411 – 413, 433 – 434 single indicator approach 402
self-esteem 341 – 342 single nucleotide polymorphisms (SNPs) 134,
self-expression 454 225 – 226
self-handicapping behaviors 88 single parenthood 338 – 339, 382
self-interests 6 Sinuhe 292
self-ratings 179 Six Culture Study of Socialization 449, 454
self-regulation 151, 183, 400, 463 skill-based mechanisms 410 – 411
self-worth 147 skills formation 502
SEM (structural equation modeling) 125, 358 skin-to-skin contact 238
sensitivity 135 – 137, 141 – 142, 403, 407, 427 Skrundz, M. 236
sensory changes 36 – 40 slaves/slavery 293, 300, 336 – 337
sensory mechanisms 43 – 44 SLC6A4 gene 134
sensory processing 267 sleep 83, 106, 341, 476
separation anxiety 88 slow thinking 256
separation of offspring 108 – 110, 338 small apes 80, 82 – 96, 93 – 94
serotonin 34, 52 – 53, 185 smiling 86 – 88, 254, 263
serotonin-transporter-linked polymorphic region smoking 187, 194, 232, 364
128, 134 – 137, 140 SNPs (single nucleotide polymorphisms) 134,
servants 292, 301 225 – 226
SES. see socioeconomic status (SES) sociable-assertive behavior 451 – 452
settler societies 336 social-behavioral functioning 453
sex 8, 11 – 12, 51 – 52, 422 social benefits models 434
sex allocation theory 184 social brain 256 – 257, 274 – 275
sex-cell size 7 social capital 423, 433, 436 – 437, 494
sex differences 6 – 8, 183 – 184 social causation models 430
sex hormones 223 social change 464 – 465
sex-related hormones 223 social-cognitive abilities 453
Sexton, H. 407 social cohesion 11
sexual dimorphism 183 social complexity 15
sexual selection 7 – 8 social construction 288 – 291, 303 – 308, 311 – 313
sexual swelling 95 – 96 social context model 126
shaming 309, 321 – 322, 342, 459 social control 376 – 377
Shanahan, L. 486 social cues 18 – 20
Shanghai 483 social drift hypothesis 435, 438
shantytowns 475, 490 social groups 9 – 10, 84 – 85
shared culture 450 – 451 social history 287, 321
Sheard, N.: Maternal Behavior in the Rat 33 social initiative 453 – 455
Sheehan, T. 47 social integration 492 – 494
sheep 38 – 40 social intelligence 10 – 12, 15
Sheridan, S. 408 sociality 10 – 12
Shipreh 297 socialization 377 – 378, 422, 449 – 459, 463 – 465
Shrestha, S. 449 – 450 social motivation 139
shyness 150 social networks 479, 492
shyness-inhibition 454 – 455 social neuroendocrinology 220 – 249
siamangs 85, 93 – 95 social processes 274, 373, 376, 379 – 380, 388, 391
sibling adoption studies 127 – 132 social relationships 432 – 433
sibling comparisons 364 – 365 social responsiveness 97
siblings 132, 174, 298, 486 social risk models 434
Sick Individuals and Sick Populations (Rose) 349 social stratification 422
Sigel, I. 403 social support 15 – 17, 178, 191, 376 – 379, 389, 486
Sigman, M. 478 social transmission model 137
537
Index
socio-assays 174 nurturant behaviors and 30; perceived, objective,

