Professional Documents
Culture Documents
6e
Robert J. Brooker
TABLE OF CONTENTS vii
10.4 Sizes of Eukaryotic Genomes and 14.3 Regulation of the trp Operon 349 18.3 Bacteriophage λ Reproductive
Repetitive Sequences 235 14.4 Translational and Posttranslational Cycle 444
10.5 Structure of Eukaryotic Chromosomes in Regulation 353 18.4 HIV Reproductive Cycle 450
Nondividing Cells 237 14.5 Riboswitches 354
10.6 Structure of Eukaryotic Chromosomes
During Cell Division 243 19 GENE MUTATION AND DNA
15
REPAIR 461
GENE REGULATION IN
11
EUKARYOTES I:
19.1 Effects of Mutations on Gene Structure
TRANSCRIPTIONAL AND
DNA REPLICATION 252 TRANSLATION
and Function 462
REGULATION 361 19.2 Random Nature of Mutations 468
11.1 Structural Overview of DNA 19.3 Spontaneous Mutations 470
Replication 252 15.1 Regulatory Transcription Factors 362 19.4 Induced Mutations 475
11.2 Bacterial DNA Replication: The 15.2 Chromatin Remodeling, Histone 19.5 DNA Repair 479
Formation of Two Replication Forks at Variants, and Histone
the Origin of Replication 256 Modification 369
11.3 Bacterial DNA Replication: Synthesis of
New DNA Strands 259
11.4 Bacterial DNA Replication: Chemistry
15.3 DNA Methylation 376
15.4 Insulators 378
20 RECOMBINATION,
IMMUNOGENETICS, AND
TRANSPOSITION 491
15.5 The ENCODE Project 379
and Accuracy 266 20.1 Homologous Recombination 491
15.6 Regulation of Translation 380
11.5 Eukaryotic DNA Replication 268 20.2 Immunogenetics 497
20.3 Transposition 499
PA R T I V
MOLECULAR PROPERTIES
16 GENE REGULATION IN
EUKARYOTES II:
EPIGENETICS 388
PA R T V
OF GENES 278
16.1 Overview of Epigenetics 388 GENETIC TECHNOLOGIES 511
12.1
12.2
Overview of Transcription 278
Transcription in Bacteria 281
16.4 Epigenetics and Environmental
Agents 400
16.5 Role of Epigenetics in Cancer 405
21 MOLECULAR
TECHNOLOGIES 511
12.3 Transcription in Eukaryotes 286 21.1 Gene Cloning Using Vectors 512
21.2 Polymerase Chain Reaction 519
12.4
12.5
RNA Modification 291
A Comparison of Transcription
and RNA Modification in Bacteria
17 NON-CODING RNAs 411
21.3
21.4
DNA Sequencing 524
Gene Mutagenesis 526
and Eukaryotes 300 17.1 Overview of Non-coding RNAs 412 21.5 Blotting Methods to Detect Gene
17.2 Non-coding RNAs: Effects on Chromatin Products 529
BACTERIA 336
23.4 Physical Mapping via Cloning and DNA 25.6 Personalized Medicine 634 28.1 Overview of Complex and Quantitative
Sequencing 570 Traits 707
23.5 Genome-Sequencing Projects 574
23.6 Metagenomics 582 26 DEVELOPMENTAL
GENETICS 643
28.2 Statistical Methods for Evaluating
Quantitative Traits 709
28.3 Polygenic Inheritance 712
24
26.1 Overview of Animal Development 643 28.4 Identification of Genes that Control
GENOMICS II: FUNCTIONAL
GENOMICS, PROTEOMICS, AND 26.2 Invertebrate Development 647 Quantitative Traits 715
BIOINFORMATICS 589 26.3 Vertebrate Development 659 28.5 Heritability 717
26.4 Plant Development 662 28.6 Selective Breeding 722
24.1 Functional Genomics 590
29
26.5 Sex Determination in Animals 666
24.2 Proteomics 595
27
24.3 Bioinformatics 600 EVOLUTIONARY GENETICS 732
25
brookergenetics6e
MEDICAL GENETICS AND 27.5 Migration 692
CANCER 611 Appendix B
27.6 Nonrandom Mating 692
Solutions to Even-Numbered
25.1 Inheritance Patterns of Genetic 27.7 Sources of New Genetic Variation 694 Problems and All Comprehension
Diseases 612 and Concept Check Questions B-1
25.2 Detection of Disease-Causing Alleles via Glossary G-1
Haplotypes 618
Index I-1
DEDICATION
I
students may be provided with online lectures, “flipping the class-
room” typically gives students more responsibility for understanding
the textbook material on their own. Along these lines, Genetics:
Analysis & Principles, Sixth Edition, is intended to provide students
n the sixth edition of Genetics: Analysis & Principles, the with a resource that can be effectively used outside of the classroom.
content has been updated to reflect current trends in the field. In Here are several of the key pedagogical features:
addition, the presentation of the content has been improved in a
∙
NEW! A new feature called Genetic TIPS provides a
way that fosters active learning. As an author, researcher, and
teacher, I want a textbook that gets students actively involved in consistent approach to help students solve problems in
learning genetics. To achieve this goal, I have worked with a genetics. This approach has three components. First, the
talented team of editors, illustrators, and media specialists who student is made aware of the Topic at hand. Second, the
have helped me to make the sixth edition of Genetics: Analysis & question is evaluated with regard to the Informaiton that is
Principles a fun learning tool. available to the student. Finally, the student is guided
Overall, an effective textbook needs to accomplish four through one or more Problem-Solving Strategies to tackle
goals.14 First, it needsCHAP toT provide
E R 1 :: comprehensive,
OVERVIEW OF GENETICS accurate, and up- the question.
to-date content in its field. Second, it needs to expose students to
the techniques and skills they will need to become successful in
that field. words,
Third, what an effective
scientifictextbook
question should
was thehave pedagogical
researcher trying
features, such to answer?
as formative assessment, that foster student learn- GENETIC TIPS THE QUESTION: All of the Genetic TIPS
begin with a question. As an example, let’s consider the following
ing. And 3. finally,
Next, the figure follows
it should inspirethe experimental
students so theysteps wantthe to scientist
pursue question:
that field as took to test the
a career. Thehypothesis.
hard workEach thatfeatured
has gone experiment con-
into the sixth The coding strand of DNA in a segment of a gene is as follows:
edition oftains two parallel
Genetics: Analysisillustrations labeled has
& Principles Experimental
been aimed Levelat ATG GGC CTT AGC. This strand carries the information to make a
achieving and Conceptual
all four of theseLevel. goals!The Experimental Level helps you region of a polypeptide with the amino acid sequence, methionine-
to understand the techniques followed. The Conceptual glycine-leucine-serine. What would be the consequences if a mutation
Level helps you to understand what is actually happening changed the second cytosine (C) in this sequence to an adenine (A)?
at each step in the procedure.
