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Neuromethods 122

Lesley J. Rogers
Giorgio Vallortigara Editors

Lateralized
Brain Functions
Methods in Human
and Non-Human Species
NEUROMETHODS

Series Editor
Wolfgang Walz
University of Saskatchewan
Saskatoon, SK, Canada

For further volumes:

http://www.springer.com/series/7657
Lateralized Brain Functions

Methods in Human and Non-Human Species

Edited by

Lesley J. Rogers
School of Science and Technology, University of New England, Armidale, Australia

Giorgio Vallortigara
Centre for Mind/Brain Sciences, University of Trento, Trento, Trento, Italy
Editors
Lesley J. Rogers Giorgio Vallortigara
School of Science and Technology Centre for Mind/Brain Sciences
University of New England University of Trento
Armidale, Australia Trento, Trento, Italy

ISSN 0893-2336 ISSN 1940-6045 (electronic)

Neuromethods
ISBN 978-1-4939-6723-0 ISBN 978-1-4939-6725-4 (eBook)
DOI 10.1007/978-1-4939-6725-4

Library of Congress Control Number: 2016956253

© Springer Science+Business Media LLC 2017

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Preface

Brief History of the Field and the Role of Animal Models

Beginning with the first discoveries of lateralized brain function and behavior in two avian
species and rats, made in the 1970s and early 1980s (1–3), investigation of lateralization in
nonhuman species has become a burgeoning field of research, expanding to include a wide
number of vertebrate species (4–9) and, more recently, even to include invertebrate species
(10, 11).
The availability of animal models for studying lateralization has produced two impor-
tant changes. Firstly, it has promoted a resurgence of interest in a research area that previ-
ously had been confined to human neuropsychology and neurology and had shown signs
of declining interest because it appeared to be incapable of tackling the most basic issues,
namely the causes of lateralization of the brain and its biological function (12). The enter-
ing of animal biology into the study of lateralization has meant not only the availability of
new system models with better accessibility to neural structures and functions but also dif-
ferent ways of considering the development of lateralization and its origins and biological
significance in evolutionary terms (13). In regard to the latter, for example, theoretical
tools from game theory and population genetics have prompted unexpected advances in
our understanding of the evolution of brain asymmetry (14–18).
A second important change, which ultimately provided the impetus for this book, relied
on the availability of animal models that allowed scientists to develop and use new tech-
niques for studying brain and behavioral lateralization. In addition, it is recognized that
these animal models are potentially able to enhance understanding of individual differences
in behavior, including aberrant or unusual behavioral and neural conditions in humans, as
well as in nonhuman species (19–21). Many different experimental approaches have been
applied, each adapted to reveal lateralization in different species or to do so more precisely
in species already known to be lateralized. This has led to a broad range of techniques with
varying success in revealing lateralization of both brain function and brain structure.
Research in the neuroscience of lateralization in humans has, of course, also progressed
in recent years, especially since neuroimaging and neurostimulation techniques have
become a source of new data on lateralization. Interestingly, however, the development of
techniques to be used with animal model systems has also had an influence on research of
lateralization in humans. An example is provided by the ethological methods developed to
investigate lateralization in everyday behavior in nonhuman species, methods that are now
used widely in humans as well (22, 23).
Now that the field has been established it seems timely to publish in one volume the
various methods used to measure lateralization in the different species. Although functional
lateralization has been investigated in humans for many decades and in nonhuman species
increasingly so over the last two or three decades, the methods used have not previously
been gathered together in one volume. Therefore, we invited leaders in the field to write
chapters on the methods they have used to investigate functional lateralization in a range of

v
vi Preface

different species, including humans. We hope that by collecting these contributions together
in one volume we are able to assist newcomers to the field. Also by presenting the methods
adopted to investigate lateralization in a broad range of species, we hope to stimulate new
research and provide a basis from which hypotheses can be tested and compared across
species.
New methods developed to test different species need to take into account species dif-
ferences in sensory and motor systems. We expect this line of study to continue as research-
ers explore the evolution of brain and behavioral lateralization (24). In fact, study of a wide
range of vertebrate species has led to the realization that a basic pattern of asymmetry of
function is common across vertebrate species (25). The right hemisphere is specialized to
attend to novel stimuli, including predators, to control social behavior, recognize faces, and
process global information using spatial cues (26). The left hemisphere is specialized for
focused attention needed to perform learned tasks, to follow rules, and to categorize stim-
uli (9). This division of function is present in humans and other vertebrate species.
Important to understanding lateralization is knowledge of how it develops. For this
direction of research a few key species have been the focus of detailed study, including
domestic chicks (e.g., 27–29), pigeons (30), and zebrafish (31–34). These species were
chosen as models for studying the development of lateralization because their stages of
development are known rather precisely and because sensory inputs can be manipulated
with the aim of determining the role of sensory experience in the development of lateraliza-
tion. In all three of these species, there is clear evidence that exposing the developing
embryo to light is essential for the development of visual lateralization. For example, light
exposure of avian embryos during the final stages of development before hatching stimu-
lates the right eye but not the left because the embryo’s head is twisted to the side so that
the left eye is occluded by the embryo’s body (8). This difference in stimulation of the left
and right eyes at a time when connections between eye and brain first become functional is
essential in establishing certain visual lateralities, a fact known because these functions are
not lateralized if the embryos are incubated in the dark during this critical stage of develop-
ment (see Chap. 19).
To discover the causation of each type of functional lateralization is another approach
taken by researchers of brain lateralization. This has led to detailed examination of the
structural differences between the left and right hemispheres, examined in both humans
and other species, as well as hemispheric differences in connections between neurons and
in neurotransmitter concentrations.
Of course, detailed knowledge of the function of lateralization is essential. What are the
advantages of having a lateralized brain and, on the contrary, what disadvantages might be
apparent? In this direction of investigation, it is important to consider two distinct types of
lateralization: individual and population lateralization (24). Individual lateralization is that
present in individual members of a species but favoring the left in some individuals and the
right side in others, resulting in no directional bias within the group or population.
Population lateralization, also referred to as directional lateralization, is present when the
majority of individuals are lateralized in the same direction. Handedness in humans is an
example of the latter, as also is the lateralization of visual processing in birds that we have
discussed above.
In this volume we have included some chapters with methods that are simple to apply
and others that require more sophisticated techniques, often newly available (e.g., Chap. 11
on optogenetics). Each set of methods can lead to discovery of different levels or types of
 Preface vii

lateralization and, contrary to one approach superseding another, all methods can be com-
plementary in advancing understanding.
Chapters 1–6 address measuring lateralization by scoring behavior induced by inputs to
one or the other side of the brain and in a range of species. These chapters include some of
the classic methods developed by experimental psychologists to deal with hemispheric spe-
cialization, such as tachistoscopic viewing and dichotic listening (Chap. 1 by Ocklenburg),
which have been improved to the highest level of technical precision and sophistication,
and the study of split-brain patients of the clinical neuropsychological tradition (Chap. 2 by
Fabri, Foschi, Pierpaoli, and Polonara). The roots of all these methods can be traced back
to the early sensory physiology and psychology of the nineteenth century. In fact, it was
Gustav Fechner, the founder of psychophysics, who made the first inquiry about the pos-
sible outcome of disconnecting the two cerebral hemispheres: “The two cerebral hemi-
spheres, while beginning with the same moods, predispositions, knowledge, and memories,
indeed the same consciousness generally, will [when divided through the middle] thereafter
develop differently according to the external relations into which each will enter” (Gustav
Theodor Fechner, 1860, in Zangwill 1974 (35)).
New methods have come from ethology and include measurements of eye preferences
(particularly in animals with laterally placed eyes and complete decussation at the optic
chiasma) and ear preferences (Chap. 3 by Rogers), as well as preferences in nostril use and
olfactory stimulation (Chap. 4 by Siniscalchi). Behavioral methods have been developed to
study lateralization in invertebrates (Chap. 6 by Frasnelli); thanks to the limited number of
neurons in these model species, the combination of behavioral analyses together with the
sophisticated molecular and genetic techniques available for some invertebrate species (e.g.,
the fruit fly) research on invertebrates promises the possibility of important breakthroughs
in the study of brain asymmetries in the years to come. In fact, the study of hand and limb
preferences, which has been traditionally the province of an allegedly unique phenomenon,
i.e., human handedness, has been deeply challenged by the mounting evidence from studies
on nonhuman primates and other species, both mammals and birds, showing a variety of
asymmetries in limb usage (36). Here the availability of precise techniques of recording the
use of the limbs in natural and seminatural conditions may prove to be crucial in comparing
strength and direction of handedness in different species (Chap. 5 by Forrester).
Chapters 7–11 cover neurobiological methods used to reveal lateralization. Again,
these include both well-established techniques such as lesion studies (Chap. 7 by Manns),
electrophysiology and pharmacology (Chap. 8 by McCabe), tract tracing (Chap. 9 by
Stöckens and Güntürkün), and early gene expression (Chap. 10 by Patton, Uysal, Kellog,
and Shimizu), as well as the new optogenetic methods that allow selective activation or
blocking of specific circuits or synapses in the left or right side of the brain (Chap. 11 by
El-Gaby, Kohl, and Paulsen).
Chapters 12–15 address imaging techniques, electroencephalographic techniques, and
transcranial stimulation to reveal lateralization. The mixture of human and nonhuman ani-
mal research here is apparent, with noninvasive techniques such as transcranial magnetic
stimulation (Chap. 12 by Cattaneo) and electroencephalographic stimulation (Chap. 13 by
Mazza and Pagno) mainly used for research in humans, fMRI imaging used to compare
human and nonhuman primates species (Chap. 14 by Hopkins and Phillips), and other
more recent methods with much finer spatial and temporal resolution, such as two-photon
microscopy, confined at present to system models such as insects (Chap. 15 by Paoli,
Andrione, and Haase).
viii Preface

For a long time the genetic basis of lateralization was limited to the study of inheritance
of handedness in humans (37). New genetic techniques in studying lateralization in humans
(Chap. 16 by Paracchini and Scerri), zebrafish (Chap. 17 by Duboué and Halpern), and C.
elegans (Chap. 18 by Vidal and Hobert) are covered in Chaps. 16–18 describing powerful
techniques to address the role of genes in the establishment and development of brain
asymmetry.
Last, but by no means least, are Chaps. 19–21 covering methods used to study the
development of lateralization and to do so by manipulation of sensory exposure (Chap. 19
by Chiandetti), hormone levels (Chap. 20 by Beking, Geuze, and Groothuis), and model
systems for the study of lateralized development (Chap. 21 by Blackiston and Levin).
Overall we believe that this collection of papers can provide a state-of-the-art collection
of methods currently in use for investigating brain and behavioral asymmetries, thereby
nurturing the next generation of scientists in this field. The latter scientists will certainly
make further progress, both in methods and theory, in the years to come. Our hope is that
this book will develop the field to such an extent that it will lead to a need for a second edi-
tion covering a new collection of methods.