socio-cognitive functioning 452 and biology of 195 – 196; prenatal 173 – 184; of
sociodramatic behavior 454 refugees 338; United States and 343 – 344
socioecological perspective 452 – 454 stress hormones 167 – 168, 186, 223, 223
Socioeconomic Index of Occupations (SEI) 423 – 434 Stroop Test 149, 186
socioeconomic status (SES) 421 – 447; culture structural equation modeling (SEM) 125, 358
and 456; education and 330; environment and structured parenting 138
479 – 487, 490, 493; genetics and 137; illegitimacy styles, of parenting 426
and 339; modern history and 320, 324, 335 – 337; Styron, W.: Sophie’s Choice 475
neighborhoods and 372 – 391; neuroendocrinology subcortical areas 257
and 230 – 231; parental educational attainment and subcortical mammalian network 235
401 – 402, 405 – 413 subjective pleasure 267
socioemotional development 137, 251 – 252, 462 – 464 submissive behaviors 95
somatosensory functions 39 – 40, 43 – 45 sub-Saharan Africa 328
Sophie’s Choice (Styron) 475 subsidized housing 377
South America 452 substance use 148
South Korea 330, 448, 455, 458 – 460, 463 – 465 suckling 33 – 35, 38 – 41, 83, 89, 92, 96 – 97
Soviet Union 328, 452, 464 Suh, E. 47
spadefoot toads 167 Sullivan, W. 496
Spain 458, 484 – 485 Sumatra 94
Sparta 300, 305, 308 – 309, 317 Sumer 291, 295
species-typical patterns 5 – 7, 18, 24, 79 – 80 Sundquist, J. 153
speech 251 – 254, 406, 428 – 429, 432 Sundquist, K. 153
sperm, production of 7 Suomi, S. 99
spillover effect 487 Super, C. 449 – 452, 476
spirituality, feminist 291 superparenting 23, 383
Spock, B. 327; The Common Sense Book of Baby and supportive parenting 136, 386 – 388, 450, 483 – 488
Child Care 334 surrogate parenthood 290
spousal support 486 – 487 surveillance 490 – 492
spurious correlates 362, 430, 435 – 436 Survey of Income and Program Population 490
squirrel monkeys 100 – 105, 109 sustenance 477 – 479
starlings 184 swaddling 305, 308 – 309, 341, 495
Starr, R. 486 Sweden 148, 151 – 153, 194, 198, 364
state authority 324, 333 – 334 sympathetic-adrenal-medullary (SAM) 231 – 232
state regulation 83 – 84 sympathetic nervous system 180, 230
statistical approaches 125 – 127, 134 – 135, 190, synaptic brain growth 13
373, 424 synaptic plasticity 109
Stattin, H. 387 synchronized capabilities 196
Steinberg, L. 493 synchrony 222, 225 – 226, 229 – 235
stepfamilies 21 – 22, 132, 146, 313 Syria 291, 338
Stern, J. 39 Syrian refugees 272, 474
steroid hormones 34, 180, 229
Stewart, M. 126 – 127 Tacitus 287, 313
Stewart, P. 181 tactile cues 273
still-face phase 227, 230 tagging 265 – 268
stimulation 350 – 351, 480 – 483 Tai Forest 110
stimulatory parenting 225 tail tuffs 90 – 92
Stoicism 313 Tai National Park 87
Stone, L. 322 – 323 Taiwan 342, 455, 459 – 461
Strange Situation 136 Tale of the Shipwrecked Sailor 292
Strepsiades 306 – 307 Talmudic era 297 – 300
stress: assisted reproductive technologies (ART) and Tamang, B. 449 – 450, 459
195; biomarkers of 237; childhood psychosocial Tamang 459
197 – 198; co-adaptation and 196; defined 178; tamarins 101 – 105, 110
depression and 128; environment and 480, 485, Tang, S. 408 – 410
496 – 497; hormones and 229 – 233, 236 – 237; Tanjung Puting 84
long-term consequences of 108 – 109; maternal Tanzania 478
168, 177 – 184; morning sickness and 193 – 194; TAP (Texas Adoption Project) 125
538
Index
TAPS (Twins, Adoptees, Peers, and Siblings) transgenes 57