FLIPPING
4. The raw data THE for eachCLASSROOM
experiment are then presented. T OPIC: What topic in genetics does this question address? The
5. Last, an interpretation of the data is offered within the text. topic is gene expression. More specifically, the question is about
A recent trend in science education is the phenomenon that is some- the relationship between a gene sequence and the genetic code.
The rationale behind this approach is that it enables you to see the
timesexperimental
called “flipping the classroom.” This phrase refers to the idea
process from beginning to end. As you read through
that some of the activities that usedwillto be done I NFORMATION: What information do you know based on the
the chapters, the experiments help youintoclass
see theare relationship
now done
question and your understanding of the topic? In the question,
outside of class, and vice versa.
between science and scientific theories. For example, instead of spending
the entire As class time lecturing over textbook and other materials, you are given the base sequence of a short segment of a gene and
a student of genetics, you will be given the opportunity told that one of the bases has been changed. From your understanding
sometoofinvolve
the classyourtimemind is in
spent engaging students
the experimental process. in various
As you are activi-
read- of the topic, you may remember that a polypeptide sequence is
ties, ing
such anas problem solving,
experiment, you mayworking through
find yourself case about
thinking studies, and
different determined by reading the mRNA (transcribed from a gene) in
designing experiments.
approaches This approach
and alternative is called
hypotheses. activepeople
Different learning.can For
view groups of three bases called codons.
manytheinstructors,
same datathe and classroom
arrive at hasverybecome
differentmore learner centered
conclusions. As you
rather teacherthrough
progress centered. theAexperiments
learner-centeredin thisclassroom
book, youprovides
will enjoy a P ROBLEM-SOLVING S TRATEGY: Compare and contrast.
rich genetics
environment far morein which
if youstudents can interact
try to develop your own withskills
eachatother and
formulat- One strategy to solve this problem is to compare the mRNA
withingtheirhypotheses,
instructors.designingInstructorsexperiments,
and fellow students often provide
and interpreting data. sequence (transcribed from this gene) before and after the mutation:
Also, some
formative of the questions in the
assessment—immediate problem
feedback thatsets are aimed
helps at refin-
each student Original: AUG GGC CUU AGC
ing these
understand skills.
if his or her learning is on the right track. Mutant: AUG GGC AUU AGC
Finally,
What are some it is worthwhile
advantages to of point
activeoutlearning?
that science is a social
Educational
discipline. As you develop your skills at scrutinizing
studies reveal that active learning usually promotes greater learning experiments, ANSWER: The mutation has changed the sequence of bases in the
gains.it In
is fun to discuss
addition, activeyour ideasoften
learning with focuses
other people,
on skillincluding
developmentfellow mRNA so that the third codon has changed from CUU to AUU.
students and faculty members. Keep in mind that you do not need Because codons specify amino acids, this may change the third
rather than on the memorization of facts that are easily forgotten.
to “know all the answers” before you enter into a scientific discus- amino acid to something else. Note: If you look ahead to Chapter 13
Students become trained to “think like scientists” and to develop a
(see Table 13.1), you will see that CUU specifies leucine, whereas
skill sion. Instead,
set that enables it isthem
moretorewarding to view
apply scientific science asAan
reasoning. ongoing
common AUU specifies isoleucine. Therefore, you would predict that the mu-
and never-ending dialogue.
concern among instructors who are beginning to try out active learn- tation would change the third amino acid from leucine to isoleucine.
ing is that they think they will have less time to teach and therefore
will cover
Genetic less material.
TIPS Will However,
HelpthisYou maytonot be the case.
Improve Although
Your
Problem-Solving Skills Throughout Chapters 2 through 29, each chapter will contain sev- ix
As your progress through this textbook, your learning will involve eral Genetic TIPS. Some of these will be within the chapter itself
two general goals: and some will precede the problem sets that are at the end of each
x PREFACE
∙
Genes → Traits: Because genetics is such a broad discipline,
ranging from the molecular level to populations, many SIGNIFICANT CONTENT CHANGES
instructors have told us that it is a challenge for students to IN THE SIXTH EDITION
see both “the forest and the trees.” It is commonly mentioned
that students often have trouble connecting the concepts they ∙
NEW! A new problem-solving feature called Genetic TIPS
have learned in molecular genetics with the traits that occur has been added to the sixth edition. The Genetic TIPS are
at the level of a whole organism (i.e., What does found within each chapter and three or four are found at the
transcription have to do with blue eyes?). To try to make this end of each chapter.
connection more meaningful, certain figure legends in each ∙
NEW! The topic of Epigenetics has been expanded to a
chapter, designated Genes → Traits, remind students that whole chapter, which is now Chapter 16.
molecular and cellular phenomena ultimately lead to the ∙
NEW! A new chapter on non-coding RNA has been added,
traits that are observed in each species (see Figure 14.8). which is Chapter 17. This long-overdue chapter is in
∙
Learning Outcomes: Each section of every chapter begins response to a remarkable explosion in our appreciation for
with a set of learning outcomes. These outcomes help the roles of non-coding RNAs in many aspects of molecular
students understand what they should be able to do once they biology. Note: Although two new chapters have been added
have mastered the material in that section. to this edition, the overall page length of the sixth edition is
∙
Formative Assessment: When students are expected to learn not longer than the fifth edition.
textbook material on their own, it is imperative that they are ∙
NEW! CRISPR-Cas systems: The role of the CRISPR-Cas
regularly given formative assessment so they can gauge system in providing prokaryotes with a genome defense
whether they are mastering the material. Formative mechanism is described in Chapter 17, and its use by
assessment is a major feature of this textbook and is bolstered researchers to mutate genes is described in Chapter 21.
by Connect—a state-of-the art digital assignment and
assessment platform. In Genetics: Analysis & Principles, Sixth
Examples of Specific Content Changes
Edition, formative assessment is provided in multiple ways.
to Individual Chapters
1. As mentioned, a new feature called Genetic TIPS is ∙
Chapter 2. Mendelian Inheritance: Several Genetic TIPS
aimed at helping students refine their problem solving
have been added to help students work through problem-
skills.
solving strategies involving Mendelian inheritance.