Armidale, Australia Lesley J. Rogers

Trento, Italy Giorgio Vallortigara

References

1. Nottebohm F (1971) Neural lateralization of brain asymmetries. Cambridge University

vocal control in a passerine bird. I. Song. J Exp
Zool 177:229–261.
2. Rogers LJ, Anson JM (1979) Lateralisation of
function in the chicken forebrain. Pharmacol
Biochem Behav 10:679–686.
3. Denenberg VH (1981) Hemispheric laterality
in animals and the effects of early experience.
Behav Brain Sci 4:1–49.
4. Bisazza A, Rogers LJ, Vallortigara G (1998)
The origins of cerebral asymmetry: a review of
evidence of behavioural and brain lateralization
in fishes, amphibians, and reptiles. Neurosci
Biobehav Rev 22:411–426.
5. Vallortigara G, Rogers LJ, Bisazza A (1999)
Possible evolutionary origins of cognitive brain
lateralization. Brain Res Rev 30:164–175.
6. Vallortigara G (2000) Comparative neuropsy-
chology of the dual brain: a stroll through left
and right animals’ perceptual worlds. Brain
Lang 73:189–219.
7. Vallortigara G, Chiandetti C, Sovrano VA
(2011) Brain asymmetry (animal). Wiley
Interdiscip Rev Cogn Sci 2:146–157.
doi:10.1002/wcs.100.
8. Rogers LJ, Andrew RJ (eds) (2002)
Comparative vertebrate lateralization.
Cambridge University Press, Cambridge.
9. Rogers LJ, Vallortigara G, Andrew RJ (2013).
Divided brains: the biology and behaviour of
Press, Cambridge.
10. Frasnelli E, Vallortigara G, Rogers LJ (2012)
Left-right asymmetries of behaviour and ner-
vous system in invertebrates. Neurosci
Biobehav Rev 36:1273–1291.
11. Frasnelli E, Haase A, Rigosi E, Anfora G,
Rogers LJ, Vallortigara G (2014). The bee as a
model to investigate brain and behavioural
asymmetries. Insects 5:120–138. doi:10.3390/
insects5010120.
12. Efron R (1990) The decline and fall of hemi-
spheric specialization. Lawrence Erlbaum
Associates, Hillsdale.
13. Rogers LJ, Vallortigara G (2015) When and
why did brains break symmetry? Symmetry
7:2181–2194.
14. Ghirlanda S, Frasnelli E, Vallortigara G (2009)
Intraspecific competition and coordination in
the evolution of lateralization. Philos Trans R
Soc Lond B Biol Sci 364:861–866.
15. Ghirlanda S, Vallortigara G (2004) The evolu-
tion of brain lateralization: a game theoretical
analysis of population structure. Proc Biol Sci
271:853–857.
16. Raymond M, Pontier D, Dufour AB, Moller A
(1996) Frequency-dependent maintenance of
left handedness in humans. Proc Biol Sci
263:1627–1633. doi:10.1098/rspb.1996.0238.
17. Vallortigara G (2006) The evolutionary psy-
chology of left and right: costs and benefits of

lateralization. Dev Psychobiol 48:418–427.
doi:10.1002/dev.20166.
18. Abrams DM, Panaggio MJ (2012) A model
balancing cooperation and competition can
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nance of left-handed athletes. J R Soc Interface
9:2718–2722. doi:10.1098/rsif.2012.0211.
19. Concha ML, Bianco IH, Wilson SW (2012)
Encoding asymmetry within neural circuits.
Nat Rev Neurosci 13:832–843.
20. Duboc V, Dufourcq P, Blader P, Roussingé M
(2015) Asymmetry of the brain: development
and implications. Ann Rev Genet
49:647–672.
21. Branson NJ, Rogers LJ (2006) Relationship
between paw preference strength and noise
phobia in Canis familiaris. J Comp Psychol
120(3):176–183.
22. Forrester GS, Pegler R, Thomas MA, Mareschal
D (2014) Handedness as a marker of cerebral
lateralization in children with and without
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23. Marzoli D, Tommasi L (2009) Side biases in
humans (Homo sapiens): three ecological stud-
ies on hemispheric asymmetries.
Naturwissenschaften 96(9):1099–1106.
24. Vallortigara G, Rogers LJ (2005) Survival with
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25. MacNeilage PF, Rogers LJ, Vallortigara G
(2009) Origins of the left and right brain. Sci
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26. Rosa Salva O, Regolin L, Mascalzoni E,
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doi:10.3819/ccbr.2012.70006.
27. Andrew RJ (1991) The nature of behavioral
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chick as a model, Oxford University Pres,
Oxford. pp. 536–554.
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28. Vallortigara G, Cozzutti C, Tommasi L, Rogers
LJ (2001) How birds use their eyes: opposite
left-right specialisation for the lateral and fron-
tal visual hemifield in the domestic chick. Curr
Biol 11:29–33.
29. Rogers LJ (1996) Behavioral, structural and
neurochemical asymmetries in the avian brain:
a model system for studying visual develop-
ment and processing. Neurosci Biobehav Rev
20:487–503.
30. Güntürkün O (2006) Avian cerebral asymme-
tries: the view from the inside. Cortex
42:104–106.
31. Barth KA, Miklosi A, Watkins J, Bianco IH,
Wilson SW, Andrew RJ (2005) fsi zebrafish
show concordant reversal of viscera, neuro-
anatomy and subset of behavioural responses.
Curr Biol 15:844–850.
32. Concha ML, Burdine RD, Russell C, Schier
AF, Wilson SW (2000) A nodal signaling path-
way regulates the laterality of neuroanatomical
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33. Gamse JT, Thisse C, Thisse B, Halpern ME
(2003) The parapineal mediates left-right
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34. Sovrano VA, Andrew RJ (2006) Eye use dur-
ing viewing a reflection: behavioural lateralisa-
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35. Zangwill OL (1974) Consciousness and the
cerebral hemispheres. In: Dimond SJ,
Beaumont JG (eds) Hemisphere function in
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36. Versace E, Vallortigara G (2015) Forelimb
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Contents

Preface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v
Contributors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xiii

PART I BEHAVIORAL METHODS

1 Tachistoscopic Viewing and Dichotic Listening . . . . . . . . . . . . . . . . . . . . . . . . 3
Sebastian Ocklenburg
2 Split-Brain Human Subjects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Mara Fabri, Nicoletta Foschi, Chiara Pierpaoli, and Gabriele Polonara
3 Eye and Ear Preferences . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Lesley J. Rogers
4 Olfactory Lateralization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
Marcello Siniscalchi
5 Hand, Limb, and Other Motor Preferences. . . . . . . . . . . . . . . . . . . . . . . . . . . 121
Gillian S. Forrester
6 Lateralization in Invertebrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
Elisa Frasnelli

PART II NEUROBIOLOGICAL METHODS

7 Unilateral Lesions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
Martina Manns
8 Pharmacological Agents and Electrophysiological Techniques . . . . . . . . . . . . . 251
Brian McCabe
9 Tract Tracing and Histological Techniques . . . . . . . . . . . . . . . . . . . . . . . . . . . 277
Felix Ströckens and Onur Güntürkün
10 Brain Mapping Using the Immediate Early Gene Zenk . . . . . . . . . . . . . . . . . . 313
Tadd B. Patton, Ahmet K. Uysal, S. Leilani Kellogg, and Toru Shimizu
11 Optogenetic Methods to Study Lateralized Synaptic Function. . . . . . . . . . . . . 331
Mohamady El-Gaby, Michael M. Kohl, and Ole Paulsen

PART III ELECTROENCEPHALOGRAPHIC, IMAGING

AND NEURO-STIMULATION METHODS

12 Transcranial Magnetic Stimulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 369

Luigi Cattaneo
13 Electroencephalographic Asymmetries in Human Cognition . . . . . . . . . . . . . . 407
Veronica Mazza and Silvia Pagano
14 Noninvasive Imaging Technologies in Primates . . . . . . . . . . . . . . . . . . . . . . . . 441
William D. Hopkins and Kimberley A. Phillips

xi
xii Contents

15 Imaging Techniques in Insects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 471

Marco Paoli, Mara Andrione, and Albrecht Haase

PART IV GENETIC TECHNIQUES

16 Genetics of Human Handedness and Laterality . . . . . . . . . . . . . . . . . . . . . . . . 523
Silvia Paracchini and Tom Scerri
17 Genetic and Transgenic Approaches to Study Zebrafish Brain Asymmetry
and Lateralized Behavior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 553
Erik R. Duboué and Marnie E. Halpern
18 Methods to Study Nervous System Laterality in the Caenorhabditis
elegans Model System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 591
Berta Vidal and Oliver Hobert

PART V DEVELOPMENT OF LATERALIZATION

19 Manipulation of Strength of Cerebral Lateralization
via Embryonic Light Stimulation in Birds . . . . . . . . . . . . . . . . . . . . . . . . . . . . 611
Cinzia Chiandetti
20 Investigating Effects of Steroid Hormones on Lateralization
of Brain and Behavior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 633
Tess Beking, Reint H. Geuze, and Ton G. G. Groothuis
21 Reversals of Bodies, Brains, and Behavior . . . . . . . . . . . . . . . . . . . . . . . . . . . . 667
Douglas J. Blackiston and Michael Levin

Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 695
Contributors

MARA ANDRIONE • Centre for Mind/Brain Sciences, University of Trento, Rovereto, Italy
TESS BEKING • Clinical and Developmental Neuropsychology, University of Groningen,
Groningen, The Netherlands; Behavioral Biology, Groningen Institute for Evolutionary
Life Sciences, University of Groningen, Groningen, The Netherlands
DOUGLAS J. BLACKISTON • Biology Department, Center for Regenerative
and Developmental Biology, Tufts University, Medford, MA, USA
LUIGI CATTANEO • Dipartimento di Neuroscienze, Biomedicina e Movimento, Sezione di
Fisiologia e Psicologia, University of Verona, Verona, Italy
CINZIA CHIANDETTI • Department of Life Sciences, University of Trieste, Trieste, Italy
ERIK R. DUBOUÉ • Department of Embryology, Carnegie Institution for Science,
Baltimore, MD, USA
MOHAMADY EL-GABY • MRC Brain Network Dynamics Unit, University of Oxford,
Oxford, UK; Department of Physiology, Development and Neuroscience, University
of Cambridge, Cambridge, UK
MARA FABRI • Dipartimento di Medicina sperimentale e clinica, Sezione di Neuroscienze e
Biologia cellulare, Università Politecnica delle Marche, Ancona, Italy
GILLIAN S. FORRESTER • Department of Psychological Science – Birkbeck, University
of London, London, UK
NICOLETTA FOSCHI • Dipartimento di Scienze Cliniche Specialistiche ed
Odontostomatologiche, Sezione di Scienze Radiologiche, Università Politecnica delle
Marche, Ancona, Italy
ELISA FRASNELLI • Department of Psychology, College of Life and Environmental Sciences,
University of Exeter, Exeter, UK
REINT H. GEUZE • Clinical and Developmental Neuropsychology, University of Groningen,
Groningen, The Netherlands
TON G.G. GROOTHUIS • Behavioral Biology, Groningen Institute for Evolutionary Life
Sciences, University of Groningen, Groningen, The Netherlands
ONUR GÜNTÜRKÜN • Department of Psychology, Institute of Cognitive Neuroscience and
Biopsychology, Ruhr-University Bochum, Bochum, Germany
ALBRECHT HAASE • Centre for Mind/Brain Sciences, University of Trento, Rovereto, Italy;
Department of Physics, University of Trento, Trento, Italy
MARNIE E. HALPERN • Department of Embryology, Carnegie Institution for Science,
Baltimore, MD, USA
OLIVER HOBERT • Department of Biological Sciences, Howard Hughes Medical Institute,
Columbia University, New York, NY, USA
WILLIAM D. HOPKINS • Division of Developmental and Cognitive Neuroscience, Yerkes
National Primate Research Center, Atlanta, GA, USA
LEILANI S. KELLOGG • Department of Psychology, University of South Florida, Tampa, FL, USA
MICHAEL M. KOHL • Department of Physiology, Anatomy and Genetics, University of
Oxford, Oxford, UK