study 142 transgenic mice mutants 33
Tardif, T. 404 transience 433
task orientation 137, 448 trauma 233 – 238, 487 – 488
Tauret 293 traumatic head injuries 480
Taylor, A. 496 trial-and-error learning 90, 452
TBED-C (Twin Study of Behavioral and Emotional triplets 101
Development in Children) 143 Trivers, R. 3 – 8
TCHAD (Twin Study of Child and Adolescent Trommsdorff, G. 449 – 450
Development) 148 – 149 Tronick, E. 495
TD (typically developing children) 233 – 234 Troughton, E. 126 – 127
teats 39 – 40 Trpc2 38
technology 451, 487 – 491 tsak vocalizations 106
TEDS (Twin’s Early Development Study) 135 – 139, Tuanan 84
142 – 144 Tullia 315
television 434, 487 – 488, 491 Turkey 451, 458, 465, 474
temperament 110, 127 – 128 Turkheimer, E. 153
temporal lobes 257 twin-difference method 138
teratogens 187 – 191 Twin/Offspring Study in Sweden (TOSS) 145 – 151
territorial behavior 85, 227 twins 101, 130
testes, descent of 99 Twins, Adoptees, Peers, and Siblings (TAPS)
testosterone (T) 220 – 222, 228 – 229 study 142
Texas Adoption Project (TAP) 125 twins-as-children designs 130
Texas Twin Project 146 twins-as-parents designs 130
Thailand 458 Twin’s Early Development Study (TEDS) 135 – 139,
thermal cues 40 – 41 142 – 144
Thinking Fast and Slow (Kahneman) 256 twin studies 21, 124 – 155, 129
Thomas, R. 495 Twin Study of Behavioral and Emotional
Thompson, W.: Behavior Genetics 124 – 125 Development in Children (TBED-C) 143
Three Character Classic 462 Twin Study of Child and Adolescent Development
three-generation households 486 (TCHAD) 148 – 149
Thucydides 287 type 2 diabetes 187
tickling 88 typically developing children (TD) 233 – 234
Tigris-Euphrates River Valley 295 Tyson, D. 409
timidity 36
timing 168, 271, 354 – 356 Ueno, A. 89
Timoxena 312 ultrasonic calls 40
Tinbergen, N. 253 uncles 16, 132
titi monkeys 101 – 105 uncorrelated risk 356
tobacco 187, 194 undernutrition 170, 184 – 186, 478
toddlerhood 141, 304 – 309. see also early childhood; ungulates 38 – 40
infancy UNICEF 351
toilet training 306, 422 uninvolved parenting 144
Tokyo Twin Cohort Project 138, 144 United Arab Emirates 342
Tokyo Twin Cross-Sectional Survey 144 United Kingdom 135, 146, 190 – 192, 261, 273,
tomb paintings 292 – 294 350 – 351
tool manufacture 90 United Nations 475
topical regions 495 United States: culture in 449, 454 – 460, 464;
Tortilla Flats (Boyle) 475 environment and 474 – 476, 479, 483 – 484, 499;
TOSS (Twin/Offspring Study in Sweden) 145 – 151 evolutionary perspectives and 22 – 23; modern
touch 225 – 226, 233, 263 – 264, 273 history and 321 – 346; neighborhoods and
touch cues 39 – 40 372 – 373, 375 – 386, 391; parental educational
toxins, prenatal 187 – 191 attainment in 404, 408 – 409; socioeconomic status
toys and dolls 293 – 294, 308, 434, 481, 499 (SES) in 421 – 426, 439
TRACKS twin study 136 – 139 urban environments 390, 451 – 452, 480, 496
traditional cultures 7, 14, 17, 20 – 23 urbanization 327, 451 – 452, 464 – 465
transcription regulation 170 U.S. Bureau of Labor Statistics 499
transduction 38 uterine environment 174 – 177
539
Index
valence effect 263 – 267, 273 Wandel, M. 478