2. Each section of every chapter ends with multiple-choice ∙
Chapter 3. Chromosome Transmission During Cell Division
questions. Also, compared with the previous edition, many
and Sexual Reproduction: The discussion of the random
chapters in the sixth edition are divided into more sections
alignment of homologs during metaphase of meiosis I was
that are shorter in length. Formative assessment at the end
expanded.
of each section allows students to evaluate their mastery of ∙
Chapter 4. Extensions of Mendelian Inheritance: The topic
the material before moving on to the next section.
of gene interaction was streamlined to focus primarily on
3. Most figures have Concept Check questions so students
examples in which the underlying molecular mechanisms are
can determine if they understand the key points in the
known.
figure. ∙
Chapter 5. Non-Mendelian Inheritance:A common
4. Extensive end-of chapter questions continue to provide
misconception among students is that you can use a Punnett
students with feedback regarding their mastery of the
square to deduce nonMendelian inheritance patterns.
material.
Throughout the chapter, this misconception has been laid to
5. The textbook material is supported by digital learning
rest, and students are given effective strategies to predict
tools found in Connect. Questions and activities are
offspring genotypes and phenotypes.
assignable in Connect, and students also have access to ∙
Chapter 6. Genetic and Linkage Mapping in Eukaryotes:
our valuable adaptive study tool, SmartBook. With this
When looking at experiments involving linkage, student
tool, students are repeatedly given questions regarding
often find it very difficult to identify the recombinant
the textbook material, and depending on their answers,
offspring. In various parts of the chapter, a strong effort has
they may advance ahead in their reading, or they are
been made to make it clear that recombinant offspring have
given specific advice on what textbook material to go
inherited a chromosome that is the product of a crossover.
back and review.
Along these same lines, a new figure (see Figure 6.6) has
Overall, the pedagogy of Genetics: Analysis & Principles, been added involving the experiments of Curt Stern showing
sixth edition, has been designed to foster student learning. Instead of that recombinant offspring carry chromosomes that are the
being a collection of facts and figures, Genetics: Analysis & Prin- product of a crossover. Also, Figure 6.8 has been revised to
ciples, Sixth Edition, by Rob Brooker, is intended to be an engaging emphasis this point.
and motivating textbook in which formative assessment allows stu- ∙
Chapter 7. Genetic Transfer and Mapping in Bacteria:
dents to move ahead and learn the material in a productive way. We Figure 7.13 is a new figure showing the increase in methicillin
welcome your feedback so we can make future editions even better! resistance in certain Staphylococcus aureus strains.
PREFACE xi
∙
Chapter 8. Variation in Chromosome Structure and Number: ∙
Chapter 21. DNA Technologies: A new subsection has
Several Genetic TIPS have been added to help students solve been added on gene mutagenesis, which includes a
problems that involve changes in chromosome structure and description of the Crispr-Cas system for inactivating
chromosome number. and mutating genes.
∙
Chapter 9. Molecular Structure of DNA and RNA: The ∙
Chapter 22. Biotechnology: Several Genetic TIPS have been
section on the discovery of the DNA double helix has been added to help students appreciate the uses of molecular
streamlined to focus on the key experiments. techniques in biotechnology.
∙
Chapter 10. Chromosome Organization and Molecular ∙
Chapter 23. Genomics I: Analysis of DNA: The information
Structure: The topic of bacterial chromosome structure has has been updated regarding completed genome sequences
been updated, which includes a new figure (see Figure 10.3) and other aspects of genomics.
and a discussion of microdomains. ∙
Chapter 24. Genomics II: Functional Genomics, Proteomics,
∙
Chapter 11. DNA Replication: A new figure has been added and Bioinformatics: A new subsection has been added
on the initiation of DNA replication in eukaryotes (see on the method called RNA-Seq (see Figure 24.3). The
Figure 11.20). Bioinformatics section has been reorganized with an emphasis
∙
Chapter 12. Gene Transcription and RNA Modification: The on gene prediction and homology.
information on alternative splicing has been moved to this ∙
Chapter 25. Medical Genetics and Cancer: Several
chapter. Genetic TIPS have been added to help students
∙
Chapter 13. Translation of mRNA: Several Genetic TIPS understand how mutations play a role in certain
have been added to help students understand the relationship diseases, including cancer.
between the genetic code and the synthesis of polypeptides. ∙
Chapter 26. Developmental Genetics: The information on
∙
Chapter 14. Gene Regulation in Bacteria: The information Hox genes in development, and the role of the SRY gene is
on catabolite activator protein has been updated. human sex determination, have been updated.
∙
Chapter 15. Gene Regulation in Eukaryotes I: Transcriptional ∙
Chapter 27. Population Genetics: The topic of inbreeding
and Translation Regulation: The material on eukaryotic gene has been expanded.
regulation is now divided into two chapters. Chapter 15 ∙
Chapter 28. Complex and Quantitative Traits: The topic of
focuses on transcriptional and translational regulation. the identification of QTLs is now found in its own
∙
Chapter 16. Gene Regulation in Eukaryotes II: Epigenetics: subsection.
This topic has now been expanded to an entire chapter. A ∙
Chapter 29. Evolutionary Genetics: The cladistics method
new subsection has been added on the role of epigenetics in for constructing a phylogenetic tree is compared with the
vernalization, which is the process in which some plant UPGMA method.
species require an exposure to cold in order to flower the
following spring. Also, a new section has been added on the Suggestions Welcome!
intriguing topic of paramutation.
