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Contents
PART I ● Patterns and Processes of Becoming:
A Framework for Understanding Animal Development
1 and Teratology 36
The Making of a Body and a Field
Introduction to Developmental Biology
“How Are You, You?” Comparative Embryology and
the Questions of Developmental Biology 2
The Cycle of Life 7
An animal’s life cycle 7
A flowering plant’s life cycle 8
Example 1: A Frog’s Life 8
Gametogenesis and fertilization 9
Cleavage and gastrulation 9
Organogenesis 11
Metamorphosis and gametogenesis 11
Example 2: Even a Weed Can Have
a Flower-Full Life 12
Reproductive and gametophytic phases 12
Embryogenesis and seed maturation 12
Vegetative phases: From sporophytic growth
to inflorescence identity 14
An Overview of Early Animal Development 14
Patterns of cleavage 14
Gastrulation: “The most important time
in your life” 17
The primary germ layers and early organs 17
Understanding cell behavior in the embryo 19
A Basic Approach to Watch Development 20
Approaching the bench: Find it, lose it, move it 20
Direct observation of living embryos 21
Dye marking 22
Genetic labeling 23
Transgenic DNA chimeras 24
Evolutionary Embryology 25
Understanding the tree of life to see
our developmental relatedness 28
The developmental history of land plants 32
Personal Significance: Medical Embryology
Genetic malformations and syndromes 36
Disruptions and teratogens 36
Coda 37
1
2
Specifying Identity
Mechanisms of Developmental Patterning 39
Levels of Commitment 40
Cell differentiation 40
Cell fate maturation 40
Autonomous Specification 41
Cytoplasmic determinants and autonomous
specification in the tunicate 42
Conditional Specification 44
Cell position matters: Conditional specification
in the sea urchin embryo 45
Syncytial Specification 48
Opposing axial gradients define position 49
3
Differential Gene Expression
Mechanisms of Cell Differentiation 55
Defining Differential Gene Expression 56
A Quick Primer on the Central Dogma 56
Evidence for Genomic Equivalence 57
Anatomy of the Gene 59
Chromatin composition 59
Exons and introns 60
Major parts of a eukaryotic gene 61
viii   
Contents
The transcription product and how it is processed 62
Noncoding regulatory elements: The on, off, and
dimmer switches of a gene 62
Mechanisms of Differential Gene Expression:
Transcription 67
Epigenetic modification: Modulating access to genes 67
Transcription factors regulate gene transcription 72
The gene regulatory network: Defining
an individual cell 78
Mechanisms of Differential Gene Expression:
Pre-messenger RNA Processing 79
Creating families of proteins through alternative
pre-mRNA splicing 80
Mechanisms of Differential Gene Expression:
mRNA Translation 82
Differential mRNA longevity 82
Stored oocyte mRNAs: Selective inhibition of
mRNA translation 83
Ribosomal selectivity: Selective activation of
mRNA translation 84
microRNAs: Specific regulation of mRNA translation
and transcription 84
Control of RNA expression by cytoplasmic
localization 87
Mechanisms of Differential Gene Expression:
Posttranslational Protein Modification 88
Coda 95
4 Meristem cells of the Arabidopsis thaliana embryo
Cell-to-Cell Communication
Mechanisms of Morphogenesis 99
A Primer on Cell-to-Cell Communication 100
Adhesion and Sorting: Juxtacrine Signaling and
the Physics of Morphogenesis 101
Differential cell affinity 101
The thermodynamic model of cell interactions 103
Cadherins and cell adhesion 103
The Extracellular Matrix as a Source of
Developmental Signals 106
Integrins: Receptors for extracellular matrix
molecules 108
The Epithelial-Mesenchymal Transition 108
Cell Signaling 109
Induction and competence 109
Paracrine Factors: Inducer Molecules 114
Morphogen gradients 114
Signal transduction cascades: The response
to inducers 116
Fibroblast growth factors and the RTK pathway 116
FGFs and the JAK-STAT pathway 118
The Hedgehog family 119
The Wnt family 123
The TGF-β superfamily 126
Other paracrine factors 127
The Cell Biology of Paracrine Signaling 132
Focal membrane protrusions as signaling sources 134
Juxtacrine Signaling for Cell Identity 137
The Notch pathway: Juxtaposed ligands and receptors
for pattern formation 137
Paracrine and juxtacrine signaling in coordination:
Vulval induction in C. elegans 138
5
Stem Cells
Their Potential and Their Niches 143
The Stem Cell Concept 144
Division and self-renewal 144
Potency defines a stem cell 145
Stem Cell Regulation 146
Pluripotent Cells in the Embryo 148
and beyond 148
Cells of the inner cell mass in the mouse embryo 151
Adult Stem Cell Niches in Animals 153
Stem cells fueling germ cell development in
the Drosophila ovary 153
Adult Neural Stem Cell Niche of the V-SVZ 155
The neural stem cell niche of the V-SVZ 155
The Adult Intestinal Stem Cell Niche 160
Clonal renewal in the crypt 161
Stem Cells Fueling the Diverse Cell Lineages
in Adult Blood 162
The hematopoietic stem cell niche 163
The Mesenchymal Stem Cell: Supporting a Variety
of Adult Tissues 165
Regulation of MSC development 166
The Human Model System to Study Development
and Disease 167
Pluripotent stem cells in the lab 167
Induced pluripotent stem cells 170
Organoids: Studying human organogenesis
in a culture dish 173

PART II ● Gametogenesis and Fertilization: The Circle of Sex
6 7
Sex Determination and Gametogenesis 179
Sex Determination 179
Chromosomal Sex Determination 180
The Mammalian Pattern of Sex Determination 180
Gonadal sex determination in mammals 182
Secondary sex determination in mammals:
Hormonal regulation of the sexual phenotype 186
Chromosomal Sex Determination in Drosophila 191
Sex determination by dosage of X 191
The Sex-lethal gene 191
Doublesex: The switch gene for sex determination 193
Environmental Sex Determination 195
Gametogenesis in Animals 196
PGCs in mammals: From genital ridge to gonads 197
Meiosis: The intertwining of life cycles 199
Spermatogenesis in mammals 202
Oogenesis in mammals 204
Sex Determination and Gametogenesis
in Angiosperm Plants 206
Sex Determination 206
Gametogenesis 209
Pollen 211
The ovule 211
PART III ● Early Development: Cleavage, Gastrulation, and Axis Formation
8 and ctenophores 248
Snails, Flowers, and Nematodes
Different Mechanisms for Similar Patterns
of Specification 247
A Reminder of the Evolutionary Context That
Built the Strategies Governing Early
Development 248
Contents   ix
Fertilization Beginning a New Organism 215
Structure of the Gametes 216
Sperm 216
The egg 218
Recognition of egg and sperm 221
External Fertilization in Sea Urchins 221
Sperm attraction: Action at a distance 221
The acrosome reaction 223
Recognition of the egg’s extracellular coat 224
Fusion of the egg and sperm cell membranes 224
Prevention of polyspermy: One egg, one sperm 224
Activation of egg metabolism in sea urchins 228
Fusion of genetic material in sea urchins 233
Internal Fertilization in Mammals 234
Getting the gametes into the oviduct:
Translocation and capacitation 234
In the vicinity of the oocyte: Hyperactivation, directed
sperm migration, and the acrosome reaction 236
Recognition at the zona pellucida 236
Gamete fusion and the prevention of polyspermy 237
Activation of the mammalian egg 239
Fusion of genetic material 239
Fertilization in Angiosperm Plants 241
Pollination and beyond: The progamic phase 241
Pollen germination and tube elongation 242
Pollen tube navigation 242
Double fertilization 243
Coda 244
The diploblastic animals: Cnidarians
The triploblastic animals: Protostomes
and deuterostomes 248
What’s to develop next 250
Early Development in Snails 250
Cleavage in Snail Embryos 251
Maternal regulation of snail cleavage 252
Axis determination in the snail embryo 259
Gastrulation in Snails 262
x   
Contents
The Nematode C. elegans 263
Cleavage and Axis Formation in C. elegans 264
Rotational cleavage of the egg 264
Anterior-posterior axis formation 265
Dorsal-ventral and right-left axis formation 266
Control of blastomere identity 267
Gastrulation of 66 Cells in C. elegans 270
9 Fate maps and the determination
The Genetics of Axis Specification
in Drosophila 273
Early Drosophila Development 275
Fertilization 275
Cleavage 276
The mid-blastula transition 277
Gastrulation 278
The Genetic Mechanisms Patterning
the Drosophila Body 281
Segmentation and the Anterior-Posterior
Body Plan 282
Maternal gradients: Polarity regulation
by oocyte cytoplasm 283
The anterior organizing center: The Bicoid and
Hunchback gradients 286
The terminal gene group 287
Summarizing early anterior-posterior axis
specification in Drosophila 288
Segmentation Genes 288
Segments and parasegments 288
The gap genes 289
The pair-rule genes 291
The segment polarity genes 292
The Homeotic Selector Genes 295
Generating the Dorsal-Ventral Axis 297
Dorsal-ventral patterning in the oocyte 297
Generating the dorsal-ventral axis
within the embryo 298
Axes and Organ Primordia: The Cartesian
Coordinate Model 299
10
Sea Urchins and Tunicates
Deuterostome Invertebrates 303
Early Development in Sea Urchins 304
Early cleavage 304
Blastula formation 306
of sea urchin blastomeres 306
Gene regulatory networks and skeletogenic mesenchyme
specification 307
Specification of the vegetal cells 310
Sea Urchin Gastrulation 311
Ingression of the skeletogenic mesenchyme 311
Invagination of the archenteron 315
Early Development in Tunicates 318
Cleavage 318
The tunicate fate map 319
Autonomous and conditional specification of tunicate
blastomeres 320
11
Amphibians and Fish 325
Early Amphibian Development 325
Fertilization, Cortical Rotation, and Cleavage 326
Unequal radial holoblastic cleavage 328
The mid-blastula transition: Preparing
for gastrulation 329
Amphibian Gastrulation 329
Epiboly of the prospective ectoderm 330
Vegetal rotation and the invagination
of the bottle cells 331
Involution at the blastopore lip 334
Convergent extension of the dorsal mesoderm 336
Progressive Determination of
the Amphibian Axes 339
Specification of the germ layers 339
The dorsal-ventral and anterior-posterior axes 340
The Work of Hans Spemann and Hilde Mangold:
Primary Embryonic Induction 340
Molecular Mechanisms of Amphibian Axis
Formation 342
How does the organizer form? 343