Valentin, S. 476 Wang, L. 462
values 425, 450 Wang, Q. 449 – 450, 459
Van Batenburg-Eddes, T. 364 warfare 10, 298 – 299, 311, 321, 337 – 338,
Vandermaas-Peeler, M. 483 474, 487
variability 110 wariness 137 – 138
variable number tandem repeats (VNTRs) 134 warmth 126 – 128, 137 – 139, 142 – 146, 150 – 151;
variants 134 – 137 culture and 455 – 458, 461 – 465; environment
vasopressin (AVP) 33 – 36, 46, 220 – 224, 227 – 228 and 487; neighborhood and 378, 386;
VBM (voxel-based morphometry) 270 parental educational attainment and 403, 407;
ventral bed nucleus of the stria terminalis (vBNST) socioeconomic status (SES) and 427, 430
46 – 53, 58 War on Poverty 421 – 423
ventral pallidum (VP) 52 war trauma 233 – 238
ventral riding 87, 105 Wasserman, G. 485
ventral stimulation 44 Way Canguk Research Area 94
ventral striatum 49, 52 – 53 weaning 17, 41, 79, 85, 90 – 92, 99 – 101, 104 – 106,
ventral tegmental area (VTA) 47 – 53, 235 294, 298, 305, 422
ventral trunk 39 – 40 Weaver, A. 104
ventral-ventral contact 82 Weisfeld, G. 264
ventromedial nucleus of the hypothalamus (VMH) welfare state 333
46 – 49 well-being 375 – 381, 385, 390, 402, 432, 497
ventromedial prefrontal cortex (vmPFC) 235 – 236 Wellman, H. 404
Venus fertility figurines 290 – 291 Wells, N. 500
verbal markers 459 Western cultures 8, 321 – 324, 448, 457 – 466
Vernet-Maury, E. 40 Western history 289 – 291, 324 – 327, 338 – 339
Vernon-Feagans, L. 494 wet nurses 294, 298, 305, 309
versusadaptation 181 – 182 Whang, S. 455 – 456
vervets 17, 97 – 100 whistle vocalizations 106
victimization 146 White House conference on child health and
vigilance 79, 220, 225 – 227, 232, 235 protection (1936) 421
violence, inter-partner 479 White samples 154
virgin animals 33 – 40, 43 – 45 Whiteside-Mansell, L. 485
virtual twins 142 Whiting, B. 449
visual acuity 86 Whiting, J. 449, 495
visual cues 40, 253 Wiesner, B.: Maternal Behavior in the Rat 33
VMH (ventromedial nucleus of the hypothalamus) Wiley, A. 496
46 – 49 Williams, T. 184
vmPFC (ventromedial prefrontal cortex) 235 – 236 Willoughby, M. 494
VNTRs (variable number tandem repeats) 134 Wilson, M. 22
vocalizations and vocal cues 18, 87 – 89, 93 – 96, wire mothers 30
101, 104 – 106, 251 – 252, 459; early detection of within-pair estimates 364
264 – 265; maternal behavior and 37; parental brain women. see females; mothers
and 260 – 261 women’s magazines 325
voltammetry 50 – 51 Wood, L. 464
vomeronasal system 40, 46 Woodworth, G. 126 – 127
von Ranke, L. 287 Wörmann,V. 458
Vortruba-Drzal, E. 482 worms 294
voxel-based morphometry (VBM) 270 Wu, H. 462
VP (ventral pallidum) 52 Wu, P. 459
VTA (ventral tegmental area) 47 – 53, 235 Wu, R. 53
Vygotsky, L. 452 – 453 Wynne-Edwards, K. 184
Waddington, C. 169 – 171 Yamamoto,Y. 457 – 458

Waffarn, F. 183 Yaman, A. 458
Waldron, M. 153 Yates, W. 126 – 127
Wales 138 Yerkes Research Center of Emory University
Wallenius, M. 501 83 – 87, 108
Wamba 82 Yim, I. 196
540
Index
YMCA 328 Zhou, N. 457 – 458

YouGov Omnibus Parents Survey 499 – 500 Zinacantec Maya 452
Young, K. 266 – 268 zinc sulfate 38
Youngstrom, E. 485 Zolkoski, S. 488
youth culture 330 zygosity 130
Yovsi, R. 465 zygotes 169
541

Handbook of Parenting - Volume 2 - Biology and Ecology of Parenting

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Handbook of Parenting - Volume 2 - Biology and Ecology of Parenting

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Edited by Marc H. Bornstein

Preface to the Third Edition ix

1 The Evolution of Parenting and Evolutionary Approaches to Childrearing 3

2 Psychobiology of Maternal Behavior in Nonhuman Mammals 30

3 Parenting in Nonhuman Primates 78

4 Genetics and Parenting 123

5 Prenatal Parenting 166

6 The Social Neuroendocrinology of Human Parenting 220

7 Neurobiology of Human Parenting 250

8 Ancient History of Parenting 287

9 Modern History of Parenting 320

10 Epidemiology of Parenting 349

11 Neighborhoods and Parenting 371

12 Parent Education Attainment and Parenting 400

13 Socioeconomic Status and Parenting 421

14 Culture and Parenting 448

15 Environment and Parenting 474

BIOLOGY AND ECOLOGY OF PARENTING is Volume 2 of the Handbook. For parent-

Kim A. Bard is a Professor of Comparative Developmental Psychology in the Department of