xiii
xiv Contributors

MICHAEL LEVIN • Biology Department, Center for Regenerative and Developmental

Biology, Tufts University, Medford, MA, USA
MARTINA MANNS • Department of Biopsychology, Institute of Cognitive Neuroscience,
Ruhr-University Bochum, Bochum, Germany
VERONICA MAZZA • Centre for Mind/Brain Sciences (CIMeC), University of Trento,
Rovereto, Italy; IRCSS San Giovanni di Dio, Fatebenefratelli, Brescia, Italy
BRIAN MCCABE • Sub-Department of Animal Behavior, University of Cambridge,
Cambridge, UK
SEBASTIAN OCKLENBURG • Institute of Cognitive Neuroscience and Biopsychology,
Ruhr-University Bochum, Bochum, Germany
SILVIA PAGANO • Centre for Mind/Brain Sciences (CIMeC), University of Trento,
Rovereto, Italy
MARCO PAOLI • Centre for Mind/Brain Science, University of Trento, Rovereto, Italy
SILVIA PARACCHINI • School of Medicine, University of St. Andrews, North Haugh, Scotland
TADD B. PATTON • Department of Psychological Sciences, Augusta University, Augusta,
GA, USA
OLE PAULSEN • Department of Physiology, Development and Neuroscience, University of
Cambridge, Cambridge, UK
KIMBERLEY A. PHILLIPS • Department of Psychology, Trinity University, San Antonio,
TX, USA; Southwest National Primate Research Center, Texas Biomedical Research
Institute, San Antonio, TX, USA
CHIARA PIERPAOLI • Dipartimento di Medicina sperimentale e clinica, Sezione di
Neuroscienze e Biologia cellulare, Università Politecnica delle Marche, Ancona, Italy
GABRIELE POLONARA • Centro Epilessia, Clinica di Neurologia, Azienda Ospedaliera-
Universitaria Umberto I, Ancona, Italy
LESLEY J. ROGERS • School of Science and Technology, University of New England,
Armidale, NSW, Australia
TOM SCERRI • Walter and Eliza Hall, Institute of Medical Research, Parkville, VIC,
Australia
TORU SHIMIZU • Department of Psychology, University of South Florida, Tampa, FL, USA
MARCELLO SINISCALCHI • Department of Veterinary Medicine – Section of Behavioral
Sciences and Animal Bioethics, University of Bari “Aldo Moro,”, Bari, Italy
FELIX STRÖCKENS • Department of Psychology, Institute of Cognitive Neuroscience and
Biopsychology, Ruhr-University Bochum, Bochum, Germany
AHMET K. UYSAL • Department of Psychology, University of South Florida, Tampa, FL, USA
GIORGIO VALLORTIGARA • Centre for Mind/Brains Sciences, University of Trento,
Rovereto, Italy
BERTA VIDAL • Department of Biological Sciences, Howard Hughes Medical Institute,
Columbia University, New York, NY, USA
 Part I

Behavioral Methods
 Chapter 1

Tachistoscopic Viewing and Dichotic Listening

Sebastian Ocklenburg

Abstract
While advanced neuroimaging methods such as fMRI provide a reliable way to determine individual later-
alization of function, these methods are costly and not readily available to every scientist interested in
investigating functional hemispheric asymmetries in humans. Behavioral methods of testing humans pro-
vide cheaper and easily administered alternatives to fMRI scans and are still widely used in lateralization
research today. In the following chapter, two key methods will be reviewed: divided visual field paradigms
based on tachistoscopic viewing and the dichotic listening task.

Key words Dichotic listening, Auditory system, Divided visual field paradigm, Tachistoscopic view-
ing, Visual system, Laterality, Hemispheric asymmetries

1 Tachistoscopic Viewing

1.1 Introduction There are several methodological options for the researcher to
determine, in humans, whether an individual is left- or right-
dominant for a specific cognitive function. Apart from invasive pro-
cedures such as the Wada test [1], electrophysiological methods like
EEG [2] and advanced neuroimaging methods such as PET [3] or
fMRI [4] provide reliable ways of determining individual lateraliza-
tion of cognitive functions. Unfortunately, these methods have in
common that they require more or less costly equipment such as
MRI scanners or EEG systems, which are not readily available to
every scientist interested in investigating functional hemispheric
asymmetries in humans. Moreover, some of these methods are not
suitable for specific groups of patients (e.g. patients with a pace-
maker cannot be tested in a MRI scanner) or may cause discomfort
in some participants (e.g. a fMRI scan can be perceived as very
unpleasant by claustrophobic individuals due to the narrowness of
the scanner tube). By contrast, behavioral measures of hemispheric
asymmetries have no contraindications and are cheap and readily
available to any researcher. Divided visual field paradigms based on
tachistoscopic viewing comprise one of the major groups of behav-
ioral tests used in laterality research (see Table 1 for key findings).
Lesley J. Rogers and Giorgio Vallortigara (eds.), Lateralized Brain Functions: Methods in Human and Non-Human Species,
Neuromethods, vol. 122, DOI 10.1007/978-1-4939-6725-4_1, © Springer Science+Business Media LLC 2017

3
4 Sebastian Ocklenburg

Table 1
Key papers for divided visual field paradigms based on tachistoscopic viewing

Study Finding
[5] Description of the first tachistoscope
[6] Used the divided visual field technique to assess interhemispheric transfer time
[7] Showed that a split-brain patient could name stimuli presented in the right but not in the left
visual half-field, indicating left hemispheric dominance for language production and
highlighting the role of the corpus callosum in cognition
[8] Showed that female sex hormones modulate performance in divided visual field tasks for
lexical decision, figural comparison, and face discrimination
[9] Overview about the findings obtained with the divided visual field paradigm and other
techniques in split brain patients
[10] Comprehensive review of the methodology of divided visual field paradigms
[11] Found positive correlations between divided visual field lateralization and brain activation
asymmetries during picture naming and word naming in the MRI scanner, proving that
divided visual field paradigms can be a reliable predictor of brain activation asymmetries

Fig. 1 A tachistoscope (Faculty of Psychology, Ruhr-University Bochum)

The term “tachistoscopic” (from Greek “tachistos”: very rapid

and “skopein”: to view) refers to presentation of visual stimuli for a
precisely controlled period of time [12, 13]. Typically, this period of
time is very short, e.g. in the millisecond range. Historically, tachis-
toscopic presentation of visual stimuli was performed using a family
of scientific instruments called tachistoscopes [14] (see Fig. 1).
Today it is usually performed using a standard PC and monitor.
 Tachistoscopic Viewing and Dichotic Listening 5

Fig. 2 Neuroanatomical organization of the visual system in humans. For central

fixation, information in the left visual field is processed by the right hemisphere
and information in the right visual field by the left hemisphere

How can tachistoscopic presentation of stimuli be helpful when

investigating lateralization? Due to the structure of the mammalian
visual system (see Fig. 2), a stimulus that is presented in the left
visual field is initially processed by the right hemisphere and vice
versa. Nasal portions of left and right visual input are fed into con-
tralateral hemispheres; distal portions are relayed into ipsilateral
hemisphere. Thus, by presenting visual information in only one
visual field, this information is initially only processed by the contra-
lateral hemisphere. By comparing performance on left versus right
visual field trials, the experimenter can statistically evaluate perfor-
mance differences for a specific set of stimuli between the left and
the right hemisphere. Typically, hemispheric asymmetries are
reflected by a processing advantage (e.g. faster reaction times or
higher accuracy) in the contralateral visual field (e.g. a left visual
6 Sebastian Ocklenburg

field advantage for tasks that require discrimination of complex fig-

ures or faces, or a right visual field advantage for verbal tasks [8]).
Unfortunately, stimulus processing in only one hemisphere
cannot be retained for long after onset of stimulus presentation.
On the one hand, subjects might perform saccades, that is, rapid
lateral eye movements that change the position of the foveae rela-
tive to the stimuli, possibly resulting in bilateral stimulation. On
the other hand, information transfer via the corpus callosum is
likely to take place soon after initial information processing, possi-
bly reducing hemispheric asymmetries due to bilateral stimulus
processing. Therefore tachistoscopic stimulus presentation (less
than 200 ms, ideally shorter) is a must when using the divided
visual field technique to investigate hemispheric asymmetries in
healthy humans.

1.2 Methods In general, participants should have unimpaired or corrected-to-

normal vision in both eyes, should be able to keep their head rela-
1.2.1 Prerequisites
tively motionless over prolonged periods of time while fixating the
to Use the Task
fixation cross, and should be able to press response buttons in
order to log reactions to the stimuli. Also, participants should not
have any neurological damage in the visual system. For this reason,
the task might not be optimal for certain clinical groups, and
administration in younger children might also be complicated.
Some authors recommend testing only right-handed participants
[10], at least when investigating differences between left- and
right-handers is not part of the study design. This recommenda-
tion is based on the fact that left-handers can show different pat-
terns of lateralization than right-handers in a number of cognitive
systems, e.g. for language [15]. Thus, including this group could
introduce unwanted variance in the data. Also, participants’ sex
and age should be carefully balanced, as it has been shown that
both can possibly influence performance in divided visual field par-
adigms [16, 17]. Moreover, many psychiatric disorders such as
schizophrenia [18] have been found to have an impact on perfor-
mance in divided visual field paradigms. Hence, as long as the
impact of pathological processes on lateralization is not within the
focus of the study, it is advised to test neurologically and psycho-
logically healthy participants.