Functions of the organizer 348
Induction of neural ectoderm and dorsal mesoderm:
BMP inhibitors 348
Conservation of BMP signaling during
dorsal-ventral patterning 351
Regional Specificity of Neural Induction along
the Anterior-Posterior Axis 351
Specifying the Left-Right Axis 355
Early Zebrafish Development 356
Zebrafish Cleavages: Yolking Up the Process
Gastrulation and Formation of the Germ Layers
Progression of epiboly 361
Internalization of the hypoblast 362
The embryonic shield and the neural keel 363
Dorsal-Ventral Axis Formation 364
The fish blastopore lip 365
Teasing apart the powers of Nodal and BMP during axis
determination 365
Left-Right Axis Formation 367
PART IV ● Building with Ectoderm:
The Vertebrate Nervous Systyem and Epidermis
13
Neural Tube Formation and Patterning 401
Transforming the Neural Plate into a Tube:
The Birth of the Central Nervous System
Primary neurulation 404
Secondary neurulation 412
Patterning the Central Nervous System 413
The anterior-posterior axis 413
The dorsal-ventral axis 415
Opposing morphogens 416
All Axes Come Together 419
Contents   xi
12
Birds and Mammals 369
Early Development in Birds 371
Avian Cleavage 372
Gastrulation of the avian embryo 372
Axis specification and the avian “organizer” 377
358
Left-right axis formation 379
361
Early Development in Mammals 380
Mammalian cleavage 380
Trophoblast or ICM? The first decision of the rest of your
life 382
Mammalian gastrulation 383
Mammalian axis formation 388
Twins 394
Coda 396
14
Brain Growth 421
Neuroanatomy of the Developing
403 Central Nervous System 422
The cells of the developing central nervous system 422
Tissues of the developing central nervous system 422
Developmental Mechanisms
Regulating Brain Growth 427
Neural stem cell behaviors during division 427
Neurogenesis: Building from the bottom up
(or from the inside out) 428
Glia as scaffold for the layering of the cerebellum
and neocortex 430
Signaling mechanisms regulating development
of the neocortex 431
Development of the Human Brain 434
Fetal neuronal growth rate after birth 434
Hills raise the horizon for learning 435
Genes for brain growth 437
Changes in transcript quantity 438
Teenage brains: Wired and unchained 438
xii   
Contents

Local and long-range guidance molecules:

15 The street signs of the embryo 470
Repulsion patterns: Ephrins and semaphorins 470
How Did the Axon Cross the Road? 471
Neural Crest Cells and …Netrin 472
Axonal Specificity 441 Slit and Robo 473
The Travels of Retinal Ganglion Axons 475
The Neural Crest 441 Growth of the retinal ganglion axon to
Regionalization of the Neural Crest 443 the optic nerve 476
Growth of the retinal ganglion axon
Neural Crest: Multipotent Stem Cells? 444
through the optic chiasm 476
Specification of Neural Crest Cells 446
Target Selection: “Are We There Yet?” 477
Neural Crest Cell Migration: Epithelial to Chemotactic proteins 477
Mesenchymal and Beyond 447 Target selection by retinal axons:
Delamination 448 “Seeing is believing” 478
The driving force of contact inhibition 450
Synapse Formation 480
Collective migration 451
Migration Pathways of Trunk Neural Crest Cells 452
The ventral pathway 453
The dorsolateral pathway 455 16
Cranial Neural Crest 457
The “Chase and Run” Model 459
Ectodermal Placodes and
An elaborate collaboration of pushes and pulls 459 the Epidermis 485
Neural Crest-Derived Head Skeleton 461
Cranial Placodes: The Senses of Our Heads 486
Cardiac Neural Crest 462 Cranial placode induction 487
Establishing Axonal Pathways Otic-epibranchial development: A shared
in the Nervous System 464 experience 488
The Growth Cone: Driver and Engine Morphogenesis of the vertebrate eye 493
of Axon Pathfinding 465 Formation of the eye field: the beginnings
Rho, Rho, Rho your actin filaments down of the retina 494
the signaling stream 466 The lens-retina induction cascade 495

Axon Guidance 467
The Intrinsic Navigational Programming
of Motor Neurons 467
Cell adhesion: A mechanism to grab the road
The Epidermis and Its Cutaneous Appendages 498
Origin of the epidermis 498
The ectodermal appendages 499
469 Signaling pathways you can sink your teeth into 501
Ectodermal appendage stem cells 502

PART V ● Building with Mesoderm and Endoderm: Organogenesis

17
Paraxial Mesoderm
The Somites and Their Derivatives
Cell Types of the Somite 510
Establishing the Paraxial Mesoderm and
Cell Fates along the Anterior-Posterior Axis 511
Specification of the paraxial mesoderm 511
Spatiotemporal collinearity of Hox genes
determines identity along the trunk 513
507 Somitogenesis 516
Axis elongation: A caudal progenitor zone
and tissue-to-tissue forces 516
How a somite forms: The clock-wavefront model 520

Linking the clock-wavefront to Hox-mediated axial
identity and the end of somitogenesis 526
Sclerotome Development 528
Vertebrae formation 528
Tendon formation: The syndetome 532
Dermomyotome Development 534
Determination of the central dermomyotome
Determination of the myotome 535
18
Intermediate and Lateral Plate Mesoderm
Heart, Blood, and Kidneys 541
Intermediate Mesoderm: The Kidney 542
Specification of the Intermediate Mesoderm:
Pax2, Pax8, and Lim1 544
Reciprocal Interactions of Developing
Kidney Tissues 545
Mechanisms of reciprocal induction 546
Lateral Plate Mesoderm: Heart and
Circulatory System 550
Heart Development 552
A minimalist heart 552
Formation of the heart fields 552
Specification of the cardiogenic mesoderm 554
Migration of the cardiac precursor cells 555
Initial heart cell differentiation 557
Looping of the heart 557
Blood Vessel Formation 559
Vasculogenesis: The initial formation
of blood vessels 559
Angiogenesis: Sprouting of blood vessels
and remodeling of vascular beds 561
Hematopoiesis: Stem Cells and Long-Lived
Progenitor Cells 563
Sites of hematopoiesis 563
The bone marrow HSC niche 565
Coda 567
Contents   xiii
19
Development of the Tetrapod Limb 571
535 Limb Anatomy 572
The Limb Bud 572
Hox Gene Specification of Limb Skeleton Identity 574
From proximal to distal: Hox genes in the limb 574
Determining What Kind of Limb to Form
and Where to Put It 576
Specifying the limb fields 576
Induction of the early limb bud 577
Outgrowth: Generating the Proximal-Distal Axis
of the Limb 582
The apical ectodermal ridge 582
Specifying the limb mesoderm:
Determining the proximal-distal polarity 584
Turing’s model: A reaction-diffusion mechanism
of proximal-distal limb development 587
Specifying the Anterior-Posterior Axis 591
Sonic hedgehog defines a zone of polarizing activity 591
Specifying digit identity by Sonic hedgehog 591
Sonic hedgehog and FGFs: Another positive
feedback loop 594
Hox genes are part of the regulatory network specifying
digit identity 595
Generating the Dorsal-Ventral Axis 599
Cell Death and the Formation of Digits
and Joints 600
Sculpting the autopod 600
Forming the joints 601
Evolution by Altering Limb Signaling Centers 602
20
The Endoderm Tubes and Organs for
Digestion and Respiration 607
The Pharynx 609
The Digestive Tube and Its Derivatives 611
Specification of the gut tissue 612
Accessory organs: The liver, pancreas, and gallbladder 614
The Respiratory Tube 619
Epithelial-mesenchymal interactions and
the biomechanics of branching in the lungs 620
xiv   
Contents
PART VI ● Postembryonic Development
21
Metamorphosis
The Hormonal Reactivation of Development
Amphibian Metamorphosis 626
Morphological changes associated
with amphibian metamorphosis 627
Hormonal control of amphibian metamorphosis 629
Regionally specific developmental programs 631
Metamorphosis in Insects 632
Imaginal discs 633
Hormonal control of insect metamorphosis 636
The molecular biology of 20-hydroxyecdysone
activity 638
Determination of the wing imaginal discs 639
22
Regeneration
The Development of Rebuilding 643
Defining The Problem of Regeneration
Regeneration, a Recapitulation of Embryonic
Development? 645
An Evolutionary Perspective on Regeneration
Regenerative Mechanics 650
Plant Regeneration 650
A totipotent way of regenerating 650
A plant’s meri-aculous healing abilities 652
Whole Body Animal Regeneration 656
Hydra: Stem cell-mediated regeneration, orphallaxis,
and epimorphosis 656
Stem cell-mediated regeneration in flatworms 659
Tissue-Restricted Animal Regeneration 668
Salamanders: Epimorphic limb regeneration 668
Defining the cells of the regeneration blastema 668
Luring the mechanisms of regeneration
from zebrafish organs 673
Regeneration in Mammals 679
Compensatory regeneration in the mammalian liver 679
625 The spiny mouse, at the tipping point between
scar and regeneration 681
23
Development in Health and Disease
Birth Defects, Endocrine Disruptors,
and Cancer 685
The Role of Chance 686
Genetic Errors of Human Development 686
The developmental nature of human syndromes 686
Genetic and phenotypic heterogeneity 687
Teratogenesis: Environmental Assaults
on Animal Development 688
Alcohol as a teratogen 690
Retinoic acid as a teratogen 694
Endocrine Disruptors: The Embryonic Origins
of Adult Disease 695
644
Diethylstilbestrol (DES) 697
Bisphenol A (BPA) 698
Atrazine: Endocrine disruption through
647 hormone synthesis 700
Fracking: A potential new source of
endocrine disruption 701
Transgenerational Inheritance of Developmental
Disorders 702
Cancer as a Disease of Development 703
Development-based therapies for cancer 708
Coda 709
 Contents  xv