It seems very appropriate to use the word evolution to describe the
∙
Chapter 17. Non-coding RNA: This new chapter begins
continued development of this textbook. I welcome any and all
with an overview of the general functions of non-coding
comments. The refinement of any science textbook requires input
RNAs, and then the subsequent sections explore certain
from instructors and their students. These include comments re-
topics in greater detail, such as their role in chromatin
garding writing, illustrations, supplements, factual content, and
modification, transcription, translation, protein targeting, and
topics that may need greater or less emphasis. You are invited to
genome defense (e.g., the CRISPR-Cas system).
contact me at:
∙
Chapter 18. Genetics of Viruses: The material on the
integration of phage λ has been added to this chapter, along Dr. Rob Brooker
with a brief discussion of Zika virus. Also, information on Dept. of Genetics, Cell Biology, and Development
the origin of HIV and the occurrence of HIV infection University of Minnesota
worldwide and in the US has been updated. 6-160 Jackson Hall
∙
Chapter 19. Gene Mutation and DNA Repair: The information 321 Church St.
on the mismatch repair system has been updated. Minneapolis, MN 55455
∙
Chapter 20. Recombination, Immunogenetics, and brook005@umn.edu
Transposition: Section 20.2 has been revised to focus on 612-624-3053
immunogenetics.
®
Required=Results
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McGraw-Hill Connect®
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Connect is a teaching and learning platform
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Connect Insight is Connect’s new one-
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Adaptive
THE ADAPTIVE
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SmartBook’s adaptive technology provides precise,
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xiv PREFACE
REVIEWERS Reggie Cobb, Nash Community College Thomas Peavy, California State University–
Dan Choffnes, Carthage College Sacramento
Previous editions Laurie Cotroneo, Southern Utah University Rongsun Pu, Kean University
Robert S. Dotson, Tulane University Robert Rutledge, DeSales University
Amy Abdulovic-Cui, Augusta State
Robert Hinrichsen, Indiana University of Julian Shull, Appalachian State University
University
Pennsylvania Jeffry Shultz, Louisiana Tech University
Steve Alas, California State Polytechnic
Margaret Hollingsworth, University at Ronald Wagner, Central Washington
University, Pomona
Buffalo University
Harvey Babich, Stern College for Women
Mitrick Johns, Illinois University Carey Waldburger, William Paterson
Laura Hill Bermingham, University of
Ekaterina Kaverina, East Tennessee State University
Vermont
University Gary L. Walker, Appalachian State
Hector Biliran Jr., Xavier University of
Jesse Mager, University of Massachusetts University
Louisiana
Norah R. McCabe, Washington State Jessica Wooten, The University of Findlay
Mirjana Milosevic Brockett, Georgia
University
Institute of Technology
R. Deborah Overath, Texas A&M
Madhusudan Choudhary, Sam Houston
University–Corpus Christi
State University
PA RT I I N T RO D U C T I O N
CHAPTER OUTLINE
1.1 The Molecular Expression
of Genes
1.2 The Relationship Between
Genes and Traits
1.3 Fields of Genetics
1.4 The Science of Genetics
1
CC (for “carbon copy”
or “copy cat”), the first
cloned pet. In 2002, the
cat shown here was
produced by cloning,
a procedure described
in Chapter 22.
© Corbis
OVERVIEW OF GENETICS
Hardly a week goes by without a major news story involving a Studying the human genome allows us to explore fundamen-
genetic breakthrough. The increasing pace of genetic discoveries tal details about ourselves at the molecular level. The results of the
has become staggering. The Human Genome Project is a case in Human Genome Project and the 1000 Genomes Project have shed
point. This project began in the United States in 1990, when the considerable light on basic questions, like how many genes we
National Institutes of Health and the Department of Energy have, how genes direct the activities of living cells, how species
joined forces with international partners to decipher the massive evolve, how single cells develop into complex tissues, and how
amount of information contained in our genome—the DNA defective genes cause disease. Furthermore, such understanding
found within all of our chromosomes (Figure 1.1). Remarkably, may lend itself to improvements in modern medicine by leading to
in only a decade, they determined the DNA sequence (the order better diagnoses of diseases and the development of new treat-
of the bases A, T, G, and C) of over 90% of the human genome. ments for them.
The completed sequence, published in 2003, has an accuracy The journey to unravel the mysteries within our genes has
greater than 99.99%; less than one mistake was made in every involved the invention of many new technologies. For example, re-
10,000 base pairs! searchers have developed genetic techniques to produce medicines,
In 2008, a more massive undertaking, called the 1000 Ge- such as human insulin, that would otherwise be difficult or impos-
nomes Project, was launched to establish a detailed understand- sible to make. Human insulin is synthesized in strains of Esche-
ing of human genetic variation. In this international project, richia coli bacteria that have been genetically altered by the addition
researchers set out to determine the DNA sequence of at least of genes that encode the polypeptides that form this hormone. The
1000 anonymous participants from around the globe. In 2015, bacteria are grown in a laboratory and make large amounts of hu-
the sequencing of over 2500 genomes was described in the jour- man insulin. As discussed in Chapter 22, the insulin is purified and
nal Nature. administered to many people with insulin-dependent diabetes.
1
2 C H A P T E R 1 :: OVERVIEW OF GENETICS
Chromosomes
DNA, the molecule of life
Cell
The adult human body
is composed of trillions
of cells.
C G
T A
T A
• 2 meters of DNA
G
C G
T A
T A
• Approximately 22,000
T A
genes coding for
A T
C G
proteins that perform
A T
most life functions
DNA
• Approximately 3 billion
DNA base pairs per set mRNA
of chromosomes,
containing the bases A,
T, G, and C
Amino acid
FI G U RE 1.1 The human genome. The human genome is a complete set of human chromosomes. People have two sets of chromosomes—one
set from each parent—which are found in the cell nucleus. The Human Genome Project revealed that each set of chromosomes is composed of a DNA
sequence that is approximately 3 billion nucleotide base pairs long. Estimates suggest that each set contains about 22,000 different genes that encode
proteins. As discussed later, most genes are first transcribed into mRNA and then the mRNA is used to make proteins. This figure emphasizes the DNA
found in the cell nucleus. Humans also have a small amount of DNA in their mitochondria, which has also been sequenced.
CONCEPT CHECK: How might a better understanding of our genes be used in the field of medicine?
a striking example in which scientists introduced a gene from CONCEPT CHECK: What ethical issues may be associated with human cloning?
1.1 THE MOLECULAR EXPRESSION OF GENES 3
common thread that explains the existence of life and its conti-
nuity from generation to generation. For most students, this
chapter should serve as an overview of topics they have learned
in other introductory courses such as General Biology. Even so,
it is usually helpful to see the “big picture” of genetics before
delving into the finer details that are covered in Chapters 2
through 29.