1.2.2 Set-Up Compared to many other methods to assess individual lateralization,

implementation of divided visual field tasks is comparably easy and
cost-efficient. The basic set-up requires only a standard PC or other
type of personal computer, a monitor, a reaction device, and a chin
rest. For stimulus presentation, a large number of different software
tools can be used, ranging from specialized stimulus presentation
software such as E-Prime 2 (Psychology Software Tools, Inc.,
Sharpsburg, USA; http://www.pstnet.com), SuperLab 5 (Cedrus
Corporation, San Pedrom, USA; http://www.superlab.com) or
 Tachistoscopic Viewing and Dichotic Listening 7

Presentation (Neurobehavioral Systems, Inc., Berkeley, USA;

https://www.neurobs.com) to programming languages such as
Matlab (The MathWorks, Inc., Natick, USA; http://de.mathworks.
com/products/matlab/).
With regard the monitor, most research labs using visual para-
digms have traditionally used CRT (cathode-ray tube) monitors
due to superior temporal and spatial acuity which allows more reli-
able and precise onsets and offsets of visually displayed stimuli
compared to modern LCD (liquid crystal display) monitors (see
[19] for an in-depth investigation of this issue). However, since
most hardware producers have ceased production of CRT moni-
tors, and since increasingly accurate LCD monitors have become
available, more and more researchers have taken interest in using
this kind of monitor. Importantly, there seem to be large differ-
ences in suitability for vision research between different producers
and product types. For researchers interested in acquiring a new
monitor for setting up divided visual field paradigms, Ghodrati
et al. [20] provide a detailed review of the suitability of several
models of LCD monitors for vision research.
With regard to the reaction device, it has been argued that
devices connected to the computer via a serial port (such as most
specialized response boxes) are preferable to standard interface
such as USB (Universal Serial Bus) keyboards or mouse devices, as
USB can introduce a slight “lag” between the actual reaction time
and the time the reaction is recognized by the computer. Moreover,
internal wiring might cause huge differences in reaction timing
between different mouse models. Plant et al. [21] reported inter-
individual differences between different types of computer mouse
devices ranging up to 61 ms in timing error. Thus, devices measur-
ing reaction time have to be carefully chosen and tested before the
first participants can be run. As mentioned above, divided visual
field paradigms rely on presentation of visual information to only
one visual field, which results in this information being processed
only by the contralateral hemisphere. Proper administration of
such paradigms critically depends on immobilizing the head, and
instructing participants to keep body and head still during the
whole experiment. Typically, immobilization of the head is realized
using a chin rest (see Fig. 3) or a combined chin and forehead rest.
The distance between chin rest and monitor depends on the
size and dimensions of the monitor as well as on the size of the
stimuli and their distance from the fixation cross (more on this
topic in the following sections). In previous divided visual field
studies by the author of this chapter, the chin rest was placed at a
viewing distance of 57 cm from the monitor [22]. This distance is
commonly used because in this case a stimulus with the length of
1° of visual angle is 1 cm long, as the stimulus size on the monitor
is the tangent of the stimulus size in visual angle multiplied by the
distance between participant and monitor.
8 Sebastian Ocklenburg

Fig. 3 A participant demonstrating the use of a chin rest

Another factor that has to be taken into account is the testing

room. As analysis of left–right differences is central to divided
visual field paradigms, great care should be taken to avoid any lat-
eralized distraction in the testing room, such as one-sided light
sources or decorative elements. In general, a very plain room that
does not contain visual distractions is preferable. Also, dimming
the light during the task may aid in avoiding further distraction.

1.2.3 Fixation Control In order for a divided visual field paradigm to work it is essential
that stimuli supposed to be processed by the left hemisphere are
only perceived in the right visual field and vice versa. Experimentally,
this is obtained by presenting the stimuli at a specific distance from
the center of the screen. This distance differs depending on the
distance of the participant’s eyes from the screen (more on this
topic below). For this principle to work, it is essential that the par-
ticipant fixates the center of the screen throughout the experimen-
tal trial without moving his/her eyes left- or rightward. To ensure
midpoint fixation, each trial in a divided visual field paradigm starts
with the presentation of a fixation cross, with the participant being
instructed to fixate this cross throughout the whole experiment.
The fixation cross should have high visibility (e.g. black on white
background or white on black background). While many published
papers do not contain information about the size of the fixation
cross, a height and width of 1° of visual angle seems to be a com-
mon size [23]. The duration for which the fixation cross is pre-
sented on its own before a stimulus is presented varies greatly
between studies but in general seems to range from 1000 ms [24]
to 2000 ms [22].
While the fixation cross is a critical part of divided visual field
paradigms, some authors have provided evidence that simply
instructing participants to fixate the cross might not be optimal to
control for central fixation, as participants might fixate to the left or
the right of the cross on some trials [25]. Several other measures
have been suggested to ensure proper fixation control [10]. For
 Tachistoscopic Viewing and Dichotic Listening 9

example, Bourne and Hole [26] presented a single upper-case con-

sonant instead of a fixation cross for 750 ms before stimulus presen-
tation and asked the participants to verbally report this letter. Trials
in which the letter was reported incorrectly were not included in the
subsequent analysis, assuming that the participant had not properly
fixated the center of the screen in this trial. While this method argu-
ably provides a better way to control fixation than simple instruc-
tions, it increases cognitive demand and essentially creates a dual-task
paradigm, which might not be desirable for the researcher. To avoid
this issue, different methods to directly control the participant’s eye
movements have been applied. For example, Marzi and Berlucchi
[27] ensured lateralization of visual information to one hemisphere
by observing the participant’s eyes from behind a mirror that was
positioned over the fixation point. All trials in which fixation was not
maintained throughout stimulus presentation were excluded from
later analysis. Since this approach depends to a high degree on the
experimenter’s subjective assessment, more objective eye movement
measurements are preferable. For example, Meyer and Federmeier
[28] used a commercially available eye-tracking system (Applied
Science Laboratories Model 504 High-Speed Eye-Tracking System)
to track eye-movements. If participants moved their eyes from the
central fixation cross during stimulus presentation, the trial was
excluded from later analysis. The overall number of trials excluded
by this procedure was about 12 %.
Additional options to control for eye movements are available
when the participant is tested within an EEG setting. If dedicated
eye electrodes are used, automated artifact rejection procedures
can be used to exclude trials in which horizontal or vertical eye
movements occurred as determined by on amplitude changes in
the eye electrodes. For example, Lange et al. [29] excluded all tri-
als containing horizontal eye movements with an EOG (electro-
oculogram) criterion of 15 μV and vertical eye movements with an
EOG criterion of 50 μV. If no dedicated eye electrodes are used,
another possibility to control for saccadic eye movements is calcu-
lating an independent component analysis (ICA) on the EEG data
and afterwards identifying (and removing) the component(s)
reflecting saccadic eye movements. By manually adding trigger
points/markers for the typical box-shaped peaks of the saccade
component and exporting the time points to compare them with
the data from the divided visual field paradigm, the researcher
could exclude post hoc all trials in which saccades occurred
(Ocklenburg et al., unpublished data).

1.2.4 Stimulus Although protocols differ between studies, all divided visual field
Presentation in the Left paradigms have in common that at some point after initial presenta-
or Right Visual Half Field tion of the fixation cross a stimulus is presented unilaterally, in either
the left or the right visual field. In order to ensure that unilateral
stimulus presentation leads to (initial) unihemispheric processing in
10 Sebastian Ocklenburg

the contralateral hemisphere, three important factors need to be

controlled for: duration of stimulus presentation, size of the stimu-
lus, and its distance from the fixation cross. With regard to duration
of stimulus presentation, one should generally aspire to present the
stimulus as briefly as possible to prevent saccadic eye movements
towards the stimulus that would interfere with unilateral stimulus
presentation. After onset of a target (which in this case would be the
laterally presented stimulus), it takes on average 200 ms for the eyes
to start moving towards the target [30]. Thus, it is not advisable to
use stimulus durations longer than 200 ms in divided visual field
paradigms. Ideally, stimulus durations of 150 ms or less are chosen,
as some saccades can have latencies shorter than 200 ms [10].
Stimulus size and distance from the fixation cross are factors
that are somewhat interdependent. It is important to note that the
division between left and right visual field is not clear-cut, and that
the fovea is likely to be bilaterally represented and that a so-called
bilateral strip between 0.5° and 3° of the fovea projects to both
hemispheres [31]. Thus, in order to ensure unilateral stimulus pre-
sentation to the left or right hemisphere, the inside edge of a stim-
ulus should be presented at less than 2°, but better 2.5–3° away
from the central fixation cross [10]. For example, Saban-Bezalel
and Mashal [32] presented the inside edges of their stimuli 2.8°
away from the central fixation cross, while Bourne and Hole [26]
placed the inside edges of their stimuli 4° and Marzi and Berlucchi
[27] 5° from the center of the screen. Stimulus size in divided
visual field paradigms is limited by the fact that peripheral vision is
less accurate with greater eccentricity [33]. Thus, stimuli should be
kept as small as possible without compromising visibility. Most
authors keep stimulus size below 10° of visual angle [34].
Horizontally presented words are probably the most com-
monly used stimulus type in divided visual field tasks. Typically,
two main types of cognitive tasks can be performed [35], either
lexical decisions (participants have to decide whether a string of
letters represents a word or a meaningless non-word) or semantic
decisions (participants have to identify whether a word belongs to
a certain category or not). More complicated verbal stimuli have
also been used, e.g. Gold et al. [36] conducted a divided visual
field paradigm with two word metaphors. In addition to verbal
stimuli, several authors used facial stimuli [26, 34, 37, 38], emo-
tional stimuli [39–42] or complex visual figures [8, 43], but in
principle any type of visual stimulus can be used.
After stimulus presentation, most experiments involve a blank
screen or the fixation cross that is presented during the response
period. However, it has been argued that presenting a pattern mask
might be advantageous as it prevents afterimages that could con-
found stimulus duration. For instance, Michaáowski and Króliczak
[44] presented stimuli for 200 ms and then replaced them with
black and white high-contrast pattern masks, which were presented
for another 200 ms.
 Tachistoscopic Viewing and Dichotic Listening 11

After the end of the stimulus presentation segment, divided

visual field paradigms typically contain a segment in which the par-
ticipant is supposed to react to the stimuli in some way, e.g. by
pressing a button to indicate whether a stimulus belonged to a
certain category. It is important to balance use of left and right
hand for responding, e.g. by asking the participant to switch the
hand that is used to respond after a certain number of trials, in
order to prevent systematic effects of motor preferences on the
results. Also, researchers should be careful about the spatial con-
figuration of the response buttons, as it has been shown that
response times are shorter when stimulus location and response
button location match as compared to when they do not match
(the so-called “Simon effect”) [45]. Thus, using the left and right
arrow keys on a standard keyboard (or any type of button configu-
ration in which one button is on the left and the other one on the
right) might not be optimal. Instead, it is advisable to use buttons
that are above and below each other. Since subjects have to fixate
the fixation cross and cannot gaze on the response buttons, most
divided visual field paradigms rely on only a few, typically two,
response alternatives. This is done because more response alterna-
tives might introduce a memory component or lead to unwanted
head or eye movements when subjects check the response keys.
Typical types of responses are categorization (e.g. word/non-word
or normal/altered faces) as used by Hausmann and Güntürkün
[8], or Go/No-go [46]. Some older studies [47] also used oral
responses (e.g. participants verbally reported what they had seen),
but this approach has largely been discontinued because accurate
measurement of reaction time is more difficult here than for overt
motor responses such as button presses.
The number of trials in a divided visual field paradigm typically
depends on the task used (e.g. a Go/No-go task typically needs
more trials than a categorization task, since the Go and No-go
conditions have different probabilities). The number of trials
should, however, in general be rather large to allow for reliable
observation. Based on earlier studies [48, 49], Hunter and
Brysbaert [50] concluded that divided visual field tasks should at
least contain 150 observations. In general, equal numbers of left
and right visual field trials should be presented, and presentation of
these two trial types should be randomized in order to prevent
anticipatory eye movements. To further prevent any other antici-
patory effects, it is advisable to jitter the inter-stimulus interval.
Divided visual field paradigms can be quite challenging for
participants due to rapid presentation of stimuli and the unusual
requirement to keep the head still and fixate the fixation cross for a
prolonged period time. It is therefore advisable to start every
experimental session with 10–20 training trials under supervision
of the experimenter. Also, since keeping head and body still during
all experimental trials can be quite difficult for some participants,
12 Sebastian Ocklenburg

an adequate number of breaks in between experimental blocks in

which the participants are allowed to relax their head and body is
advised. While the author is not aware of any scientific investiga-
tion of break timing during divided visual field paradigms, a break
about every 10 min, with participants themselves deciding when to
carry on, appears to be advisable based on personal experience.