PART VII ● Development in Wider Contexts

Mechanisms of Evolutionary Change 742
24 Heterotopy 742
Heterochrony 744
Heterometry 745
Development and the Environment Heterotypy 746
Biotic, Abiotic, and Symbiotic Regulation Developmental Constraints on Evolution 747
of Development 711 Physical constraints 748
Morphogenetic constraints 748
Developmental Plasticity: The Environment as Pleiotropic constraints and redundancy 748
an Agent in Producing Normal Phenotypes 711
Diet-induced polyphenisms 713
Ecological Evolutionary Developmental
Biology 749
Predator-induced polyphenisms 715
Temperature as an environmental agent 716 Plasticity-First Evolution 749
Reaction norms in plants 718 Genetic assimilation in the laboratory 750
Larval settlement 720 Genetic assimilation in natural environments 751
Stress as an agent: The hard life of Selectable Epigenetic Variation 752
spadefoot toads 720
Evolution and Developmental Symbiosis 754
Developmental Symbioses 721 The evolution of multicellularity 755
Developmental symbioses in plants 722 The evolution of placental mammals 756
Mechanisms of developmental symbiosis: Getting the
Coda 756
partners together 724
Developmental symbiosis in the mammalian
intestine 727 Appendix A-1
Coda 731
Glossary G-1

25 Index I-1

Development and Evolution

Developmental Mechanisms of Evolutionary
Change 735

The Developmental Genetic Model of

Evolutionary Change 756
Preconditions for Evolution:
The Developmental Structure of the
Genome 736
Modularity: Divergence through dissociation 736
Molecular parsimony: Gene duplication
and divergence 739
Preface: Thinking Grandly
about Developmental Biology

With biology going into smaller and smaller realms, it is sometimes good to
contemplate the grand scheme of things rather than the details, to “seat thyself
sultanically among the moons of Saturn” (in Herman Melville’s phrase). It is
good, for instance, to get a perspective of developmental biology from outside
the discipline rather than from inside it.

Remembering the Field’s Interdisciplinary Foundations

Developmental biology, history tells us, is an interdisciplinary field that is at the
foundations of biology. Indeed, before the word biology came to be used, the liv-
ing world was characterized as that part of the world that was developing. The
organizers of the first meeting (in 1939) of the Growth Society, which was the
precursor of the Society for Developmental Biology, claimed that development
must be studied by combining the insights of numerous disciplines, including ge-
netics, endocrinology, biochemistry, physiology, embryology, cytology, biophysics,
mathematics, and even philosophy. Developmental biology was to be more than
embryology. It also included stem cells, which were known to generate the adult
blood, and regeneration, which was seen to be the re-activation of developmental
processes and which was critical for healing in vertebrates and for reproduction of
hydra, flatworms, and numerous other invertebrates. The first articles published
in the journal Developmental Biology showcased embryology, regeneration, and
stem cells, and the different ways of studying them.
Throughout this new 12th edition you will see a return to some of these found-
ing ideas of interdisciplinary developmental biology, namely regeneration, mor-
phomechanics, plants, and the genetic control of development.
Indeed, regeneration has historically been a major part of developmental biol-
ogy, for it is a developmental phenomenon that can be readily studied. Experimen-
tal biology was born in the efforts of eighteenth-century naturalists to document
regeneration and to examine how it was possible. The regeneration experiments of
Tremblay (hydras), Réaumur (crustaceans), and Spallanzani (salamanders) set the
standard for experimental biology and for the intelligent discussion of one’s data.
More than two centuries later, we are beginning to find answers to the great
problems of both embryology and regeneration. Indeed, the conclusions of one
support the research of the other. We may soon be able to alter the human body
so as to permit our own limbs, nerves, and organs to regenerate. Severed limbs
could be restored, diseased organs could be removed and regrown, and nerve
cells altered by age, disease, or trauma could once again function normally. The
ethical issues this would exacerbate are only beginning to be appreciated. But if
we are to have such abilities, we first have to understand how regeneration oc-
curs in those species that have this ability. Our new knowledge of the roles that
paracrine factors and physical factors play in embryonic organ formation, plus
recent studies of stem cells and their niches, has propelled what Susan Bryant has
called “a regeneration renaissance.” Since “renaissance” literally means “rebirth,”
and since regeneration can be seen as a return to the embryonic state, the term
is apt in many ways.
xviii   
Preface

Notice that biophysics was also an early part of the mix of developmental biol-
ogy. This area, too, is having a renaissance. The physical connections between
cells, the strength of their bonding, and the tensile strength of the material sub-
strates of the cells are all seen to be critical for normal development. Physical
forces are necessary for sperm-egg binding, gastrulation, heart development,
gut development, the branching of the kidney and lung epithelia, and even the
development of tumors. Physical forces can direct the development of stem cells
toward particular fates, and they can determine which part of the body is left
and which is right. The patella of our kneecap doesn’t form until we put pres-
sure on it by walking. In many cases, physical forces can direct gene expression.
Lev Beloussov, a pioneer in this area, has called this the “morphomechanics of
development.”
Another area that was prominently represented in the early programs of devel-
opmental biology was plant development. Plant development had much in common
with regeneration, as “adult” plants could redevelop entire parts of their bodies.
Whereas in animal biology the study of development diverged from the study
of physiology, that separation was not evident in plant biology. Moreover, while
many animals quickly set aside a germline that was to become the sperm or eggs,
this was not the case in plants. Such comparisons between plants and animals are
now present throughout this text, and they serve to highlight the fundamental
developmental processes that are present across phyla and even kingdoms of life.
But the genes remain the center of focus in developmental biology. And the
more we learn about them, the more interesting and complex these genes be-
come. New advances in “single cell transcriptomics” have given us an amazing
privilege—the ability to look at the gene expression patterns of individual cells
as they develop. An individual’s cells may all have the same genes, but their dif-
ferent positions in the embryo cause different genes to be active in each cell. It’s a
symphony of relationships, each cell providing the context for another. If develop-
ment is the performance, then the genome is the script or score. As anyone who
has gone to concerts knows, different bands perform the same score differently,
and the same band will play the same song differently on two successive nights.
Environment is also critical—hence, the new interest in plasticity and symbiosis
in development.
Developmental biology has also taken on a new role in science. More than
any other biological science, it demonstrates the critical importance of processes
as opposed to entities. In many organisms, the same process can be done by
different molecules. “It’s the song, not the singer,” say Doolittle and Booth, and
we can be thankful that there are redundant pathways in development—if one
pathway fails, another is often able to take over its function. The entity/process
split in developmental biology mirrors the particle/wave dichotomy in physics.
It is a “both, and” situation, rather than an “either/or” situation. In 1908, the
Scottish physiologist J. S. Haldane said, “That a meeting point between biology
and physical science may at some time be found, there is no doubting. But we
may confidently predict that if that meeting-point is found, and one of the two
sciences is swallowed up, that one will not be biology.” Developmental biology
may well solve the longstanding mysteries of physics.

New to the Twelfth Edition

In this current volume, we have attempted to track this amazing fulfillment of
the early promises of developmental biology. To this end, the book has undergone
its own morphogenesis.

Plant development covered throughout

We have now incorporated plant material into the relevant chapters. Instead of
segregating plant developmental biology into a single (and often unassigned)
 Preface   xix

chapter, we have integrated essential plant biology into the chapters on cell speci-
fication, gene regulation, cell communication, gamete production, fertilization,
axis determination, organ formation, and regeneration.

Upgraded and expanded chapter on regeneration

We have also expanded the chapter on regeneration, which we are proud to say
offers a unique summary of the field. It both captures the fascinating problems
of post-embryonic development that regeneration seems to solve and provides
a logical framework for the known mechanisms of regeneration, based on an
organism’s degree of regenerative capacity. We feel that this chapter will be an
excellent place for anyone interested in this area to start.

Updates throughout all chapters

All of the chapters have received important updates, from the introductory chap-
ter’s broader evolutionary perspective to new material on the morphomechanics
of development during Drosophila gastrulation and the formation of mammalian
lungs. Special consideration was also given to the increasing use of whole-ge-
nome, transcriptomic approaches, which are dramatically shaping our under-
standing of cell differentiation.

A new, student-centered approach

From a pedagogical standpoint, it is also good to get an outside perspective of
how students are learning developmental biology—the perspective of the student
experience. For decades, it has been the responsibility of textbooks like ours to be
the most comprehensive sources for the field’s foundational content. Although
this responsibility still remains, the reality is that students are inundated with an
overwhelming myriad of sources vying for their attention. If there was ever a time
a student of developmental biology needed a guidebook to navigate through this
dense and diverse ecosystem of texts, online resources, and infinitely expand-
ing scientific literature, the time is now and the guidebook this new volume of
Developmental Biology.