Nucleus
Each Cell Contains Many Different F I G URE 1 . 4 Molecular organization of a living cell. Cellular
Proteins That Determine Cell Structure structures are constructed from smaller building blocks. In this example,
DNA is formed from the linkage of nucleotides to produce a very long
and Function
macromolecule. The DNA associates with proteins to form a chromosome.
To a great extent, the characteristics of a cell depend on the types The chromosomes are located within a membrane-bound organelle called
of proteins that it makes. The entire collection of proteins that a the nucleus, which, along with many different types of organelles, is
cell makes at a given time is called its proteome. The range of found within a complete cell.
functions among different types of proteins is truly remarkable. photo: © Biophoto Associates/Science Source
Some proteins help determine the shape and structure of a given CONCEPT CHECK: Is DNA a small molecule, a macromolecule, or an organelle?
1.1 THE MOLECULAR EXPRESSION OF GENES 5
similar to the way that groups of letters of the alphabet represent CONCEPT CHECK: Which types of macromolecules are found in chromosomes?
words. For example, the “meaning” of the sequence of bases
ATGGGCCTTAGC differs from that of TTTAAGCTTGCC.
DNA sequences within most genes contain the information to
as a karyotype. The DNA of an average human chromosome is an
direct the order of amino acids within polypeptides according to
extraordinarily long, linear, double-stranded structure that con-
the genetic code. In the code, a three-base sequence specifies
tains well over a hundred million nucleotides. Along the immense
one particular amino acid among the 20 possible choices. One
length of a chromosome, the genetic information is parceled into
or more polypeptides form a functional protein. In this way, the
functional units known as genes. An average-sized human chro-
DNA can store the information to specify the proteins made by
mosome is expected to contain about 1000 different protein-
an organism.
encoding genes.
DNA Sequence Amino Acid Sequence
ATG GGC CTT AGC Methionine Glycine Leucine Serine The Information in DNA Is Accessed During
TTT AAG CTT GCC Phenylalanine Lysine Leucine Alanine the Process of Gene Expression
To synthesize its proteins, a cell must be able to access the informa-
In living cells, the DNA is found within large structures known as tion that is stored within its DNA. The process of using a gene se-
chromosomes. Figure 1.5 is a micrograph of the 46 chromosomes quence to affect the characteristics of cells and organisms is referred
contained in a cell from a human male; this type of image is known to as gene expression. At the molecular level, the information
6 C H A P T E R 1 :: OVERVIEW OF GENETICS
members of the same species is very common. For example, CONCEPT CHECK: Why do these two frogs look so different?
some people have brown hair and others have blond hair; some
petunias have white flowers and others have purple flowers. 2. Major alterations can also occur in the structure of a
These are examples of genetic variation. This term describes chromosome. A large segment of a chromosome can be
the differences in inherited traits among individuals within lost, rearranged, or reattached to another chromosome.
a population. 3. Variation may also occur in the total number of chromo-
In large populations that occupy a wide geographic range, somes. In some cases, an organism may inherit one too
genetic variation can be quite striking. Morphological differ- many or one too few chromosomes. In other cases, it may
ences have often led geneticists to misidentify two members of inherit an extra set of chromosomes.
the same species as belonging to separate species. As an exam-
ple, Figure 1.8 shows two dyeing poison frogs that are members Variations of sequences within genes are a common source of ge-
of the same species, Dendrobates tinctorius. They display dra- netic variation among members of the same species. In humans,
matic differences in their markings. Such contrasting forms familiar examples of variation involve genes for eye color, hair
within a single species are termed morphs. You can easily imag- texture, and skin pigmentation. Chromosome variation—a change
ine how someone might m istakenly conclude that these frogs are in chromosome structure or number (or both)—is also found, but
not members of the same species. this type of change is often detrimental. Many human genetic dis-
Changes in the nucleotide sequence of DNA underlie the orders are the result of chromosomal alterations. The most com-
genetic variation that we see among individuals. Throughout this mon example is Down syndrome, which is due to the presence of
textbook, we will routinely examine how variation in the genetic an extra chromosome (Figure 1.9a). By comparison, chromosome
material results in changes in an organism’s traits. At the molecu- variation in plants is common and often leads to plants with supe-
lar level, genetic variation can be attributed to different types of rior characteristics, such as increased resistance to disease. Plant
modifications. breeders have frequently exploited this observation. Cultivated
varieties of wheat, for example, have many more chromosomes
1. Small or large differences can occur within gene sequences. than the wild species (Figure 1.9b).
When such changes initially occur, they are called gene
mutations. Mutations result in genetic variation in which
Traits Are Governed by Genes
a gene is found in two or more alleles, as previously
described in Figure 1.7. In many cases, gene mutations and by the Environment
alter the expression or function of a protein that a gene In our discussion thus far, we have considered the role that genes
encodes. play in determining an organism’s traits. Another critical factor is
1.2 THE RELATIONSHIP BETWEEN GENES AND TRAITS 9
1 2 3 4 5 6 7 8 1 2 3 4 5 6 7 8
9 10 11 12 13 14 15 16 9 10 11 12 13 14 15 16
17 18 19 20 21 22 XX 17 18 19 20 21 22 X
FI G U RE 1.1 0 The complement of human chromosomes in somatic cells and gametes. (a) A schematic drawing of the 46 chromosomes
of a human. With the exception of the sex chromosomes, these are always found in homologous pairs. (b) The chromosomal composition of a gamete,
which contains only 23 chromosomes, one from each pair. This gamete contains an X chromosome. Half of the gametes from human males contain a
Y chromosome instead of the X chromosome.
CONCEPT CHECK: The leaf cells of a corn plant contain 20 chromosomes each. How many chromosomes are found in a gamete made by a corn plant?