1.2.5 Dependent Typically, two dependent variables are analyzed in divided visual
Variables and Statistical field paradigms, that is, accuracy or error rate (in %), and median or
Analysis mean reaction time (in ms). Should there be indication for a speed-
accuracy tradeoff (e.g. some participants were very slow yet accu-
rate, while others were fast and made many errors), it could also be
advisable to additionally calculate so-called inverse efficiency scores
(reaction time divided by percentage of correct responses), a mea-
sure integrating the two parameters [51]. These values then can be
analyzed using repeated measures analysis of variance (ANOVAs),
with visual field (left and right) as within-subjects factor and all
additional factors of interest as additional within-subject (e.g. for
experimental conditions) or between-subjects factors (e.g. when
testing patients vs. controls). If the results of a divided visual field
paradigms need to be correlated with other variables, it may also
make sense to calculate a so-called laterality quotient (LQ), e.g. by
using the following formula LQ = [R − L]/[R + L] × 100, with R
indicating the value of the dependent variable for right-visual field
presentation and L indicating the value of the dependent variable
for left-visual field presentation.

1.3 Notes The obvious advantage of divided visual field paradigms is that
they offer a cheap, fast and noninvasive method of determining the
dominant hemisphere for a specific task. However, that does not
mean they are easy to implement. Great care needs to be taken to
avoid measurement errors. This is illustrated by a meta-analysis of
divided visual field studies by Voyer [49], who assessed reliability
and found that, on average, it was 0.56 for verbal tasks, implying
moderate reliability, but only 0.28 for nonverbal tasks. Importantly,
divided visual field tasks will yield reliable data, if they are conducted
correctly. This assumption has been supported by a study by
Hunter and Brysbaert [50], who compared left-handers’ reaction
time lateralization in two different divided visual field tasks (word
and picture naming) to brain activation asymmetry during a men-
tal word generation task in the fMRI scanner. The authors found
significant positive correlations between lateralization in both
divided visual field paradigms and the fMRI task (picture naming:
r = 0.77; word naming: r = 0.63), and concluded that divided visual
field paradigms can be used as a reliable predictor of language-
related brain activation asymmetries when designed carefully. Some
years later, the group of Brysbaert continued this research in a
larger sample [11], again reporting positive correlations between
 Tachistoscopic Viewing and Dichotic Listening 13

divided visual field lateralization and brain activation asymmetries

(picture naming: r = 0.65 and word naming: r = 0.64). Detailed
analysis of different participant groups showed that divided visual
field paradigms can mainly predict language lateralization in the
fMRI tasks when participants show a consistent pattern in both
divided visual field tasks, but that their predictive power is not as
good for subjects with inconsistent patterns.
One important caveat when conducting divided visual field
paradigms concerns data interpretability. While the findings of
Hunter and Brysbaert [50] and Van der Haegen et al. [11] show
positive correlations between reaction time laterality in divided
visual field paradigms and language-related brain activation asym-
metries, the researcher has to keep in mind that reaction times in
this kind of task are a rather indirect and coarse measure of brain
activation. While they might provide a general idea of whether the
left or right hemisphere is dominant for a certain task, further
insight into the relation between performance and brain activation
is difficult to support with clear evidence from the literature. For
example, it is largely unclear how the degree of lateralization in
reaction times or accuracy in these tasks relates to the magnitude of
brain activation asymmetries. Aside from the extent of brain activa-
tion asymmetries, behavioral performance measures can be modu-
lated by several other variables (e.g. by participants’ motivation,
general intelligence, cognitive control functions, fatigue, and many
others). Thus, one should be careful not to over-interpret the
results yielded by divided visual field paradigms.
This issue can be addressed by combining the divided visual
field technique with other techniques. For example, it can be com-
bined with electrophysiological techniques like EEG (electroen-
cephalogram, see Chap. 13), allowing for deeper insights into the
temporal dynamics of the neuronal process underlying task perfor-
mance [23, 39, 46, 52–55]. Moreover, they have also been used in
the fMRI scanner [56, 57], rendering it possible to relate behav-
ioral findings more directly to brain activation patterns.
One interesting tool for researchers aiming to introduce stu-
dents to the divided visual field technique is the so-called Lateralizer
software package [58] (http://cognitrn.psych.indiana.edu/
CogsciSoftware/Lateralizer/index.html) which allows students to
program and conduct divided visual field paradigms with different
types of stimuli, such as faces, words or hierarchical stimuli. Motz
et al. [58] reported positive learning outcomes as compared to
traditional lecture-based courses, suggesting that this software
package might be an interesting addition to classes about hemi-
spheric asymmetries.

1.4 Findings In general, divided visual field paradigms are not a commonly used
on the Evolution method in comparative neuroscience studies investigating lateral-
of Lateralization ization in nonhuman model species. Some work in this regard has
14 Sebastian Ocklenburg

Fig. 4 Examples for hierarchical stimuli used by Hopkins [59]. Left panel: Local
processing, Right panel: Global processing

been done in primates. Hopkins [59] used a divided visual half-

field paradigm to investigate lateralization of global and local pro-
cessing of hierarchical compound stimuli (see Fig. 4) in the
chimpanzee (Pan troglodytes). For reaction times, he found a right
visual field/left hemisphere advantage for local processing but no
visual field differences for global processing. Since the results for
local processing resembled findings in humans [60], Hopkins [59]
concluded that the chimpanzee data suggests homologous lateral-
ization in chimpanzees and humans. Similarly, using a conditional
matching-to-sample task in a divided-field paradigm, Dépy et al.
[61] found that both humans and baboons showed better distance
processing with the left compared to the right hemisphere.

2 Dichotic Listening

2.1 Introduction In addition to divided visual field paradigms, the dichotic listening
task has been the main workhorse paradigm in behavioral laterality
research on humans for more than 50 years [62] (see Table 2 for
key findings).
Based on a procedure to test attention in air traffic controllers
developed by Broadbent [77], Kimura [63, 64] published two
landmark papers in which she reported using headphones to acous-
tically present two different digits simultaneously to the left and
right ears of her participants, and asking them to report what they
heard (see [78] for a historical perspective of this early dichotic
listening research). Kimura [63, 64] showed for the first time that
there is an asymmetry of reports for right versus left ear stimuli in
most participants, the so-called right-ear advantage (REA). While
these early studies used digits as stimuli, most recent dichotic lis-
tening studies use a variation of the so-called consonant–vowel
(CV) syllables dichotic listening paradigm [65, 66]. In this
paradigm, participants wear headphones, and two different
 Tachistoscopic Viewing and Dichotic Listening 15

Table 2
Key papers for the dichotic listening task

Study Finding
[63, 64] Developed dichotic listening procedure with digits as stimuli and showed right ear advantage
for the first time
[65, 66] Developed the classic consonant–vowel version of the dichotic listening task
[67, 68] Introduction of the forced-attention version of the dichotic listening task, adding the
“forced right” and “forced” left condition to the task to assess the impact of cognitive
control on dichotic listening performance
[69] Study showing that determination of the speech-dominant hemisphere with dichotic
listening overlaps to 92 % with the results of the Wada test, proving that dichotic listening
is a reliable method to assess the language dominant hemisphere
[70] Meta-analysis showing significantly decreased lateralization in schizophrenia in studies using
the consonant–vowel or fused word dichotic listening tasks
[71] Review article highlighting the role of the corpus callosum for both bottom-up and
top-down stimulus process during dichotic listening
[72] Comprehensive fMRI study with 113 participants investigating the neural correlates of
forced-attention dichotic listening
[73, 74] Introduction of the smartphone-based iDichotic app that allows testing participants with
the forced attention version of the dichotic listening paradigm outside traditional
laboratory settings
[75] Large-scale study (1782 participants) showing that sex differences in dichotic listening are
age-dependent and small. Male younger adults showed greater asymmetry than female
younger adults, but no sex differences were found for children or older adults
[76] Large-scale study (3680 participants) showing that the right ear advantage for dichotic
stimuli increases over 60 years of age

acoustic CV stimuli (e.g. BA and GA) are presented at the same

time (see Fig. 5). Participants are asked to indicate which stimulus
he or she has heard. Typically, most participants report more syl-
lables presented to the right ear than to the left ear. This is thought
to reflect left-hemispheric dominance for speech processing. While
sensory input from the ears is transmitted to both auditory corti-
ces, the contralateral projections are stronger and may inhibit ipsi-
lateral projections so that input from the right ear is mostly
processed by the left hemisphere, and vice versa [79].
As of 2015, the search term‚ “dichotic listening” yields more
than 2800 publications in the scientific search engine PubMed
(http://www.ncbi.nlm.nih.gov/pubmed), probably more than
for any other experimental paradigm used to assess lateralization.
Its success is also reflected in the fact that Hugdahl [80] published
an edited book on dichotic listening, and that several researchers
have authored review articles covering the task in general [79,
16 Sebastian Ocklenburg

Fig. 5 Schematic representation of the consonant vowel syllable dichotic listen-

ing task. Input from the left ear is mainly processed by the right hemisphere and
input from the right ear is mainly processed by the left hemisphere

81–83], the role of attention [84], or the role the corpus callosum
in dichotic listening [71, 85]. Moreover, several review articles
cover alterations of dichotic listening performance following path-
ological conditions such as Alzheimer’s disease [86], depression
[87], learning disabilities [88], stroke [89], or schizophrenia [70,
90]. Taken together, the dichotic listening task has proven to be
one of the most versatile tasks in laterality research and is a useful
addition to the methodological repertoire for every scientist inter-
ested in investigating hemispheric asymmetries.