• Focused and streamlined coverage. Over the years, as new knowledge has
grown, so has our own textbook, which was reaching a size that might
itself trigger student overload and defeat the purposes of engagement
and deep learning. The information bombarding students is not going
away; therefore, they need not only access to the information but also a
clear guide that fosters movement from the essential ideas to the com-
plex mechanisms and finally to inclusive invitations that welcome their
research in this field. We have both reduced and reorganized the content
in each chapter to achieve a clear and supportive lattice so that both the
professor and the student can more easily navigate the increasing vol-
ume and complexity of developmental biology.
• Innovative pedagogy: Empowering students to craft their own learning. The
first material students will encounter in each section of a chapter rep-
resents the most essential content. We have introduced a new element
called “Further Development,” which highlights content we feel repre-
sents some of the more complex ideas in the field. In addition, students
will also come across invitations to view some Further Developments
online. These online topics represent fantastic opportunities for students
to further develop their understanding of developmental biology along
paths of their own interest—paths of investigation that professors can
have confidence match the standards of quality seen throughout the
textbook (unlike some other online sources). The special in-text fea-
tures of previous editions—Dev Tutorials, Developing Questions, Next
Step Investigations, and citations throughout—are still in place to play
xx   
Preface

important roles in empowering students to take that final leap to engage

with the developmental biology literature. To better support students’
use of the research literature, we now include a new Appendix focused
on how to find and analyze research articles in developmental biology.

Thanks to this new organization of content, professors and students will now
be in complete control of what level of material may be most appropriate. We are
proud to introduce Developmental Biology 12e, as it still provides direct access to
all levels of the content but without diluting its quality and the overall learning
experience.

Acknowledgments
First, the two authors gratefully acknowledge their mutual respect for one an-
other and for the enjoyment of each other’s work. Michael wants the community
to know that Scott has been most accepting and welcoming to new ideas and
that his enthusiasm for producing the best product has not wavered any day
of any edition. Scott wants the community to know that he is thrilled with the
new ideas that Michael has brought to the book and that Michael’s commitment
to undergraduate education is second to none.
Second, we are thrilled to acknowledge the importance of Mary Stott Tyler to
this book. The winner of the Viktor Hamburger Education Award and the author
of Fly Cycle, Differential Expressions, The Developmental Biology Vade Mecum, and
Inquiry Biology, Mary has been a mixture of author, editor, and curator of contents
for this 12th edition, helping us decide “what to leave in/what to leave out.” As we
added plant studies to the book and had to remove other studies, Mary’s insight
and vision for the finished book was essential.
If science is like a balloon expanding into the unknown—and the larger the
balloon, the more points in contact with the unknown—then developmental
biology has contacted an astounding number of unknowns. The accuracy and
coverage of the 12th edition owes much to the work of the many expert reviewers
who took the time to provide respectful formal and informal feedback throughout
the process (see list). The organization of these reviews was consistently executed
by Lauren Cahillane, Nina Rodriguez-Marty and Katie Tunkavige—thank you
for making this important part possible. This 12th edition is particularly unique
as it marks the new incorporation of plant developmental biology. There were
numerous reviewers who offered their expertise in select chapters, thank you to
all. Special thanks, however, go to Anna Edlund and Marta Laskowski for their
reviews of the plant content. They were very patient with us, and any misunder-
standings are those of the authors.
This edition also marks a dramatic change to the publishing of Developmental
Biology. With the retirement of Andy Sinauer, Sinauer Associates has become an
imprint of Oxford University Press. Our book overlaps these two periods, and has
seen the change of managers, art directors, and our long-time editor. We thank
both Sinauer Associates and Oxford University Press for their great efforts in
sustaining the book during this period of metamorphosis. We wish to especially
thank Dean Scudder for taking on the managerial tasks and allowing us to work
on new models of science education during this transition. Moreover, half-way
through production of this edition, Jason Noe of Oxford became our overseeing
editor. Such a transition and short timeline for production might rattle the best of
editors, but Jason helped to establish the best adaptable plans to keep things on
track. Sincere thanks for your efforts, Jason. Meanwhile, in the house of Sinauer,
production editors Laura Green and Kathaleen Emerson shared their expertise
and their truly collaborative insights, offering us respectful considerations during
key times that we will not forget. Thank you Laura for also sharing with us your
most valuable plant background throughout the editorial process.
 Preface   xxi

The success of this and each edition equally rests on the quality of the book’s
design and look, for which we sincerely appreciate the wonderful work Sinauer’s
art, media, and overall production team have done. The media team was headed by
Suzanne Carter and supported by the creative drive of Peter Lacey. Sincere thanks
to you both. Further thanks to the entire group at Dragonfly Media, who continue
to do a great job taking care to represent many of Michael’s original drawings
with supreme accuracy. We’d also like to thank Joan Gemme, Beth Roberge, and
Annette Rapier for their excellent design, layout, and production of this edition.
One of the long-loved hallmarks of Developmental Biology has been the incor-
poration of actual data and images that represent the science. Special thanks to
the permissions team, Mark Siddall, Tracy Marton, and Michele Beckta for their
non-stop efforts in securing the rights to these essential pieces of the book. But
of course, a new book can only reach the hands of the students with the help of
a robust and strategic sales team. Many thanks to Susan McGlew and to all the
salespersons at Oxford now helping to support this textbook.
Lastly, it needs to be acknowledged that while Scott is blissfully retired,
Michael is still working his tail off doing teaching, research, committee assign-
ments, and so forth, in addition to his strong family commitments. He would
not be able to provide the time and energy to this textbook if he did not have
the support of his own institution and students. Thank you, Smith College, for
continuing to allow Michael to produce and disseminate his Web Conferences,
Developmental Documentaries, and the Dev Tutorials freely to the community.
Most sincere thanks to Michael’s research students, who had to endure their
principle investigator being too engrossed in all things development all the time!
Know that your patience, support, and insights surely made this book possible.