Language: English
Credits: Tim Lindell, Turgut Dincer, David E. Brown, and the Online
Distributed Proofreading Team at https://www.pgdp.net
(This file was produced from images generously made
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CONTENTS
PAGE
I. Baby Biographies in General 1
II. The New-born Baby: Structure and Movements 20
III. The New-born Baby: Sensations and Consciousness 39
IV. The Earliest Developments 58
Beginnings of Emotion and Progress in Sense
V. 78
Powers
VI. Progress toward Grasping 99
She learns to grasp, and discovers the World of
VII. 118
Things
VIII. The Era of Handling Things 141
IX. The Dawn of Intelligence 161
X. Beginnings of Locomotion 182
XI. Creeping and Standing 203
Rudiments of Speech; Climbing and Progress toward
XII. 224
Walking
XIII. Walking Alone; Developing Intelligence 238
THE BIOGRAPHY OF A BABY
I
BABY BIOGRAPHIES IN GENERAL
“It is a well recognized fact in the history of science that the very
subjects which concern our dearest interests, which lie nearest our
hearts, are exactly those which are the last to submit to scientific
methods, to be reduced to scientific law. Thus it has come to pass
that while babies are born and grow up in every household, and
while the gradual unfolding of their faculties has been watched with
the keenest interest and intensest joy by intelligent and even
scientific fathers and mothers from time immemorial, yet very little
has yet been done in the scientific study of this most important of all
possible subjects—the ontogenetic evolution of the faculties of the
human mind.
“Only in the last few years has scientific attention been drawn to the
subject at all. Its transcendent importance has already enlisted many
observers, but on account of the great complexity of the phenomena,
and still more the intrinsic difficulty of their interpretation, scientific
progress has scarcely yet commenced.
“What is wanted most of all in this, as in every science, is a body of
carefully observed facts. But to be an accomplished investigator in
this field requires a rare combination of qualities. There must be a
wide intelligence combined with patience in observing and honesty in
recording. There must be also an earnest scientific spirit, a loving
sympathy with the subject of investigation, yet under watchful
restraint, lest it cloud the judgment; keenness of intuitive perception,
yet soberness of judgment in interpretation.”
I have appropriated these words of Dr. Joseph Le Conte because the
general reader is not likely to see them where they were originally
printed, in a little university study, and it is a pity to let the general
reader miss so good an introduction to the subject. Not all learned
men rate baby biography as highly as Dr. Le Conte does; but
probably all biologists do, and those psychologists who are most
strongly impressed with the evolutionary interpretation of life.
It is easy to see why one’s views of evolution affect the matter. In
botany, for instance, we do not think that we can understand the
mature plant by studying it alone, without knowledge of its
germinating period. If we omitted all study of radicle and plumule and
cotyledon, we should not only lose an interesting chapter from the
science, but even the part we kept, the classification and morphology
and physiology of the grown plant itself, would be seriously
misunderstood in some ways. So in other sciences: it is necessary to
understand how things came to be what they are, to study the
process of becoming, so to speak, before the completed result can
be understood. This is what we mean by “the genetic method” of
studying a subject.
Now, in proportion as one believes that the faculties of the human
mind unfold by evolutionary law, like a plant from the germ, he will
feel the need of studying these also genetically. As we find them in
our grown selves, they are often perplexing. What seems a single
complete, inborn faculty may really be made up of simpler ones, so
fused together by long practice that they cannot be discerned. We
know that this is the case with seeing. For instance, we give a glance
at a ball, and see its form with a single act of mind. Yet that act
became possible only after long drill in putting simpler perceptions
together. Many a test of form, turning objects over and over, passing
the hands round and round them, learning the absence of corners,
the equality of diameters, did we go through in babyhood, many an
inspection by eye, many an exercise of memory, connecting the
peculiar arrangement of light and shade with the form as felt, before
we could “see” a ball. Had this been understood in Froebel’s time, it
would have made a material difference in his suggestions as to
sense training in earliest infancy. So other powers that seem simple
and inborn may perhaps be detected in the act of forming
themselves out of simpler ones, if we watch babies closely enough,
and it may lead us to revise some of our theories about education.
There are enthusiasts, indeed, who would have us believe that child
study is going to revolutionize all our educational methods, but those
who are surest of these wonderful results, and readiest to tell
mothers and teachers what is the truly scientific thing to do with their
children, are not the ones who have done the most serious first hand
study of children. From indications so far, it is likely that the outcome
of such study will oftener be to confirm some good old-fashioned
ways of training (showing that they rested unconsciously on a sound
psychological basis) than to discover new ways. No substitute has
yet been found by scientific pedagogy for motherly good sense and
devotion.
Yet the direct study of child minds does bring out some new
suggestions of educational value, does give a verdict sometimes
between old conflicting theories, and always makes us understand
more clearly what we are doing with children. And on the purely
scientific side there is one aspect of especial interest in genetic
studies. That is, the possible light we may get on the past of the
human race.
It has long been observed that there are curious resemblances
between babies and monkeys, between boys and barbaric tribes.
Schoolboys administer law among themselves much as a tribal court
does; babies sit like monkeys, with the soles of their little feet facing
each other. Such resemblances led, long before the age of Darwin,
to the speculation that children in developing passed through stages
similar to those the race had passed through; and the speculation
has become an accepted doctrine since embryology has shown how
each individual before birth passes in successive stages through the
lower forms of life.
This series of changes in the individual is called by evolutionists the
Ontogenic Series; and the similar series through which the race has
passed in the myriads of ages of its evolution is called the
Phylogenic.
Now, of these two versions of the great world history, the phylogenic
is a worn and ancient volume, mutilated in many places, and often
illegible. The most interesting chapter of all is torn out—that which
records the passing over of man from brute to human, the beginning
of true human reason, speech, and skill. The lowest living races are
far beyond the transition line; the remains of the past can never tell
us how it was crossed, for before man could leave anything more
than bones—any products of his art, such as weapons, or signs of
fire—he had traveled a long way from his first human condition.
But from the ontogenic record no chapter can be torn out: a fresh
copy of the whole history, from alpha to omega, is written out every
time an infant is conceived, and born, and grows to manhood. And
somewhere on the way between the first cell of the embryo and
maturity each one must repeat in his own life that wonderful
transition into human intelligence. If we can thoroughly decipher tills
ontogenic record, then, what may we not hope to learn of the road by
which we human beings came?
We must not forget that the correspondence between these life
books is only a rough one. They are versions of the same world
story, but they have traveled far from their common origin, and have
become widely unlike in details. The baby has to take many short
cuts, and condense and omit inconceivably, to get through in a few
brief years a development that the race took ages for. Even the order
of development gets disarranged sometimes. For instance, primitive
man probably reached a higher development before he could talk
than babies have to now, after ages of talking ancestry: we must not
look to a child just learning to talk, to get an idea of what the minds
of men were like when they were just learning to talk. Again, the
human child is carrying on under the influence of adults an evolution
that primitive man worked out without help or hindrance from any
one wiser than himself; and that makes a great difference in the way
he does it.