2.2 Methods In order to get meaningful results from the Dichotic Listening
Task, participants’ hearing capabilities should be tested using an
2.2.1 Prerequisites
audiometer (or audiometric software on a standard PC) prior to
to Use the Task
testing. Obviously, participants who are deaf or show substantially
impaired hearing in one ear need to be excluded. For example,
Hahn et al. [91] tested participants at frequencies of 500, 1000,
1500, and 3000 Hz with a MA25 audiometer (MAICO Diagnostic
GmbH, Berlin, Germany) and included only participants with inte-
raural differences smaller than 15 dB at any of these frequencies. In
addition to the interaural difference criterion, general hearing
threshold can be used as an exclusion criterion, e.g. Hahn et al.
[92] only included participants with hearing thresholds smaller
than 20 dB on both ears. While the exact exclusion criteria differ
between studies and depend on the sample that is being tested (e.g.
when testing elderly participants, the hearing threshold criterion
should be less strict), it is generally advisable to follow a rather con-
servative approach to prevent artifacts in the data. Another impor-
tant factor is the linguistic background of the participants in relation
to the language in which the stimuli are spoken. Bless et al. [74]
 Tachistoscopic Viewing and Dichotic Listening 17

tested participants with different linguistic backgrounds (English,

Danish, Norwegian, Hindi, Chinese, Spanish) with Dichotic
Listening Tasks with different stimulus languages (English,
Norwegian, German, Estonian) and found that participants showed
stronger language lateralization when they were tested in their
native language. Thus, it is advisable to test only native speakers for
the language in which the stimuli were recorded, unless testing dif-
ferences between native and nonnative speakers in dichotic listening
performance is a specific study aim. Also, participants’ sex [75], age
[76] and handedness [93] should be considered when determining
exclusion and inclusion criteria for the test sample, as these variables
are known to modulate performance in the Dichotic Listening
Task. Moreover, many psychiatric disorders like schizophrenia [90]
and neurological diseases like Alzheimer’s disease [86] have been
found to influence performance in the Dichotic Listening Task,
typically by reducing the right ear advantage. Thus, as long as the
impact of pathological processes on lateralization is not a focus of
the study, it is advisable to test neurologically and psychologically
healthy participants.

2.2.2 Set-Up As for divided visual field paradigms, implementation of dichotic

listening tasks is comparably easy and cost-efficient. Since dichotic
listening is an auditory task, the environment in which the task is
performed should be noise-free, ideally a noise-shielded room. For
stimulus presentation, most research labs use a standard PC or
other type of personal computer-. The same software tools as for
divided visual field paradigms can be used, that is, specialized stim-
ulus presentation software such as E-Prime 2 (Psychology Software
Tools, Inc., Sharpsburg, USA; http://www.pstnet.com), SuperLab
5 (Cedrus Corporation, San Pedrom, USA; http://www.superlab.
com) or Presentation (Neurobehavioral Systems, Inc., Berkeley,
USA; https://www.neurobs.com) or programming languages like
Matlab (The MathWorks, Inc., Natick, USA; http://de.mathworks.
com/products/matlab/). In addition to classic PC-based testing,
a Dichotic Listening Application for Apple iOS Smartphones is
available (http://www.dichoticlistening.com/). Bless et al. [73]
extensively tested the feasibility of conducting dichotic listening
research with this application and concluded that it represents a
valid and reliable method for conducting dichotic listening research
both in experimentally controlled as well as uncontrolled settings.
Therefore, this application opens up interesting opportunities for
researchers looking for a way to test participants easily in their own
homes or at their bedside in a hospital/clinical setting.
For stimulus presentation, high quality over-the-ear headphones
are needed. To the author’s best knowledge, there is no published
systematic comparison of different types of headphones with regard
to usability in dichotic listening tasks. One viable option is DT 770
Pro headphones (Beyerdynamic GmbH, Heilbronn, Germany), as
Another random document with
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 Samuel Chase,
William Paca,
Maryland.
Thomas Stone,
Charles Carroll, of Carrollton.

George Wythe,
Richard Henry Lee,
Thomas Jefferson,
Virginia. Benjamin Harrison,
Thomas Nelson, jr.,
Francis Lightfoot Lee,
Carter Braxton.

William Hooper,
North Carolina. Joseph Hewes,
John Penn.

Edward Rutledge,
Thomas Heyward, jr.,
South Carolina.
Thomas Lynch, jr.,
Arthur Middleton.

Button Gwinnett,
Georgia. Lyman Hall,
George Walton.
Resolved, That copies of the Declaration be sent to the several
assemblies, conventions, and committees or councils of safety, and to
the several commanding officers of the Continental Troops: That it
be PROCLAIMED in each of the United States, and at the Head of the
Army.—[Jour. Cong., vol. 1, p. 396.]
 Articles of Confederation.

Done at Philadelphia on the 9th day of July, 1778.

[While the Declaration of Independence was under consideration
in the Continental Congress, and before it was finally agreed upon,
measures were taken for the establishment of a constitutional form
of government; and on the 11th of June, 1776, it was “Resolved, That
a committee be appointed to prepare and digest the form of a
confederation to be entered into between these Colonies;” which
committee was appointed the next day, June 12, and consisted of a
member from each Colony, namely: Mr. Bartlett. Mr. S. Adams, Mr.
Hopkins, Mr. Sherman, Mr. R. R. Livingston, Mr. Dickinson, Mr.
McKean, Mr. Stone, Mr. Nelson, Mr. Hewes, Mr. E. Rutledge, and
Mr. Gwinnett. On the 12th of July, 1776, the committee reported a
draught of the Articles of Confederation, which was printed for the
use of the members under the strictest injunctions of secrecy.
This report underwent a thorough discussion in Congress, from
time to time, until the 15th of November, 1777; on which day,
“Articles of Confederation and Perpetual Union” were finally agreed
to in form, and they were directed to be proposed to the Legislatures
of all the United States, and if approved by them, they were advised
to authorize their delegates to ratify the same in the Congress of the
United States; and in that event they were to become conclusive. On
the 17th of November, 1777, the Congress agreed upon the form of a
circular letter to accompany the Articles of Confederation, which
concluded with a recommendation to each of the several Legislatures
“to invest its delegates with competent powers, ultimately, and in the
name and behalf of the State, to subscribe articles of confederation
and perpetual union of the United States, and to attend Congress for
that purpose on or before the 10th day of March next.” This letter
was signed by the President of Congress and sent, with a copy of the
articles, to each State Legislature.
 On the 26th of June, 1778, Congress agreed upon the form of a
ratification of the Articles of Confederation, and directed a copy of
the articles and the ratification to be engrossed on parchment;
which, on the 9th of July, 1778, having been examined and the
blanks filled, was signed by the delegates of New Hampshire,
Massachusetts Bay, Rhode Island and Providence Plantations,
Connecticut, New York, Pennsylvania, Virginia, and South Carolina.
Congress then directed that a circular letter be addressed to the
States whose delegates were not present, or being present, conceived
they were not authorized to sign the ratification, informing them how
many and what States had ratified the Articles of Confederation, and
desiring them, with all convenient dispatch, to authorize their
delegates to ratify the same. Of these States, North Carolina ratified
on the 21st and Georgia on the 24th of July, 1778; New Jersey on the
26th of November following; Delaware on the 5th of May, 1779;
Maryland on the 1st of March, 1781; and on the 2d of March, 1781,
Congress assembled under the new form of government.]

ARTICLES OF CONFEDERATION.