­— M.J.F.B.
—S.F.G.
May 24, 2019
Reviewers of the Twelfth Edition
Anna Allen, Howard University
William Anderson, Harvard University
Nicola Barber, University of Oregon
Madelaine Bartlett, University of Massachusetts, Amherst
Marianne Bronner, California Institute of Technology
Timothy Brush, University of Texas, Rio Grande Valley
Blanche Capel, Duke University
Jacqueline Connour, Ohio Northern University
D. Cornelison, University of Missouri, Columbia
Dr. Angus Davidson, The University of Nottingham
Anna Edlund, Bethany College
Elizabeth D. Eldon, California State University, Long Beach
Deborah Marie Garrity, Colorado State University
Bob Goldstein, University of North Carolina
Eric Guisbert, Florida Institute of Technology
Jeff Hardin, University of Wisconsin, Madison
Richard Harland, University of California Berkeley
Marcus Heisler, The University of Sydney
Arnold G Hyndman, Rutgers University
Zhi-Chun Lai, Pennsylvania State University
Michael Lehmann, University of Arkansas
Michael Levin, Tufts University
Yuanyuan Rose Li, University of Alabama at Birmingham
Barbara Mania-Farnell, Purdue University Northwest
Adam C. Martin, Massachusetts Institute of Technology
David Matus, Stony Brook University
Roberto Mayor, University College London
Dave McClay, Duke University
Claus Nielsen, University of Copenhagen
Fred Nijhout, Duke University
Lee Niswander, University of Colorado, Boulder
Julia Oxford, Boise State University
Mark Peifer, University of North Carolina
Isabelle Peter, California Institute of Technology
Ann Rougvie, University of Minnesota
Sabrina Sabatini, Sapienza University of Rome
Thomas F. Schilling, University of California, Irvine
Nick Sokol, Indiana University
Richard Paul Sorrentino, Auburn University
Ana Soto, Tufts University
David Stachura, California State University, Chico
Claudio Stern, University College London
Andrea Streit, King’s College London
Keiko Sugimoto, RIKEN
Jonathan Sylvester, Georgia State University
Daniel E Wagner, Harvard Medical School
Zhu Wang, University of California, Santa Cruz
Paul M. Wassarman, Icahn School of Medicine at Mount Sinai
Daniel Weinstein, Queens College, CUNY
Jessica LaMae Whited, Harvard University
Jeanne Wilson-Rawls, Arizona State University
Colleen Winters, Towson University
Tracy Young-Pearse, Harvard Medical School
Media and Supplements
to accompany Developmental Biology, Twelfth Edition
For the Student
Companion Website
devbio.com
Significantly enhanced for the Twelfth Edition, and refer-
enced throughout the textbook, the Developmental Biology
Companion Website provides students with a range of en-
gaging resources to help them learn the material presented
in the textbook. The companion site is available free of
charge and includes resources in the following categories:
• Dev Tutorials: Professionally produced video
tutorials, presented by the textbook’s authors,
reinforce key concepts.
• Watch Development: Putting concepts into
action, these informative videos show real-life
developmental biology processes.
• Further Development: These extensive topics
provide more information for advanced students,
historical, philosophical, and ethical perspectives
on issues in developmental biology, and links to
additional online resources.
• Scientists Speak: In these lectures and question-
and-answer interviews, developmental biology
topics are explored by leading experts in the field.
• Flashcards: Per-chapter flashcard sets help stu-
dents learn and review the many new terms and
definitions introduced in the textbook.
• Literature Cited: Full citations are provided for
all of the literature cited in the textbook (most
linked to their PubMed citations).
• Research Guide: This illustrated and annotated
guide helps students find and comprehend
research articles in developmental biology.
For the Instructor
(Available to qualified adopters)
Instructor’s Resource Library
The Developmental Biology, Twelfth Edition Instructor’s
Resource Library includes the following resources:
• Case Studies in Dev Bio: This collection of case
study problems provides instructors with ready-
to-use in-class active learning exercises. The case
studies foster deep learning in developmental
biology by providing students an opportunity
to apply course content to the critical analysis of
data, to generate hypotheses, and to solve novel
problems in the field. Each case study includes a
PowerPoint presentation and a student handout
with accompanying questions.
• Developing Questions: Thought-provoking
questions, many with answers, references, and
recommendations for further reading, are pro-
vided so that you and your students can explore
questions that are posed throughout each chapter.
• Textbook Figures & Tables: All of the textbook’s
figures, photos, and tables are provided both in
JPEG and PowerPoint formats. All images have
been optimized for excellent legibility when pro-
jected in the classroom.
Value Options
eBook
(ISBN 978-1-60535-823-9)
Developmental Biology, Twelfth Edition is available as
an eBook, via several different eBook providers, including
RedShelf and VitalSource. Please visit the Oxford Univer-
sity Press website at oup.com/ushe for more information.
Looseleaf Textbook
(ISBN 978-1-60535-824-6)
Developmental Biology, Twelfth Edition is also available in
a three-hole punched, looseleaf format. Students can take
just the sections they need to class and can easily integrate
instructor material with the text.
Another random document with
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third millennium as a political deity, the war-god, and leader of the
people, as real a personality as Hammurabi himself. The same is
true of Asshur, once the local deity of the aboriginal land of the
Assyrians, but later raised by the imperial expansion of this people
almost to the position of a universal god, the guardian of the land,
the teacher and the father of the kings; nor can we discern that he
was ever an elemental god.
Speaking generally, in spite of many important differences, we
may regard the religious structure to which the cults of Anatolia and
Egypt belonged as morphologically the same as that which I am
defining as Hellenic. Also, among all these peoples, by the side of
the few higher deities who have developed moral personalities, we
find special elemental divinities, as, in Hellas, we find Helios and the
deities of the wind, Hephaistos the fire-god.
The distinction between the religions of the Hellenes and “the
barbarians,” which Aristophanes defines as the difference between
the worship of ideal divine personages, such as Zeus, Apollo, and
Demeter, and the direct worship of elementary powers, such as sun
and moon, is not borne out by modern research. Where we find sun-
worship or moon-worship in the East, it does not appear to have
been directed immediately to the thing itself regarded as a living or
animate body, but to a personal god of the sun or the moon—Bel,
Shamash, or Sin. We can only distinguish the Greek from the
Oriental in respect of Nature-religion by the lesser degree of
devotion that the Hellene showed to it. Only those of his divinities
whose names connoted nothing in the material or natural world,
could develop into free moral personalities, and dominate the
religious imagination of the people. Nowhere, for instance, had
Helios any high position in the Greek world except at Rhodes, where
we must reckon with pre-Hellenic, Minoan, and later with Semitic
influences. Therefore, when, shortly before and after the beginning
of the Graeco-Roman period, a wave of sun-worship welled from the
East over the West, it may have brought with it religious ideas of
high spirituality and ethical purity, yet by the race-consciousness of
the Hellenes it must have been judged to be a regress towards a
barbaric past.
 The instinct of the Greek in his creation of divine forms shows
always a bias towards the personal and the individual, an aversion to
the amorphous and vague, and herein we may contrast him with the
Persian and Egyptian. A certain minor phenomenon in these
religions will illustrate and attest this. All of them admitted by the side
of the high personal deities certain subordinate personages less
sharply conceived, divine emanations, as we may sometimes call
them, or personifications of moral or abstract ideas. Plutarch
specially mentions the Persian worship of Truth, Goodwill, Law-
abidingness, Wisdom, emanations of Ahura-Mazda, which in the
light of the sacred books we may, perhaps, interpret as the Fravashis
or Soul-powers of the High God; and in certain Egyptian myths and
religious records we hear of a personification of Truth, whose statue
is described by the same writer. But at least in the Persian system
we may suspect that such divine beings had little concrete
personality, but, rather, were conceived vaguely as daimoniac forces,
special activities of divine force in the invisible world. Now the Greek
of the period when we really know him seems to have been mentally
unable to allow his consciousness of these things or these forces to
remain just at that point. Once, no doubt, it was after this fashion that
his ancestors dimly imagined Eros, or the half-personal Curse-power
Ἀρά; but he himself could only cherish Eros under the finished and
concrete form of a beautiful personal god, and the curse was only
vitalised for him when it took on the form of the personal Erinys. This
topic is a fruitful one, and I hope to develop it on a later occasion.
It suggests what is now the next matter I wish to touch on—the
comparison of the Mediterranean religions in respect of their
anthropomorphism. Philosophically, the term might be censured as
failing to distinguish any special type of religion; for we should all
admit that man can only envisage the unseen world in forms
intelligible to his own mind and reflecting his own mental structure.
But, apart from this truism, we find that religions differ essentially and
vitally according as this anthropomorphism is vague and indefinite or
sharply defined and dominating; according as they picture the
divinity as the exact though idealised counterpart of man, and
construct the divine society purely on the lines of the human, or
refrain from doing this either through weakness and obscurity of
imagination or in deference to a different and perhaps more elevated
law of the religious intellect. Now, of the Hellenic religion no feature
is salient as its anthropomorphism, and throughout its whole
development and career the anthropomorphic principle has been
more dominating and imperious than it has ever been found to be in
other religions.11.1 At what remote period in the evolution of the
Hellenic mind this principle began in force, what were the influences
that fostered and strengthened it, in what various ways it shaped the
religious history of the Hellenic people, are questions that I may be
able to treat more in detail in the future. But there are two important
phenomena that I will indicate now, which we must associate with it,
and which afford us an illuminating point of view from which we may
contrast the Greek world and the Oriental. In the first place, the
anthropomorphic principle, combining with an artistic faculty the
highest that the world has known, produced in Greece a unique form
of idolatry; and, in the second place, in consequence chiefly of this
idolatry, the purely Hellenic religion remained almost incapable of
that which we call mysticism.
Now, much remains still to be thought out, especially for those
interested in Mediterranean culture, concerning the influence of
idolatry on religion; and not only the history, but the psychology of
religion, must note and estimate the influence of religious art. It may
well be that the primitive Greeks, like the primitive Roman, the early
Teuton, and Indo-Iranian stocks, were non-idolatrous, and this
appears to have been true to some extent of the Minoan culture.
Nevertheless, the Mediterranean area has from time immemorial
been the centre of the fabric and the worship of the eikon and the
idol. The impulse may have come from the East or from Egypt to the
Hellene; he in his turn imparted it to the Indian Aryans, as we now
know, and in great measure at least to the Roman, just as the
Assyrian-Babylonian temple-worship imparted it to the Persian.
Nowhere, we may well believe, has the influence of idolatry been so
strong upon the religious temperament as it was upon that of the
Hellenes; for to it they owed works of the type that may be called the
human-divine, which surpass any other art-achievement of man.
I can here only indicate briefly its main effects. It intensified the
perception of the real personal god as a material fact. It increased
polytheism by multiplying the separate figures of worship, often,
perhaps, without intention. It assisted the imagination to discard what
was uncouth and terrifying in the Hellenic religion, and was at once
the effect and the cause of the attachment of the Hellenic mind
towards mild and gracious types of godhead. The aniconic emblem
and uncouth fetich-formed figures were here and there retained,
because of vague ideas about luck or for superstitious fetichistic
reasons; but the beautiful idol was cherished because it could
arouse the enthusiastic affection of a sensitive people, and could
bring them to the very presence of a friendly divine person. The
saying that the Olympian deities died of their own loveliness means
a wrong interpretation of the facts and the people. But for a beautiful
idolatry, Hellenic polytheism would have passed away some