The moral of all this is that people should be very cautious indeed in
drawing parallels between the child and the race, and especially in
basing educational theories on them. But if one is cautious enough
and patient enough, there are many hints about our race history to
be found in every nursery. Some of these I shall relate in the
following chapters.
Most studies of children deal with later childhood, the school years;
and these are almost always statistical in their method, taking the
individual child very little into account. My own study has been of
babyhood, and its method has been biographical. It is hard to get
statistics about babies, scattered as they are, one by one, in different
homes, not massed in schoolrooms. Now and then a doctor has
found material for good comparative investigations, and much effort
has been spent in trying to gather up measurements of babies’
growth; but on the whole the most fruitful method so far has been the
biographical one—that of watching one baby’s development, day by
day, and recording it.
I am often asked if the results one gets in this way are not
misleading, since each child might differ greatly from others. One
must, of course, use great caution in drawing general conclusions
from a single child, but in many things all babies are alike, and one
learns to perceive pretty well which are the things. Babyhood is
mainly taken up with the development of the large, general racial
powers; individual differences are less important than in later
childhood. And the biographical method of child study has the
inestimable advantage of showing the process of evolution going on,
the actual unfolding of one stage out of another, and the steps by
which the changes come about. No amount of comparative statistics
could give this. If I should find out that a thousand babies learned to
stand at an average age of forty-six weeks and two days, I should
not know as much that is important about standing, as a stage in
human progress, as I should after watching a single baby carefully
through the whole process of achieving balance on his little soles.
Yet there are not many baby biographies in existence. There are
scarcely half a dozen records that are full and consecutive enough to
be at all entitled to the name, and even of more fragmentary ones
the number in print as separate essays is scarcely larger. A good
many more, however, have been available in manuscript to students,
and many mothers no doubt keep such little notebooks. These notes
are often highly exact and intelligent, as far as they go (I have found
this especially true of the notebooks of members of the Association
of Collegiate Alumnæ), and afford important corroborations here and
there to more continuous records.
It was the Germans who first thought baby life worth recording, and
the most complete and scientific of all the records is a German one.
The first record known was published in the last century by a
Professor Tiedemann—a mere slip of an essay, long completely
forgotten, but resuscitated about the middle of this century,
translated into French (and lately into English), and used by all
students of the subject. Some of its observations we must, with our
present knowledge, set down as erroneous; but it is on the whole
exact and valuable, and a remarkable thing for a man to have done
more than a hundred years ago.
Perhaps Darwin, in 1840, was the next person to take notes of an
infant’s development; but they were taken only incidentally to
another study, and were not published for more than thirty years
(partly in “The Expression of the Emotions in Man and Animals,”
1873, partly in a magazine article in 1877). They are scanty but
important. In the interval before they were published two or three
small records had been published in Germany, and at least one
paper, that of M. Taine, in France.
In 1881, the first edition of Professor Preyer’s “model record” was
published, and before his death, in 1897, it had reached its third
edition in Germany, and had been widely circulated in America in Mr.
Brown’s excellent translation, “The Senses and the Will,” and “The
Development of the Intellect.” It did more to stimulate and direct the
study of infancy than any other publication. It has, however, the
limitations that were to be expected from Professor Preyer’s special
training as a physiologist, and is meagre on the side of mental,
moral, and emotional development. Professor Sully’s “Extracts from
a Father’s Diary,” published in part in 1881 and 1884 and fully in
1896, is richer on these sides, and also more readable.
Within the present decade, it is worth observing, the principal
records have been American, not German, and have been written by
women. Outside of America, only men, usually university professors,
have made extended records. Professor Preyer and Professor Sully
have both appealed in vain to their countrywomen to keep such
records, holding up American women for emulation. My “Notes on
the Development of a Child” were published in 1893 and 1899. In
1896 appeared Mrs. Hall’s “The First 500 Days of a Child’s Life,” a
brief record, and confined to a short period, but a very good one, and
perhaps the best for use as a guide by any one who wishes to keep
a record and finds Preyer too technical. Mrs. Moore’s “Mental
Development of a Child” is quite as much a psychological study as a
record, but is based on full biographical notes; it will be more used
by students than general readers. Mrs. Hogan’s “A Study of a Child,”
1898, is less scholarly than the others, but has a great deal of useful
material; it does not begin at birth, however, but with the fourteenth
month.
Perhaps I should say a word here as to the way in which I came to
make a baby biography, for I am often asked how one should go to
work at it. It was not done in my case for any scientific purpose, for I
did not feel competent to make observations of scientific value. But I
had for years desired an opportunity to see the wonderful unfolding
of human powers out of the limp helplessness of the new-born baby;
to watch this fascinating drama of evolution daily, minutely, and with
an effort to understand it as far as I could, for my own pleasure and
information. I scarcely know whence the suggestion had come;
probably almost by inheritance, for my mother and grandmother had
both been in somewhat notable degree observers of the
development of babies’ minds. But, unlike them, I had the notebook
habit from college and editorial days, and jotted things down as I
watched, till quite unexpectedly I found myself in possession of a
large mass of data.
A few days after my own notes began I obtained Professor Preyer’s
record, and without it I should have found the earliest weeks quite
unintelligible. For some months my notes were largely memoranda
of the likenesses and differences between my niece’s development
and that of Preyer’s boy, and I still think this is the best way for a new
observer to get started. As time went on, I departed more and more
from the lines of Preyer’s observations, and after the first year was
little influenced by them. Later, I devoted a good deal of study to the
notes, and tried to analyze their scientific results.
There is one question that I have been asked a hundred times about
baby biography: “Doesn’t it do the children some harm? Doesn’t it
make them nervous? Doesn’t it make them self-conscious?” At first
this seemed to me an odd misapprehension—as if people supposed
observing children meant doing something to them. But I have no
doubt it could be so foolishly managed as to harm the child. There
are thousands of parents who tell anecdotes about children before
their faces every day in the year, and if such a parent turns child
student it is hard to say what he may not do in the way of dissecting
a child’s mind openly, questioning the little one about himself, and
experimenting with his thoughts and feelings. But such observing is
as worthless scientifically as it is bad for the child: the whole value of
an observation is gone as soon as the phenomena observed lose
simplicity and spontaneity. It should be unnecessary to say that no
competent observer tampers with the child in any way. If Professor
Preyer, observing the baby as he first grasps at objects, notes down
the way in which he misdirects his inexpert little hands; if Mrs. Barus
keeps record of her boy’s favorite playthings; if I sit by the window
and catch with my pencil my niece’s prattle as she plays about below
—and if these babies afterward turn out spoiled, the mischief must
be credited to some other agency than the silent notebook.