David F. Bjorklund is Professor of Psychology at Florida Atlantic University. He received a BA

Miguel Cordero is a child clinical psychologist with a MA in Behavioural Neurosciences at the

Alison S. Fleming is a Professor Emerita of Psychology at the University of Toronto. Fleming

Ming Li is Professor of Psychology at the University of Nebraska-Lincoln. Li was educated at the

Patrick O. McGowan is Associate Professor of Biological Sciences at the University of Toronto

Alyson J. Myers is a PhD candidate in Developmental Psychology at Florida Atlantic University.

Jenae M. Neiderhiser is a Distinguished Professor of Psychology and Human Development

Christine E. Parsons is an Associate Professor at the Interacting Minds Centre, Department of

Priya Rajyaguru is an Academic Clinical Fellow in Psychiatry working clinically in psychiatric

Nicholas E. Waters is a PhD candidate in Developmental Psychology at the University of Michi-

Principles of Evolution and Evolutionary Developmental Science

Evolution by Natural Selection

Principles of Evolutionary Developmental Science

Parental Investment Theory

The Environment of Evolutionary Adaptedness

What Were the Selection Pressures that Led to the Modern

The Significance of Social Intelligence

The Consequences of Delaying Development

The Importance of Paternal Investment

Evolutionary Perspectives on Attachment and Parenting

Evolutionary Adaptations that Promote Attachment

a principal evolutionary function of early experience—the first 5 to 7 years—is to induce

Evolved Mechanisms Underlying Neglect, Abuse, and Infanticide

Rather, abandonment is at one extreme of a continuum that ranges between termination of

Is “Parenting” the Right Concept to Describe Human Childrearing?

Darwin, C. (1859). The origin of species. New York: Modern Library.

Methodological Issues in the Study of Rodent Maternal Behavior

Description of Maternal Behavior at Parturition

Hormonal Effects on the Onset of Maternal Behaviors

Emotional and Anxiety Changes in the New Mother

Sensory Changes in the New Mother

Olfactory Cues Have Inhibitory Effects

Auditory and Visual Cues

Experiential Effects on the Maintenance and Retention

Behavioral Changes From Birth to Weaning

Parity and Effects of Postpartum Experiences

Sensory Mechanisms Involved in Maternal Experience

Long-Term Effects of Maternal Experience and Parity on Maternal Affect

Neuroanatomy of Maternal Behavior

Maternal “Core”: Two Antagonistic Neural Systems

Neurochemistry of Maternal Behavior

The Mesolimbic and Mesocortical Dopamine Systems

In Vivo Dopamine and Enhanced Maternal Motivation

Preface to the Third Edition ix

1 The Evolution of Parenting and Evolutionary Approaches to Childrearing 3

2 Psychobiology of Maternal Behavior in Nonhuman Mammals 30

3 Parenting in Nonhuman Primates 78

4 Genetics and Parenting 123

5 Prenatal Parenting 166

6 The Social Neuroendocrinology of Human Parenting 220

7 Neurobiology of Human Parenting 250

8 Ancient History of Parenting 287

9 Modern History of Parenting 320

10 Epidemiology of Parenting 349

11 Neighborhoods and Parenting 371

12 Parent Education Attainment and Parenting 400

13 Socioeconomic Status and Parenting 421

14 Culture and Parenting 448

15 Environment and Parenting 474

Figure 3.1 Classification of living primates frequently mentioned in this chapter.

Figure 4.1 Historical timeline of notable behavioral genetics and developmental events.

Figure 4.2 Child and parent-based twin designs and gene-environment correlation.

Figure 4.3 Traditional adoption design.

Figure 4.4 Extended Children-of-Twins design.