centuries before it did, the deities fading into alien types or becoming
fused one with the other. Nor was its force and influence exhausted
by the introduction of Christianity, for it shaped the destinies of the
Greek Church, and threw down a victorious challenge to the
iconoclastic Emperors.
If now we were to look across the Mediterranean, and could
survey the religious monuments of Persia, Assyria and Babylonia,
Phoenicia, and the Hittite people, we should find a general
acceptance of the anthropomorphic idea. The high personal deities
are represented mainly in human form, but the art is not able to
interpret the polytheistic beliefs with skilfully differentiated types. In
Chaldaic and Assyrian art one type of countenance is used for
various divinities, and this such as might inspire awe rather than
affection. And the anthropomorphism is unstable. Often animal traits
appear in parts of the divine figure. Nergal has a lion’s head; even
the warrior Marduk is invoked in the mystic incantations as “Black
Bull of the Deep, Lion of the dark house.”14.1 In fact, over a large part
of anterior Asia, anthropomorphism and theriomorphism exist side by
side in religious concept and religious art. We may say the same of
Egypt, but here theriomorphism is the dominating factor.
As regards the explanation of this phenomenon, many questions
are involved which are outside my present province. I would only
express my growing conviction that these two distinct modes of
representing the divine personage to the worshipper are not
necessarily prior and posterior, the one to the other, in the evolution
of religion. They can easily, and frequently do, coexist. The vaguely
conceived deity shifts his shape, and the same people may imagine
him mainly as a glorified man of human volition and action, and yet
think of him as temporarily incarnate in an animal, and embody his
type for purposes of worship or religious art in animal forms.
I would further indicate here what I cannot prove in detail—that
theriomorphism lends itself to mysticism, while the anthropomorphic
idolatry of Greece was strongly in opposition to it. The mystic
theosophy that pervaded later paganism, and from which early
Christianity could not escape, originated, as Reitzenstein has well
shown, mainly in Egypt, and it arose partly, I think, in connection with
the hieratic and allegorical interpretation of the theriomorphic idol.
There was nothing mystic about the Zeus of Pheidias, so far as the
form of the god was concerned. The forms were entirely adequate to
the expression of the physical, moral, and spiritual nature of the god.
The god was just that, and there was nothing behind, and, as the
ancient enthusiast avers, “having once seen him thus, you could not
imagine him otherwise.” But when a divinity to whom high religious
conceptions have already come to attach is presented, as it might be
in Egyptian religious art, with the head of a jackal or an ape, the
feeling is certain to arise sooner or later in the mind of the
worshipper that the sense-form is inadequate to the idea. Then his
troubled questioning will receive a mystic answer, and the animal
type of godhead will be given an esoteric interpretation.
Plutarch, in the De Iside et Osiride,15.1 is one of our witnesses. He
finds a profounder significance for theosophy in the beetle, the asp,
and the weasel than in the most beautiful anthropomorphic work of
bronze or marble. He here turns his back on his ancestors, and goes
over to the sect of the Egyptian mystic.
But the most curious testimony to my thesis is borne by an
inscription on an Egyptian lamp—an invocation of the God Thoth: “O
Father of Light, O Word (λόγος) that orderest day and night, come
show thyself to me. O God of Gods, in thy ape-form enter.”15.2 Here
the association of so mystic a concept as the “Logos,” the divine
Reason, an emanation of God with the form of an ape, is striking
enough, and suggests to us many reflections on the contrast
between the Egyptian theriomorphism and the human idolatry of the
Greek. The Hermes of Praxiteles was too stubborn a fact before the
people’s eyes to fade or to soar into the high vagueness of the
“Logos,” too stable in his beautiful humanity to sink into the ape.
But before leaving this subject I would point out a phenomenon in
the Hellenic world that shows the working of the same principle. The
Orphic god Dionysos-Sabazios-Zagreus was πολύμορφος, a shape-
shifter, conceived now as bull, now as serpent, now as man, and the
Orphic sects were penetrated with a mystic theosophy; and, again,
they were a foreign element embedded in Greek society and religion.
While we were dealing with the subject of anthropomorphism, we
should consider also the question of sex, for a religion that gives
predominance to the god is certain to differ in some essential
respects from one in which a goddess is supreme. Now, although the
conception of an All-Father was a recognised belief in every Greek
community, and theoretically Zeus was admitted to be the highest
god, yet we may believe Athena counted more than he for the
Athenians, and Hera more for the Argives. And we have evidence of
the passionate devotion of many urban and village communities to
the mother Demeter and her daughter Kore, to whom the greatest
mysteries of Greece, full of the promise of posthumous salvation,
were consecrated. Also, in the adjacent lands of earlier culture we
mark the same phenomenon. In Egyptian religion we have the
commanding figure of Isis, who, though by no means supreme in the
earlier period, seems to dominate the latter age of this polytheism. In
the Assyrian-Babylonian Pantheon, though the male deity is at the
head, Ishtar appears as his compeer, or as inferior only to Asshur.
Coming westward towards Asia Minor, we seem to see the goddess
overshadowing the god. On the great Hittite monument at Boghaz-
Keui, in Cappadocia, skilfully interpreted by Dr. Frazer, we observe a
great goddess with her son coequal with the Father-God. In the
lands adjacent to the coast a Mother-Goddess, sometimes also
imagined as virgin, Kybele of Phrygia, Ma of Cappadocia, Hipta of
Lydia, Astarte of Askalon, Artemis of Ephesos who was probably a
blend of Hellenic and Oriental cult-ideas, appears to have been
dominant from an immemorial antiquity; and Sir Arthur Evans has
discovered the same mysterious feminine power pre-eminent in the
Minoan religion. We may even affirm that she has ruled a great part
of the Mediterranean down to the present day.
The various questions suggested by this predominance of
goddess-worship are fascinating and subtle. The sociological one—
how far it is to be connected with a system of counting descent
through the female, with a matrilinear society—I have partly
discussed elsewhere.17.1 I may later be able to enter on the question
that is of more interest for the psychology of religion—the effect of
such worships on the religious sentiment. Here I can merely point to
the phenomenon as a natural and logical product of the principle of
anthropomorphism, but would call attention to the fact that in the
East it sometimes developed into a form that, from the
anthropomorphic point of view, must be called morbid and
subversive of this principle; for the rivalry of divine sex was here and
there solved by the fusion of the two natures in the divinity, and we
find a bisexual type—a male Astarte, a bearded Ishtar.17.2 The
healthy-minded anthropomorphism of the Hellene rejects this
Oriental extravagance.
If we now could consider in detail the various moral conceptions
attached to the high State divinities of Greece and the East, we
should be struck with a general similarity in the point of view of the
various culture-stocks. The higher deities, on the whole, are ethical
beings who favour the righteous and punish transgressors; and the
worship of Greece falls here into line with the Hebraic conceptions of
a god of righteousness. But in one important particular Hellenic
thought markedly differs from Oriental, especially the Persian. In the
people’s religion throughout Hellas the deities are, on the whole,
worshipped as beneficent, as doing good to their worshippers, so
long as these do not offend or sin against them. The apparent
exceptions are no real exceptions. Ares may have been regarded as
an evil god by the poet or the philosopher, but we cannot discover
that this was ever the view of the people who cared to establish his
cult. The Erinyes are vindictive; nevertheless, they are moral, and
the struggle between them and Apollo in the Aeschylean drama is
only the contest between a more barbaric and a more civilised
morality. In the list of Greek divine titles and appellatives, only one or
two at most can be given a significance of evil.
 Doubtless, beneath the bright anthropomorphic religion lurked a
fear of ghosts and evil spirits, and the later days of Hellenic
paganism were somewhat clouded with demonology. But the
average Greek protected himself sufficiently by purification and easy
conventional magic. He did not brood on the principle of evil or
personify it as a great cosmic power, and therefore he would not
naturally evolve a system of religious dualism, though the germs
from which this might grow may be found in Orphic tradition and
doctrine. Contrast this with the evidence from Egypt, Assyria, and
Persia. The Egyptian and Assyrian records bear strong impress of
the prominence and power of the belief in evil spirits. The high gods
of Assyria were continually being invoked and implored by the
worshipper to save him from the demons, and one of these, Ira, a
demon of pestilence, seems to have received actual worship; and
much of Egyptian private ritual was protective magic against them.
But nowhere did the power of evil assume such grand proportions
as in the old Mazdean creed of Persia, and the dualism between the
good and evil principle became here the foundation of a great
religion that spread its influence wide through the West. The religion
had its prophet, Zarathustra, in whose historic reality we ought not to
doubt. In his system the faithful Mazdean is called upon to play his
part in the struggle between Ahura-Mazda and Angra-Mainyu, and
this struggle continues through the ages till in the final cataclysm the
Daevas, or evil demons, will be overthrown. We note here that this
faith includes the idea of a final Judgment, so familiar to Judaic and
Christian thought, but scarcely to be found in the native Hellenic
religion. Further, it should be observed that the Mazdean dualism
between good and evil has nothing in common with the Platonic
antithesis between mind and sense, or St. Paul’s between spirit and
flesh, or with the hatred of the body that is expressed in Buddhism.
The good Mazdean might regard his body as good and pure, and
therefore he escaped, as by a different way did the Greek, from the
tyranny of a morbid asceticism. Only he developed the doctrine of
purity into a code more burdensome than can be found, I think,
elsewhere. The ideas of ritual-purity on which he framed this code
are found broadcast through the East and in Egypt, and appear in
the Hellenic religion also. The Greek, however, did not allow himself
to be oppressed by his own cathartic system, but turned it to
excellent service in the domain of law, as I have tried to show
elsewhere.20.1
Generally, as regards the association of religion and morality, we
find this to be always intimate in the more developed races, but our
statistics are insufficient for us to determine with certainty the
comparative strength of the religious sanction of morals in the
ancient societies of the Mediterranean. The ethical-religious force of
the Zarathustrian faith seems to approach that of the Hebraic. We
should judge it to be stronger, at least, than any that was exercised
in Hellas, for Hellas, outside the Orphic sects, had neither sacred
books of universal recognition nor a prophet. Yet all Hellenic morality
was protected by religion, and the Delphic oracle, which occasionally
was able to play the part of the father-confessor, encouraged a high
standard of conduct—as high as the average found elsewhere in the
ancient world. We may note, however, one lacuna in the Hellenic
code: neither Greek ethics, on the whole, nor Greek religion,
emphasised or exalted or deified the virtue of truth; but we hear of a
goddess of Truth in Egypt, and it becomes a cardinal tenet and a
divine force in the Zarathustrian ideal.
Again, in all ancient societies religion is closely interwoven with
political, legal, and social institutions, and its influence on these
concerns the history of the evolution of society and law. It is only in
modern society, or in a few most ideal creeds at periods of great
exaltation, that a severance is made between Cæsar and God. Save
Buddhism, the religions of the ancient societies of the East and of
Egypt were all in a sense political. Darius regards himself as
specially protected by Ahura-Mazda, and we are told by
Herodotus21.1 that in the private Persian’s prayers no separate
personal benefits were besought, but only the welfare of the King
and the Persian community. The gods of Assyria inspire counsel and
order the campaign; Shamash, the sun-god, is “Just Ruler” and the
“Lord of Law”; and Ninib is styled the god “who lays for ever the
foundation-stone of the State,” and who, like Zeus ὅριος, and the
Latin Terminus, “protects the boundaries of the cornfield.” The Syrian
goddess of Bambyke, Kybele of Phrygia, Astarte of Askalon, all wear