Even direct experimenting on a child is not so bad as it sounds.
When you show a baby his father’s photograph to see if he
recognizes it, you are experimenting on him. The only difference
between the child student’s experimenting and that which all the
members of the family are doing all day with the baby, is that the
student knows better what he is trying to find out, and that he writes
it down.
Probably women are more skillful than men in quietly following the
course of the child’s mind, even leading him to reveal himself without
at all meddling with him or marring his simplicity. It has been so in a
marked degree in the cases I have seen. But no one who has good
judgment will allow himself to spoil both the child and his own
observation; and any one who has not good judgment will find plenty
of ways to spoil a child more potent than observing him.
II
THE NEW-BORN BABY: STRUCTURE AND
MOVEMENTS.
“Its first act is a cry, not of wrath, as Kant said, nor a shout of joy, as
Schwartz thought, but a snuffling, and then long, thin, tearless á—á,
with the timbre of a Scotch bagpipe, purely automatic, but of
discomfort. With this monotonous and dismal cry, with its red,
shriveled, parboiled skin (for the child commonly loses weight the
first few days), squinting, cross-eyed, pot-bellied, and bow-legged, it
is not strange that, if the mother has not followed Froebel’s
exhortations and come to love her child before birth, there is a brief
interval occasionally dangerous to the child before the maternal
instinct is fully aroused.”
It cannot be denied that this unflattering description is fair enough,
and our baby was no handsomer than the rest of her kind. The little
boy uncle, who had been elated to hear that his niece resembled
him, looked shocked and mortified when he saw her. Yet she did not
lack admirers. I have never noticed that women (even those who are
not mothers) mind a few little æsthetic defects, such as these that
President Hall mentions, with so many counterbalancing charms in
the little warm, soft, living thing.
Nor is it women only who find the new baby enchanting—in
Germany, at least. Semmig, whose “Tagebuch eines Vaters” is one
of the earliest attempts at a record, is delighted even with the “dismal
and monotonous cry.” “Heavenly music of the first cry!” he exclaims,
“sacred voice of life, first sound of the poem of a heart, first note of
the symphony of human life, thou echo of God’s word! What sound is
like unto thee?” “Yes, it is so: the cry of the baby is music! When it is
still, especially in the night, one is uneasy; one longs for this primitive
expression of the little being, and is consoled, enraptured, when the
helpless creature breaks into loud wails, and says to us: I live, give
me what I need! Oh, cry of the baby in the night, nightingale song for
mother and father!”
Our baby was at least a handsome one from the doctor’s point of
view, strong, healthy, and well formed; and this is to be taken into
account as a determining factor in all the record that follows.
I thought that she must be out of the normal in the matter of legs, so
oddly brief were the fat little members. Afterward I learned that all
babies are built that way—and indeed that they are altogether so
different in structure from the grown man that Dr. Oppenheim, in his
book on “The Development of the Child,” comes near to saying that
we must regard the infant as a different animal form from the adult,
almost as the caterpillar is different from the butterfly. Common
speech recognizes this in the case of several of the higher animals,
naming the young form as differently as if it were a different species.
We say a colt, a calf, a puppy, a baby; not a young horse, cow, dog,
or man.
We call a baby a little copy of the man, but really if he were
magnified to man’s size and strength, we should regard him at first
glance as an idiot and monster, with enormous head and abdomen,
short legs, and no neck, not to speak of the flat-nosed, prognathous
face; and on the other hand, a baby that was really a small copy of
man’s body would seem positively uncanny. We see this in old
pictures, where the artist tried to depict babies by placing small-sized
men and women in the mother’s arms.
The middle point of the baby’s length falls a little above the navel,
the abdomen and legs together making up a little more than half the
whole length; in the man the legs alone make a trifle more than half.
In proportion to the baby’s total weight, its brain weighs seven times
as much as a grown person’s, its muscles little more than half as
much.
“The two [man and baby] do not breathe alike, their pulse rates are
not alike, the composition of their bodies is not alike.” The baby’s
body at birth is 74.7 per cent. water, ours 58.5 per cent. It is largely
due to its loose, watery structure that the baby’s brain is so heavy—
which shows the folly of trying to compare mental powers by means
of brain weights, as is so often done in discussing woman’s sphere.
As Donaldson says, if there were anything in that basis of
comparison, the new-born baby would be the intellectual master of
us all. The baby has bright red and watery marrow, instead of the
yellow, fatty substance in our bones; and its blood differs so from
ours in proportion of red and white corpuscles and in chemical make-
up as to “amount almost to a difference in kind,” says Dr.
Oppenheim, who adds that such a condition of marrow or blood, if
found in a grown person, would be considered an indication of
disease.
The organs are differently placed within the body, and even
differently formed. The bony structure is everywhere soft and
unfinished, the plates of the skull imperfectly fitted together, with
gaps at the corners; and it is well that they are, for if the brain box
were closed tight the brain within could never grow. Surgeons have
lately even made artificial openings where the skull was prematurely
perfect, to save the baby from idiocy. The bony inclosures of the
middle ear are quite unfinished, so that on the one side catarrhal
inflammations from the nose and throat travel up to the ear more
readily than in later life, while on the other side ear inflammations are
more likely to pass into the brain. The spine is straight, like an ape’s,
instead of having the double curve of human-kind, which seems to
be brought about by the pull of the muscles after we have come to
stand erect.
I have quoted these details from Oppenheim, and from Vierordt’s
and Roberts’s measurements, as given by Dr. Burk (“Growth of
Children in Height and Weight.”) Some of the figures are given
otherwise by other authorities. I might fill many pages with similar
details. Some of these differences do not disappear till full manhood,
others are gone in a few weeks after birth. And in them all there is so
constant a repetition of lower animal forms that anatomists are
brought to a confidence in the “recapitulation doctrine,” such as they
can hardly give to others by means of a few sample facts.
The most curious of all the monkey traits shown by the new-born
baby is the one investigated by Dr. Louis Robinson (“Nineteenth