To all to whom these presents shall come,

We, the undersigned, delegates of the States affixed to our names,
send greeting:
Whereas the delegates of the United States of America in Congress
assembled did, on the fifteenth day of November, in the year of our
Lord one thousand seven hundred and seventy-seven, and in the
second year of the independence of America, agree to certain Articles
of Confederation and Perpetual Union between the States of New
Hampshire, Massachusetts Bay, Rhode Island and Providence
Plantations, Connecticut, New York, New Jersey, Pennsylvania,
Delaware, Maryland, Virginia, North Carolina, South Carolina, and
Georgia, in the words following, viz:
Articles of Confederation and Perpetual Union between the States
of New Hampshire, Massachusetts Bay, Rhode Island and
Providence Plantations, Connecticut, New York, New Jersey,
Pennsylvania, Delaware, Maryland, Virginia, North Carolina,
South Carolina, and Georgia.
 Article I. The style of this Confederacy shall be, “The United
States of America.”
Article II. Each State retains its sovereignty, freedom, and
independence, and every power, jurisdiction, and right, which is not
by this confederation expressly delegated to the United States in
Congress assembled.
Article III. The said States hereby severally enter into a firm
league of friendship with each other for their common defense, the
security of their liberties, and their mutual and general welfare;
binding themselves to assist each other against all force offered to, or
attacks made upon them, or any of them, on account of religion,
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the people of each State shall have free ingress and regress to and
from any other State, and shall enjoy therein all the privileges of
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such restrictions, shall not extend so far as to prevent the removal of
property imported into any State to any other State, of which the
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restriction shall be laid by any State on the property of the United
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If any person guilty of or charged with treason, felony, or other
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Full faith and credit shall be given in each of these States to the
records, acts, and judicial proceedings of the courts and magistrates
of every other State.
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in such manner as the Legislature of each State shall direct, to meet
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power reserved to each State to recall its delegates or any of them, at
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No State shall be represented in Congress by less than two nor by
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No vessels of war shall be kept up in time of peace by any State,
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in Congress assembled can be consulted; nor shall any State grant
commissions to any ships or vessels of war, nor letters of marque or
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Congress assembled; and then only against the kingdom or state, and
the subjects thereof, against which war has been so declared, and
under such regulations as shall be established by the United States in
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which case vessels of war may be fitted out for that occasion, and
kept so long as the danger shall continue, or until the United States
in Congress assembled shall determine otherwise.
Article VII. When land forces are raised by any State for the
common defense, all officers of, or under the rank of colonel, shall be
appointed by the Legislature of each State respectively by whom such
forces shall be raised, or in such manner as such State shall direct;
and all vacancies shall be filled up by the State which first made the
appointment.
Article VIII. All charges of war, and all other expenses that shall
be incurred for the common defense or general welfare and allowed
by the United States in Congress assembled, shall be defrayed out of
a common treasury, which shall be supplied by the several States, in
proportion to the value of all land within each State, granted to, or
surveyed for, any person, as such land and the buildings and
improvements thereon shall be estimated, according to such mode as
the United States in Congress assembled shall, from time to time,
direct and appoint.
The taxes for paying that proportion shall be laid and levied by the
authority and direction of the Legislatures of the several States,
within the time agreed upon by the United States in Congress
assembled.
Article IX. The United States in Congress assembled shall have
the sole and exclusive right and power of determining on peace and
war, except in the cases mentioned in the sixth article; of sending
and receiving embassadors; entering into treaties and alliances:
Provided, That no treaty of commerce shall be made whereby the
legislative power of the respective States shall be restrained from
imposing such imposts and duties on foreigners as their own people
are subjected to, or from prohibiting the exportation or importation
of any species of goods or commodities whatsoever; of establishing
rules for deciding, in all cases, what captures on land or water shall
be legal, and in what manner prizes taken by land or naval forces in
the service of the United States, shall be divided or appropriated; of
granting letters of marque and reprisal in times of peace; appointing
courts for the trial of piracies and felonies committed on the high
seas, and establishing courts for receiving and determining finally,
appeals in all cases of captures: Provided, That no member of
Congress shall be appointed a judge of any of the said courts.
The United States in Congress assembled shall also be the last
resort on appeal in all disputes and differences now subsisting, or
that hereafter may arise between two or more States concerning
boundary, jurisdiction, or any other cause whatever; which authority
shall always be exercised in the manner following: Whenever the
legislative or executive authority or lawful agent of any State in
controversy with another, shall present a petition to Congress,
stating the matter in question, and praying for a hearing, notice
thereof shall be given by order of Congress to the legislative or
executive authority of the other State in controversy, and a day
assigned for the appearance of the parties by their lawful agents, who
shall then be directed to appoint, by joint consent, commissioners or
judges to constitute a court for hearing and determining the matter
in question; but if they cannot agree, Congress shall name three
persons out of each of the United States, and from the list of such
persons each party shall alternately strike out one, the petitioners
beginning, until the number shall be reduced to thirteen; and from
that number not less than seven nor more than nine names, as
Congress shall direct, shall, in the presence of Congress, be drawn
out by lot; and the persons whose names shall be so drawn, or any
five of them, shall be commissioners or judges, to hear and finally
determine the controversy, so always as a major part of the judges
who shall hear the cause, shall agree in the determination; and if
either party shall neglect to attend at the day appointed, without
showing reasons which Congress shall judge sufficient, or, being
present, shall refuse to strike, the Congress shall proceed to
nominate three persons out of each State, and the Secretary of
Congress shall strike in behalf of such party absent or refusing; and
the judgment and sentence of the court to be appointed in the
manner before prescribed shall be final and conclusive; and if any of
the parties shall refuse to submit to the authority of such court or to
appear or defend their claim or cause, the court shall, nevertheless,
proceed to pronounce sentence or judgment, which shall, in like
manner, be final and decisive; the judgment or sentence, and other
proceedings, being in either case transmitted to Congress, and
lodged among the acts of Congress for the security of the parties
concerned: Provided, That every commissioner, before he sits in
judgment, shall take an oath, to be administered by one of the judges
of the supreme or superior court of the State, where the cause shall
be tried, “well and truly to hear and determine the matter in
question, according to the best of his judgment without favor,
affection, or hope of reward:” Provided, also, That no State shall be
deprived of territory for the benefit of the United States.
All controversies concerning the private right of soil claimed under
different grants of two or more States, whose jurisdictions, as they
may respect such lands, and the States which passed such grants, are
adjusted, the said grants or either of them being at the same time
claimed to have originated antecedent to such settlement of
jurisdiction, shall, on the petition of either party to the Congress of
the United States, be finally determined, as near as may be, in the
same manner as is before prescribed for deciding disputes respecting
territorial jurisdiction between different States.
The United States in Congress assembled shall also have the sole
and exclusive right and power of regulating the alloy and value of
coin struck by their own authority, or by that of the respective States;
fixing the standard of weights and measures throughout the United
States; regulating the trade and managing all affairs with the
Indians, not members of any of the States: Provided, That the
legislative right of any State within its own limits, be not infringed or
violated; establishing and regulating post-offices from one State to
another, throughout all the United States, and exacting such postage
on the papers passing through the same, as may be requisite to
defray the expenses of the said office; appointing all officers of the
land forces in the service of the United States, excepting regimental
officers; appointing all the officers of the naval forces, and
commissioning all officers whatever in the service of the United
States; making rules for the government and regulation of the said
land and naval forces, and directing their operations.
The United States in Congress assembled shall have authority to
appoint a committee to sit in the recess of Congress, to be
denominated “a Committee of the States,” and to consist of one
delegate from each State, and to appoint such other committees and
civil officers as may be necessary for managing the general affairs of
the United States, under their direction; to appoint one of their
number to preside; provided that no person be allowed to serve in
the office of president more than one year in any term of three years;
to ascertain the necessary sums of money to be raised for the service
of the United States, and to appropriate and apply the same for
defraying the public expenses; to borrow money or emit bills on the
credit of the United States, transmitting every half-year to the
respective States, an account of the sums of money so borrowed or
emitted; to build and equip a navy; to agree upon the number of land
forces, and to make requisitions from each State for its quota, in
proportion to the number of white inhabitants in such State, which
requisitions shall be binding; and thereupon the Legislature of each
State shall appoint the regimental officers, raise the men, and clothe,
arm, and equip them in a soldier-like manner, at the expense of the
United States; and the officers and men so clothed, armed, and
equipped, shall march to the place appointed, and within the time
agreed on by the United States in Congress assembled; but if the
United States in Congress assembled shall, on consideration of
circumstances, judge proper that any State should not raise men, or
should raise a smaller number than its quota, and that any other
State should raise a greater number of men than the quota thereof,
such extra number shall be raised, officered, clothed, armed, and
equipped in the same manner as the quota of each State, unless the
Legislature of such State shall judge that such extra number cannot
be safely spared out of the same; in which case they shall raise,
officer, clothe, arm, and equip as many of such extra number as they
judge can be safely spared. And the officers and men so clothed,
armed, and equipped shall march to the place appointed, and within
the time agreed on by the United States in Congress assembled.
The United States in Congress assembled shall never engage in a
war, nor grant letters of marque and reprisal in time of peace, nor
enter into any treaties or alliances, nor coin money, nor regulate the
value thereof, nor ascertain the sums and expenses necessary for the
defense and welfare of the United States or any of them, nor emit
bills, nor borrow money on the credit of the United States, nor
appropriate money, nor agree upon the number of vessels of war to
be built or purchased, or the number of land or sea forces to be
raised, nor appoint a commander-in-chief of the Army or Navy,
unless nine States assent to the same; nor shall a question on any
other point, except for adjourning from day to day, be determined,
unless by the votes of a majority of the United States in Congress
assembled.
The Congress of the United States shall have power to adjourn to
any time within the year, and to any place within the United States,
so that no period of adjournment be for a longer duration than the
space of six months; and shall publish the journal of their
proceedings monthly, except such parts thereof relating to treaties,
alliances, or military operations, as in their judgment require
secrecy; and the yeas and nays of the delegates of each State on any
question, shall be entered on the journal, when it is desired by any
delegate; and the delegates of a State, or any of them, at his or their
request, shall be furnished with a transcript of the said journal,
except such parts as are above excepted, to lay before the Legislature
of the several States.
Article X. The committee of the States, or any nine of them, shall
be authorized to execute, in the recess of Congress, such of the
powers of Congress as the United States in Congress assembled, by
the consent of nine States, shall, from time to time, think expedient
to vest them with: Provided, That no power be delegated to the said
committee, for the exercise of which, by the Articles of
Confederation, the voice of nine States in the Congress of the United
States assembled is requisite.
Article XI. Canada, acceding to this confederation, and joining in
the measures of the United States, shall be admitted into, and
entitled to, all the advantages of this Union; but no other colony shall
be admitted into the same, unless such admission be agreed to by
nine States.
Article XII. All bills of credit emitted, moneys borrowed, and
debts contracted, by or under the authority of Congress, before the
assembling of the United States, in pursuance of the present
confederation, shall be deemed and considered as a charge against
the United States, for payment and satisfaction whereof the said
United States and the public faith are hereby solemnly pledged.
Article XIII. Every State shall abide by the determinations of the
United States in Congress assembled, on all questions which by this
confederation are submitted to them. And the articles of this
confederation shall be inviolably observed by every State, and the
union shall be perpetual; nor shall any alteration at any time
hereafter be made in any of them, unless such alteration be agreed to
in a Congress of the United States, and be afterwards confirmed by
the Legislatures of every State.
And whereas it has pleased the Great Governor of the world to
incline the hearts of the Legislatures we respectively represent in
Congress, to approve of, and to authorize us to ratify the said articles
of confederation and perpetual union: Know ye, That we, the
undersigned delegates, by virtue of the power and authority to us
given for that purpose, do, by these presents, in the name and in
behalf of our respective constituents, fully and entirely ratify and
confirm each and every of the said Articles of Confederation and
Perpetual Union, and all and singular the matters and things therein
contained. And we do further solemnly plight and engage the faith of
our respective constituents, that they shall abide by the
determinations of the United States, in Congress assembled, on all
questions which, by the said confederation, are submitted to them;
and that the articles thereof shall be inviolably observed by the States
we respectively represent; and that the union shall be perpetual.
In witness whereof we have hereunto set our hands, in Congress.
Done at Philadelphia, in the State of Pennsylvania, the ninth day
of July, in the year of our Lord one thousand seven hundred and
seventy-eight, and in the third year of the Independence of
America.
On the part and behalf of the State of New Hampshire.—Josiah
Bartlett, John Wentworth, jr., August 8, 1778.
On the part and behalf of the State of Massachusetts Bay.—John
Hancock, Samuel Adams, Elbridge Gerry, Francis Dana, James
Lovell, Samuel Holten.
On the part and in behalf of the State of Rhode Island and
Providence Plantations.—William Ellery, Henry Marchant, John
Collins.
On the part and behalf of the State of Connecticut.—Roger
Sherman, Samuel Huntington, Oliver Wolcott, Titus Hosmer,
Andrew Adams.
On the part and behalf of the State of New York.—Jas. Duane, Fra.
Lewis, Wm. Duer, Gouv. Morris.
On the part and in behalf of the State of New Jersey.—Jno.
Witherspoon, Nath. Scudder, Nov. 26, 1778.
On the part and behalf of the State of Pennsylvania.—Robt.
Morris, Daniel Roberdeau, Jona. Bayard Smith, William Clingan,
Joseph Reed, July 22d, 1778.
On the part and behalf of the State of Delaware.—Thos. McKean,
Feb. 13, 1779, John Dickinson, May 5, 1779, Nicholas Van Dyke.
On the part and behalf of the State of Maryland.—John Hanson,
March 1, 1781, Daniel Carroll, March 1, 1781.
On the part and behalf of the State of Virginia.—Richard Henry
Lee, John Banister, Thomas Adams, Jno. Harvie, Francis Lightfoot
Lee.
 On the part and behalf of the State of North Carolina.—John
Penn, July 21, 1778, Corns. Harnett, Jno. Williams.
On the part and behalf of the State of South Carolina.—Henry
Laurens, William Henry Drayton, Jno. Mathews, Richard Hutson,
Thomas Heyward, Jr.
On the part and behalf of the State of Georgia.—Jno. Walton, July
24, 1778, Edw. Telfair, Edw. Langworthy.
 Ordinance of 1787.