the mural crown, the badge of the city goddess. But I doubt if our
materials are as yet rich enough to inform us in what precise way
religion played a constructive part in the oldest civilisations,—
namely, those of Assyria and Egypt. We may observe that the code
of Hammurabi, our oldest legal document, is curiously secular and in
many respects modern.
The question can be most fruitfully pursued in the study of the
Greek societies; for no other religion of which we have any record
was so political as the Hellenic, not even, as I should judge, the
Roman, to which it bears the closest resemblance in this respect.
The very origin of the πόλις, the city-state, was often religious; for
the name or title of the deity often gave a name to the city, and the
temple was in this case probably the centre of the earliest residence.
In the organised and complex Greek societies, every institution of
the State—the assembly, the council, the law-courts, the agrarian
economy, all the regulations of the family and clan—were
consecrated and safeguarded by the supervision of some deity.
Often he or she was worshipped as in a literal sense the State
ancestor, and in one of the temples might be found burning the
perpetual fire which symbolised the permanence of the city’s life.
And in Greece we find a unique phenomenon, which, though small,
is of great significance—the deity might here and there be made to
take the office and title of a civic magistrate. For instance, Apollo
was στεφανηφόρος in Asia Minor cities, and in the later days of
Sparta, as the recent excavations have shown, the ghost of
Lykourgos was elected as the chief inspector of the education of the
young.
To the superficial observer, then, the Greek civic society might
appear a theocracy. But such a view would imply ignorance of the
average character of the ancient Greek world. There can be no
theocracy where there is no theocrat. In Asia Minor the priest might
be a great political power, but in Greece this was never so. Here the
political, secular, utilitarian interest dominates the religion. The high
divinities become politicians, and immersed in secular affairs, and
even take sides in the party strife, as some of the religious titles
attest. Thus Greek religion escaped morbidity and insanity,
becoming genial and human, and compensating by its adaptability to
the common needs of social life for what it lacked of mystery and
aloofness. Therefore, also, in Greek invocations and hymns we do
not often hear the echo of that sublimity that resounds in the Iranian,
Assyrian, and still more in the Hebrew liturgies.
Another interesting point of comparison is the relation of religion to
the arts and sciences. Their association may be said to have been
more intimate in Hellenism than it has been found to be in any other
creed. We can estimate what music and the drama owed to Apollo
and Dionysos, and how the life of the philosopher, artist, and poet
was considered consecrated to certain divinities. We hear of the
Delphic oracle encouraging philosophic pursuits. The name
“Museum” is a landmark in the religious history of education, and we
know that the temple of Asklepios in Kos was the cradle of the
school of modern medicine. The records of the other religions of this
area show glimpses of the same association, and more extended
research may throw further light on it. The Babylonian gods Nebo
and Ea were divinities of wisdom and the arts, and to the former,
who was the inventor of writing, the library of Ashurbanapal was
consecrated. Chaldean astronomy was evolved from their astrology,
which was itself a religious system. But demonology was stronger in
Assyria, Persia, and Egypt than in Hellas, and demonology is the foe
of science. In the Zend-Avesta the priestly medicine-man, who heals
by spell and exorcism, is ranked higher than the scientific
practitioner. A chapter might be written on the negative advantages
of Greek religion, and none was of greater moment than this—that it
had no sacred books or authoritative religious cosmogony to oppose
to the dawn and the development of scientific inquiry. Asklepios had
been a practitioner in the method of thaumaturgic cures, but he
accepted Hippokrates genially when the time came.
As regards the relation between Greek philosophy and Greek
religion, something may remain to be discovered by any scholar who
is equally familiar with both. It would be absurd to attempt to
summarise the facts in a few phrases here. I wish merely to indicate
the absence in pure Hellenic speculation of any elaborated system of
theosophy, such as the late Egyptian “gnosis,” till we come to Neo-
Platonism, when the Greek intellect is no longer pure. We discover
also a vacuum in the religious mind and nomenclature of the earlier
Greek: he had neither the concept nor any name to express the
concept of what we call “faith,” the intellectual acceptance and
confessional affirmation of certain dogmas concerning the divinity;
and in this respect he differed essentially not only from the Christian,
but also from the Iranian and Buddhistic votary.
A great part of the study of ancient religion is a study of ritual, and
it is interesting to survey the Mediterranean area, so as to discern
similarities or divergencies in the forms of religious service.
Everywhere we observe the blood-sacrifice of animals, and very
frequently the harmless offering of fruits and cereals, and now one,
now the other, in Greece as elsewhere, was regarded as the more
pious. The former is of the higher interest, for certain ideas which
have been constructive of higher religions—our own, for example—
have grown out of it. At first sight the animal oblation seems
everywhere much the same in character and significance. The
sacrificial ritual of Leviticus does not differ in any essential trait from
that which commended itself to the Greeks and the other peoples of
these lands. Certain animals are everywhere offered, at times as a
free and cheerful gift, at other times as an atonement to expiate sin
and to deprecate wrath. Certain other animals are tabooed, for
reasons that may repay searching out.
In most regions we have evidence of the practice of human
sacrifice, either as an established system or as an occasional
expedient. The motives that prompted it present an important and
intricate question to the modern inquirer. The two nations that grew
to abhor it and to protest against it were the Hebrew and the Greek,
though the latter did not wholly escape the taint of it; for he had
inherited the practice from his ancestral past, and he found it
indigenous in the lands he conquered. Repellent as the rite may be,
it much concerns the study of the religions of the cultured races.
Now, an interesting theory concerning sacrifice was expounded
and brought into prominent discussion by Professor Robertson Smith
in his Religion of the Semites, and in an earlier article in the
Encyclopædia Britannica—namely, that a certain type of ancient
sacrifice was a mystic sacramental communion, the worshipper
partaking of some sacred food or drink in which the spirit of the deity
was temporarily lodged. This mystic act, of which there is no clear
trace in the Old Testament, is reported from Egypt,25.1 and it appears
to have been part of the Attis ritual of Phrygia. We find doubtful
traces of it in the Eleusinian and Samothracian mysteries; also a
glimpse of it here and there in the public religion of Hellas. But it is
best attested as a potent force in the Dionysiac worship, especially in
a certain savage ritual that we may call Thracian, but also in the
refined and Hellenised service as well.
I cannot dwell here on the various aspects of this problem. The
Hellenic statistics and their significance I have partly collected and
estimated in a paper published some years ago.25.2 The application
of the sacramental idea to the explanation of the Eleusinian
mysteries, ingeniously attempted by Dr. Jevons, I have discussed in
the third volume of my Cults of the Greek States; and the Dionysiac
communion-service is considered at length in the fifth.
The attractiveness of the mystic appeal of the Sacrament appears
to have increased in the later days of paganism, especially in its
period of struggle with Christianity. That strangest rite of the expiring
polytheism, the ταυροβόλιον, or the baptism in bull’s blood, in the
worship of Kybele, has been successfully traced back by M. Cumont
to the worship of the Babylonian Anaitis. The sacramental concept
was the stronghold of Mithraism, but can hardly be regarded as part
of its heritage from Persia, for it does not seem to have been familiar
to the Iranian religion nor to the Vedic Indian. In fact, the religious
history of no other Aryan race discloses it with clearness, save that
of the Thraco-Phrygian and Hellenic. Was it, then, a special product
of ancient “Mediterranean” religious thought? It would be important
to know, and Crete may one day be able to tell us, whether King
Minos took the sacrament. Meantime, I would urge upon those who
are studying this phenomenon in the various religions the necessity
of precise definition, so as to distinguish the different grades of the
sacramental concept, for loose statements are somewhat rife about
it.
Apart from the ritual of the altar, there is another mode of attaining
mystic union with the divinity—namely, by means of a sacred
marriage or simulated corporeal union. This is suggested by the
initiation formulæ of the mysteries of Attis-Kybele. The cult of Kybele
was connected with that of the Minoan goddess, and the strange
legend of Pasiphae and the bull-god lends itself naturally to this
interpretation. The Hellenic religion also presents us with a few
examples of the holy marriage of the human bride with the god, the
most notable being the annual ceremony of the union of the
“Queen,” the wife of the King Archon, at Athens, with Dionysos. And
in the mysteries of later paganism, as well as in certain forms and
symbolism of early Christianity, Professor Dieterich has traced the
surviving influence of this rite.
Among all the phenomena of ritual, none are more interesting or in
their effects more momentous than the rites that are associated with
the dogma of the death of the divinity. That the high gods are
naturally mortal and liable to death is an idea that is certainly rare,
though it may be found in Egyptian and old Teutonic mythology; but
the dogma of the annual or periodic death and resurrection of the
divinity has been, and is, enacted in much peasant ritual, and
worked for the purposes of agrarian magic in Europe and elsewhere.
More rarely we find the belief attached to the mystic forms and faith
of some advanced religion, and it is specially in the Mediterranean
area where it appears in a high stage of development. It is a salient
feature of the Egyptian worship of Isis; of the Sumerian-Babylonian
ritual, in which the dead Thammuz was bewailed, and which
penetrated Syria and other parts of Asia Minor; of the worship of
Attis and Adonis in Phrygia and the Lebanon; and of certain shrines
of the Oriental Aphrodite. It is associated often with orgiastic sorrow
and ecstatic joy, and with the belief in human immortality of which
the resurrection of the deity is the symbol and the efficacious means.
This idea and this ritual appears to have been alien to the native
Hellenic religion. The Hellenic gods and goddesses do not die and
rise again.
Only in one Aryan nation of antiquity, so far as I am aware, was
the idea clear and operative—the Thraco-Phrygian, in the religion of
Dionysos-Sabazios. This alien cult, when transplanted into Greece,
retained still some savagery in the rite that enacted the death of the
god; but in the Orphic sects the ritual idea was developed into a
doctrine of posthumous salvation, from which the later pre-Christian
world drew spiritual comfort and some fertile moral conceptions. This
Thracian-Dionysiac influence in Hellas, though chastened and
sobered by the sanity of the national temperament, initiated the
Hellene into a certain spiritual mood that was not naturally evoked by
the native religion; for it brought into his polytheism a higher
measure of enthusiasm, a more ecstatic spirit of self-abandonment,
than it possessed by its own traditional bent. Many civilised religions
appear to have passed beyond the phase of orgiastic fervour. It
emerges in the old Egyptian ritual, and most powerfully in the religion
of Phrygia and of certain districts of Syria; but it seems to have been
alien to the higher Semitic and the Iranian religions, as it was to the
native Hellenic.
I have only been able here, without argument or detailed
exposition, to present a short summary of the more striking
phenomena in the religious systems of our spiritual ancestors. Many
of the problems I have stated still invite further research, which may
considerably modify our theories. I claim that the subject possesses
a masterful interest both in its own right and for the light it sheds on
ancient philosophy, ancient art, and ancient institutions. And it ought
in the future to attract more and more the devotion of some of our
post-graduate students. Much remains to be done even for the
Hellenic and Roman religions, still more for those of Egypt and
Assyria. Here, in our University of Oxford, under whose auspices the
Sacred Books of the East were translated, and where the equipment
for the study is at least equal to that of any other centre of learning,
this appeal ought not to be made in vain.
 CHAPTER II.
Statement of the Problem and the Evidence.

The subject I have chosen for this course may appear over-
ambitious; and the attempt to pass critical judgment upon the facts
that arise in this wide comparative survey may be thought
premature. For not only is the area vast, but large tracts of it are still
unexplored, while certain regions have yielded materials that are
ample and promising, but of which the true interpretation has not yet
been found. We have, for instance, abundant evidence flowing in
with ever-increasing volume of the Sumerian-Babylonian religion, but
only a portion of the cuneiform texts has as yet been authoritatively
translated and made available for the service of Comparative
Religion. The Hittite monuments are witnesses of primary value
concerning Hittite religion: but the Hittite script may reveal much
more that is vital to our view of it, and without the help of that script
we are not sure of the exact interpretation of those religious
monuments; but though we have recently heard certain encouraging
expressions of hope, the master-word has not yet been found that
can open the door to this buried treasure of knowledge. And again,
the attempt to gauge accurately the relation and the indebtedness of
Greek religion to that of the near East cannot be wholly successful,
until we know more of the Minoan-Mycenaean religion; and our hope
hangs here partly on the discovery of more monuments, but mainly, I
am convinced, on the decipherment of the mysterious Minoan
writing, to which great achievement Sir Arthur Evans’ recent work on
the Scripta Minoa is a valuable contribution. Therefore the time is
certainly not yet ripe for a final and authoritative pronouncement on
the great questions that I am venturing upon in this course.
But even the early premature attempts to solve a problem may
contribute something to the ultimate satisfying solution. And often in
the middle of our investigations, when new evidence continues to
pour in, there comes a moment when it is desirable to look around
and take stock, so to speak, to consider whether we can draw some
general conclusions with safety, or in what direction the facts appear
at this stage to be pointing. In regard to the religions of anterior Asia
and South-Eastern Europe, and the question of their relationships,
this is now, I feel, a seasonable thing to do—all the more because
the Asiatic region has been mainly explored by specialists, who have
worked, as was profitable and right, each in his special province,
without having the time or perhaps the training to achieve a
comparative survey of the whole. We know also by long experience
the peculiar dangers to which specialists are prone; in their
enthusiastic devotion to their own domain, they are apt to believe
that it supplies them with the master-key whereby to unlock many
other secret places of human history. This hope, soon to prove an
illusion, was regnant when the interpretation of the Sacred Vedic
Books was first accomplished. And now certain scholars, who are
distinguished specialists in Assyriology, are putting forward a similar
claim for Babylon, and are championing the view that the Sumerian-
Assyrian religion and culture played a dominating part in the
evolution of the Mediterranean civilisation, and that therefore much
of the religious beliefs and practices of the early Greeks and other
European stocks must be traced back to Mesopotamia as their
fountain-head.31.1 This will be encouraging to that distinguished
writer on Greek religion, Dr. O. Gruppe, who almost a generation ago
proclaimed in his Griechische Mythologie the dogma of the
emanation of all religion from a single centre, and the dependence of
Greece upon the near East.
Now there ought to be no prejudice a priori against such a theory,
which stands on a different footing from what I may call the Vedic
fallacy: and it is childish to allow to the Aryan, or any other racial
bias, any malignant influence in these difficult discussions. Those
who have worked for years upon the marvellously rich records of
Mesopotamian culture, whether at first hand or, like myself, at
second hand, cannot fail to receive the deepest impression of its
imperial grandeur and its forceful vitality, and of its intensity of
thought and purpose in the sphere of religion. Naturally, they may
feel, such spiritual power must have radiated influence far and wide
over the adjacent lands; and no one could maintain that South-
Eastern Europe was too remote to have been touched, perhaps
penetrated, by it. For we know that, under certain conditions, the
race-barrier falls down before the march of a conquering and
dominating religion. And now, in the new light of a wider historical
survey, instead of saying, as once was said, “What is more its own
than a people’s gods?” we may rather ask, “What is less its own than
a people’s gods?” always, however, remembering that race-tradition,
inherited instinctive feeling and thought, is very strong in these
matters, and that a people will, often unconsciously, cling to its
ancestral modes of religious consciousness and expression, while it
will freely borrow alien forms, names, and ritual.
The inquiry indicated by the title of these lectures is naturally
twofold; it may be applied either to the earlier or the later periods of
the Hellenic and Hellenic-Roman history. The question concerning
the later period, though much critical research is needed for its clear
solution, is far simpler and more hopeful: for the evidence is
immeasurably fuller and more precise, and historical dates and
landmarks are there to help. The history of the invasion of the West
by Mithraism has been masterfully stated by Cumont; the general
influence of the Anatolian religions upon Graeco-Roman society is
presented and estimated by the same writer in his Religious
Orientales; by Toutain, in his Les Cultes Paiens dans l’Empire
Romain; by our own scholar, Samuel Dill, in Roman Society in the
last Century of the Western Empire; and more summarily by
Salomon Reinach in his Orpheus. Therefore I am not going to
pursue the inquiry at this end, although I may have to notice and use
some of the later evidence. I am going to raise the question
concerning the very origins of the Hellenic religious system, so as to
test the recently proclaimed dogma of certain Assyriologists, and to
determine, if possible, whether the Orient played any formative part
in the organic development of Greek religion. For this is just the
question which, I venture to maintain, has never yet been critically
explored. From what I have said at the beginning, it is obvious that I
cannot promise final and proved results. It will be gain enough if we
can dimly discern something behind the veil that shrouds the origins
of things, can reach to something that has the air of a reasoned
scientific hypothesis, and still more if we can indicate the paths along
which one day light may come.
 We may then begin at once with stating more clearly what are the
necessary conditions for a successful solution of the problem. First,
we must accomplish a thorough exploration of the religions of the
Anatolian and Mesopotamian lands; secondly, we must explore the
Minoan-Mycenaean religion, and estimate the strength of its
influence on the later period; thirdly, we must be able to decide what
beliefs and practices the Hellenes brought with them from the North;
lastly, before we can hope for any precision in our results, we must
be able to answer with some degree of accuracy a burning
chronological question: What was the date of the arrival through the
Balkans from the North of those Aryan-speaking tribes that by
mingling with the Southerners formed the Hellenic people of history?
For only then shall we be able to test the whole question, by
considering the position of the Eastern powers at this momentous
epoch. The third of these inquiries, concerning the aboriginal
religious ideas of the earliest Aryan Hellenes, is perhaps the most
troublesome of all. I may venture upon it at a later occasion, but it is
far too difficult and extensive to combine it with the others in a short
treatise. Nor can I do more than touch lightly on the Minoan-
Mycenaean period; for I wish to devote the greater part of these
lectures to the comparative survey of Greece, Anatolia, and
Mesopotamia, as this task has never yet been critically performed.
Something like an attempt was made by Tiele in his Histoire des
anciennes Religions, but when that book was written much of the
most important evidence had not yet come in.
But before beginning the exploration of any large area, whether for
the purposes of Comparative Religion, archaeology, or anthropology,
we must possess or acquire certain data of ethnography and secular
history. We must, for instance, face the chronological question that I
mentioned just above, before we can estimate the formative
influences at work in the earliest phases of Hellenic development.
Recent archaeological evidence, which I cannot here discuss,
renders us valuable aid at this critical point of our inquiry. We can no
longer relegate the earliest Hellenic invasion of Greece to a very
remote period of Mediterranean history. The arguments from the
Minoan culture, combined with the still more striking evidence, of
which the value is not yet fully appreciated, obtained by the recent
excavations of the British School on the soil of Thessaly,34.1 point to
the conviction that this, the epoch-making event of the world’s
secular and spiritual life, occurred not much earlier than 1500 B.C.
On this hypothesis, our quest becomes less vague. We can consider
what influences were likely to be radiating from the East upon the
opposite shores of the Aegean during those few centuries, in which
the Hellenic tribes were passing from barbarism to culture, and the
religious beliefs and ritual were developing into that comparatively
advanced and complex form of polytheism which is presented about
1000 B.C. in the Homeric poems. By this date we may assert that
the Hellenic spirit had evolved certain definite traits and had acquired
a certain autonomous power. While continuing always to be quickly
responsive to alien influence, it would not henceforth admit the alien
product with the submissive and infantine docility of barbarians In
fact, when we compare the Homeric religion with that of the fifth
century, we feel that in this particular sphere of the social and
spiritual life the Hellene in many essentials had already come to his
own in the Homeric period. Therefore, in trying to track the earliest
streams of influences that moulded his religious consciousness,
what was operative before the tenth century is of more primary
importance than what was at work upon him afterwards.
Now the most recent researches into Mesopotamian history
establish with certainty the conclusion that there was no direct
political contact possible between the powers in the valley of the
Euphrates and the western shores of the Aegean, in the second
millennium B.C. It is true that the first Tiglath-Pileser, near the end of
the twelfth century, extended the Assyrian arms to the shores of
South-Eastern Asia, to Cilicia and Phoenicia35.1; but there does not
seem to have been any permanent Assyrian or Babylonian
settlement on this littoral. The city of Sinope in the north, which, as
the legend attests and the name that must be derived from the
Assyrian god Sin indicates, was originally an Assyrian colony, was
probably of later foundation, and geographically too remote to count
for the present inquiry. In fact, between the nascent Hellas and the
great world of Mesopotamia, there were powerful and possibly
independent strata of cultures interposing. We have to reckon first
with the great Hittite Kingdom, which included Cappadocia and