An Ordinance for the Government of the Territory of the United

States Northwest of the Ohio River. [In Congress, July 13, 1787.]
Be it ordained by the United States in Congress assembled, That
the said Territory, for the purposes of temporary government, be one
district; subject, however to be divided into two districts, as future
circumstances may, in the opinion of Congress, make it expedient.
Be it ordained by the authority aforesaid, That the estates both of
resident and non-resident proprietors in the said Territory, dying
intestate, shall descend to and be distributed among their children,
and the descendants of a deceased child, in equal parts; the
descendants of a deceased child or grandchild to take the share of
their deceased parent in equal parts among them; and where there
shall be no children or descendants; then in equal parts to the next of
kin, in equal degree; and among collaterals, the children of a
deceased brother or sister of the intestate shall have, in equal parts
among them, their deceased parents’ share; and there shall, in no
case, be a distinction between kindred of the whole and half blood;
saving in all cases to the widow of the intestate, her third part of the
real estate for life, and one-third part of the personal estate; and this
law relative to descents and dower shall remain in full force until
altered by the Legislature of the district. And until the governor and
judges shall adopt laws as hereinafter mentioned, estates in the said
Territory may be devised or bequeathed by wills in writing, signed
and sealed by him or her, in whom the estate may be, (being of full
age,) and attested by three witnesses; and real estates may be
conveyed by lease and release, or bargain and sale, signed, sealed,
and delivered by the person, being of full age, in whom the estate
may be and attested by two witnesses, provided such wills be duly
proved, and such conveyances be acknowledged, or the execution
thereof duly proved and be recorded within one year, after proper
magistrates, courts, and registers shall be appointed for that
purpose; and personal property may be transferred by delivery,
saving, however, to the French and Canadian inhabitants, and other
settlers of the Kaskaskies, Saint Vincent’s, and the neighboring
villages, who have heretofore professed themselves citizens of
Virginia, their laws and customs now in force among them, relative
to the descent and conveyance of property.
Be it ordained by the authority aforesaid, That there shall be
appointed, from time to time, by Congress, a governor, whose
commission shall continue in force for the term of three years, unless
sooner revoked by Congress; he shall reside in the district, and have
a freehold estate therein, in one thousand acres of land, while in the
exercise of his office.
There shall be appointed, from time to time, by Congress, a
secretary, whose commission shall continue in force for four years,
unless sooner revoked; he shall reside in the district, and have a
freehold estate therein, in five hundred acres of land, while in the
exercise of his office; it shall be his duty to keep and preserve the acts
and laws passed by the Legislature, and the public records of the
district, and the proceedings of the governor in his executive
department; and transmit authentic copies of such acts and
proceedings every six months to the secretary of Congress. There
shall also be appointed a court, to consist of three judges, any two of
whom to form a court, who shall have a common law jurisdiction,
and reside in the district, and have each therein a freehold estate, in
five hundred acres of land, while in the exercise of their offices, and
their commissions shall continue in force during good behavior.
The governor and judges, or a majority of them, shall adopt and
publish in the district such laws of the original States, criminal and
civil, as may be necessary, and best suited to the circumstances of the
district, and report them to Congress, from time to time, which laws
shall be in force in the district until the organization of the general
assembly therein, unless disapproved of by Congress; but afterwards
the legislature shall have authority to alter them as they shall think
fit.
The governor for the time being shall be commander-in-chief of
the militia; appoint and commission all officers in the same below
the rank of general officers. All general officers shall be appointed
and commissioned by Congress.
Previous to the organization of the General Assembly, the governor
shall appoint such magistrates and other civil officers in each county
or township as he shall find necessary for the preservation of the
peace and good order in the same. After the General Assembly shall
be organized, the powers and duties of magistrates and other civil
officers shall be regulated and defined by the said assembly; but all
magistrates and other civil officers, not herein otherwise directed,
shall, during the continuance of this temporary government, be
appointed by the governor.
For the prevention of crimes and injuries, the laws to be adopted
or made shall have force in all parts of the district, and for the
execution of process, criminal and civil, the governor shall make
proper divisions thereof; and he shall proceed from time to time, as
circumstances may require, to lay out the parts of the district in
which the Indian titles shall have been extinguished, into counties
and townships, subject, however, to such alterations as may
thereafter be made by the Legislature.
So soon as there shall be five thousand free male inhabitants of full
age in the district, upon giving proof thereof to the governor, they
shall receive authority, with time and place, to elect representatives
from their counties or townships, to represent them in the General
Assembly; Provided, That for every five hundred free male
inhabitants, there shall be one representative; and so on,
progressively, with the number of free male inhabitants, shall the
right of representation increase, until the number of representatives
shall amount to twenty-five; after which the number and proportion
of representatives shall be regulated by the Legislature: Provided,
That no Person be eligible or qualified to act as a representative
unless he shall have been a citizen of one of the United States three
years, and be a resident in the district, or unless he shall have resided
in the district three years; and in either case, shall likewise hold in
his own right, in fee simple, two hundred acres of land within the
same: Provided, also, That a freehold in fifty acres of land in the
district, having been a citizen of one of the States, and being resident
in the district, or the like free hold and two years’ residence in the
district, shall be necessary to qualify a man as an elector of a
representative.
The representatives thus elected shall serve for the term of two
years; and in case of the death of a representative, or removal from
office, the governor shall issue a writ to the county or township for
which he was a member to elect another in his stead, to serve for the
residue of the term.
The General Assembly, or Legislature, shall consist of the
governor, legislative council, and a house of representatives. The
legislative council shall consist of five members, to continue in office
five years, unless sooner removed by Congress, any three of whom to
be a quorum; and the members of the council shall be nominated
and appointed in the following manner, to wit: As soon as
representatives shall be elected, the governor shall appoint a time
and place for them to meet together, and when met, they shall
nominate ten persons, residents in the district, and each possessed of
a freehold in five hundred acres of land, and return their names to
Congress; five of whom Congress shall appoint and commission to
serve as aforesaid; and whenever a vacancy shall happen in the
council, by death or removal from office, the house of representatives
shall nominate two persons, qualified as aforesaid, for each vacancy,
and return their names to Congress; one of whom Congress shall
appoint and commission for the residue of the term. And every five
years, four months at least before the expiration of the time of
service of the members of the council, the said house shall nominate
ten persons, qualified as aforesaid, and return their names to
Congress; five of whom Congress shall appoint and commission to
serve as members of the council five years, unless sooner removed.
And the governor, legislative council, and house of representatives,
shall have authority to make laws in all cases for the good
government of the district, not repugnant to the principles and
articles in this ordinance established and declared, and all bills
having passed by a majority in the house, and by a majority in the
council, shall be referred to the governor for his assent; but no bill or
legislative act whatever shall be of any force without his assent. The
governor shall have power to convene, prorogue, and dissolve the
General Assembly when in his opinion it shall be expedient.
 The governor, judges, legislative council, secretary, and such other
officers as Congress shall appoint in the district, shall take an oath or
affirmation of fidelity and of office, the governor before the President
of Congress, and all other officers before the governor. As soon as a
Legislature shall be formed in the district, the council and house
assemble, in one room, shall have authority, by joint ballot, to elect a
delegate to Congress, who shall have a seat in Congress, with a right
of debating, but not of voting during this temporary government.
And for extending the fundamental principles of civil and religious
liberty, which form the basis whereon these republics, their laws and
constitutions, are erected; to fix and establish those principles as the
basis of all laws, constitutions, and governments, which forever
hereafter shall be formed in the said Territory; to provide, also, for
the establishment of States, and permanent government therein, and
for their admission to a share in the Federal councils on an equal
footing with the original States, at as early periods as may be
consistent with the general interest:
It is hereby ordained and declared, by the authority aforesaid,
That the following articles shall be considered as articles of compact,
between the original States and the people and States in the said
Territory, and forever remain unalterable, unless by common
consent, to wit:
Article 1. No person, demeaning himself in a peaceable and
orderly manner, shall ever be molested on account of his mode of
worship or religious sentiments, in the said Territory.
Art. 2. The inhabitants of the said Territory shall always be
entitled to the benefits of the writ of habeas corpus, and of the trial
by jury; of a proportionate representation of the people in the
Legislature, and of judicial proceedings according to the course of
the common law. All persons shall be bailable, unless for capital
offenses, where the proof shall be evident or the presumption great.
All fines shall be moderate; and no cruel or unusual punishments
shall be inflicted. No man shall be deprived of his liberty or property
but by the judgment of his peers, or the law of the land; and should
the public exigencies make it necessary for the common preservation
to take any person’s property, or to demand his particular services,
full compensation shall be made for the same. And, in the just
preservation of rights and property, it is understood and declared
that no law ought ever to be made, or have force in the said Territory,
that shall, in any manner whatever, interfere with, or affect, private
contracts or engagements, bona fide and without fraud, previously
formed.
Art. 3. Religion, morality, and knowledge, being necessary to good
government and the happiness of mankind, schools and the means of
education shall forever be encouraged. The utmost good faith shall
always be observed toward the Indians; their lands and property
shall never be taken from them without their consent; and in their
property, rights, and liberty they shall never be invaded or disturbed,
unless in just and unlawful wars authorized by Congress; but laws
founded in justice and humanity shall, from time to time, be made
for preventing wrongs being done to them, and for preserving peace
and friendship with them.
Art. 4. The said Territory, and the States which may be formed
therein, shall ever remain a part of this confederacy of the United
States of America, subject to the Articles of Confederation, and to
such alterations therein as shall be constitutionally made; and to all
the acts and ordinances of the United States in Congress assembled,
conformable thereto. The inhabitants and settlers in the Territory
shall be subject to pay a part of the Federal debts, contracted or to be
contracted, and a proportional part of the expenses of Government,
to be apportioned on them by Congress, according to the same
common rule and measure by which apportionments thereof shall be
made on the other States; and the taxes for paying their proportion
shall be laid and levied by the authority and direction of Legislatures
of the district or districts, or new States, as in the original States,
within the time agreed upon by the United States in Congress
assembled. The Legislatures of those districts, or new States shall
never interfere with the primary disposal of the soil by the United
States in Congress assembled, nor with any regulations Congress
may find necessary for securing the title in such soil to the bona fide
purchasers. No tax shall be imposed on lands the property of the
United States; and in no case shall non-resident proprietors be taxed
higher than residents. The navigable waters leading into the
Mississippi and St. Lawrence, and the carrying places between the
same, shall be common highways, and forever free, as well to the
inhabitants of the said Territory as to the citizens of the United
States, and those of any other States that may be admitted into the
confederacy, without any tax, impost, or duty therefor.
Art. 5. There shall be formed in the said Territory not less than
three, nor more than five States; and the boundaries of the States, as
soon as Virginia shall alter her act of cession, and consent to the
same, shall become fixed and established as follows, to wit: The
western State in the said territory shall be bounded by the
Mississippi, the Ohio, and Wabash Rivers; a direct line drawn from
the Wabash and Post Vincents, due north, to the territorial line
between the United States and Canada; and by the said territorial
line to the Lake of the Woods and Mississippi. The middle States
shall be bounded by the said direct line, the Wabash, from Post
Vincents to the Ohio, by the Ohio, by a direct line drawn due north
from the mouth of the Great Miami to the said territorial line, and by
the said territorial line. The eastern State shall be bounded by the
last mentioned direct line, the Ohio, Pennsylvania, and the said
territorial line: Provided, however, And it is further understood and
declared that the boundaries of these three States shall be subject so
far to be altered, that, if Congress shall hereafter find it expedient,
they shall have authority to form one or two States in that part of the
said Territory which lies north of an east and west line drawn
through the southerly bend or extreme of Lake Michigan. And
whenever any of the said States shall have sixty thousand free
inhabitants therein, such State shall be admitted, by its delegates,
into the Congress of the United States, on an equal footing with the
original States in all respects whatever; and shall be at liberty to form
a permanent constitution and State government: Provided, The
constitution and government so to be formed shall be republican,
and in conformity to the principles contained in these articles; and,
so far as it can be consistent with the general interest of the
confederacy, such admission shall be allowed at an earlier period,
and when there may be a less number of free inhabitants in the State
than sixty thousand.
Art. 6. There shall be neither slavery nor involuntary servitude in
the said Territory, otherwise than in the punishment of crimes,
whereof the party shall have been duly convicted: Provided always,
That any person escaping into the same, from whom labor or service
is lawfully claimed in any one of the original States, such fugitive