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"Ontogeny recapitulates phylogeny" was Haeckel's answer – the wrong one – to the most vexing question of nineteenth-century biology: what is the relationship between individual development (ontogeny) and the evolution of species and lineages (phylogeny)? In this, the first major book on the subject in fifty years, Stephen Jay Gould documents the history of the idea of recapitulation from its first appearance among the pre-Socratics to its fall in the early twentieth century.
Mr. Gould explores recapitulation as an idea that intrigued politicians and theologians as well as scientists. He shows that Haeckel's hypothesis – that human fetuses with gill slits are, literally, tiny fish, exact replicas of their water-breathing ancestors – had an influence that extended beyond biology into education, criminology, psychoanalysis (Freud and Jung were devout recapitulationists), and racism. The theory of recapitulation, Gould argues, finally collapsed not from the weight of contrary data, but because the rise of Mendelian genetics rendered it untenable.
Turning to modern concepts, Gould demonstrates that, even though the whole subject of parallels between ontogeny and phylogeny fell into disrepute, it is still one of the great themes of evolutionary biology. Heterochrony – changes in developmental timing, producing parallels between ontogeny and phylogeny – is shown to be crucial to an understanding of gene regulation, the key to any rapprochement between molecular and evolutionary biology. Gould argues that the primary evolutionary value of heterochrony may lie in immediate ecological advantages for slow or rapid maturation, rather than in long-term changes of form, as all previous theories proclaimed.
Neoteny – the opposite of recapitulation – is shown to be the most important determinant of human evolution. We have evolved by retaining the juvenile characters of our ancestors and have achieved both behavioral flexibility and our characteristic morphology thereby (large brains by prolonged retention of rapid fetal growth rates, for example).
Gould concludes that "there may be nothing new under the sun, but permutation of the old within complex systems can do wonders. As biologists, we deal directly with the kind of material complexity that confers an unbounded potential upon simple, continuous changes in underlying processes. This is the chief joy of our science".
1. Prospectus
Part I: Recapitulation
2. The Analogistic Tradition from Anaximander to Bonnet
The Seeds of Recapitulation in Greek Science?
Ontogeny and Phylogeny in the Conflict of “Evolution” and Epigenesis: The Idyll of Charles Bonnet
Appendix: The Revolution in “Evolution”
3. Transcendental Origins, 1793–1860
Naturphilosophie: An Expression of Developmentalism
Two Leading Recapitulationists among the Naturphilosophen: Oken and Meckel
Oken’s Classification of Animals Linear Additions of Organs
J. F. Meckel’s Sober Statement of the Same Principles
Serres and the French Transcendentalists
Recapitulation and the Theory of Developmental Arrests
Von Baer’s Critique of Recapitulation
The Direction of Development and Classification of Animals
Von Baer and Naturphilosophie: What Is the Universal Direction of Development?
Louis Agassiz and the Threefold Parallelism
4. Evolutionary Triumph, 1859–1900
Evolutionary Theory and Zoological Practice
Darwin and the Evolution of Von Baer’ Laws
Evolution and the Mechanics of Recapitulation
Ernst Haeckel: Phylogeny as the Mechanical Cause of Ontogeny
The Mechanism of Recapitulation
The American Neo-Lamarckians: The Law of Acceleration as Evolution’s Motor
Progressive Evolution by Acceleration
The Extent of Parallelism
Why Does Recapitulation Dominate the History of Life?
Alpheus Hyatt and Universal Acceleration
Lamarckism and the Memory Analogy
Recapitulation and Darwinism
Appendix: The Evolutionary Translation of von Baer’s Laws
5. Pervasive Influence
Criminal Anthropology
Racism
Child Development
Primary Education
Freudian Psychoanalysis
Epilogue
6. Decline, Fall, and Generalization
A Clever Argument
An Empirical Critique
Organs or Ancestors: The Transformation of Haeckel’s Heterochrony
Interpolations into Juvenile Stages
Introduction of Juvenile Features into the Adults of Descendants
What Had Become of von Baer’s Critique?
Benign Neglect: Recapitulation and the Rise of Experimental Embryology
The Prior Assumptions of Recapitulation
Wilhelm His and His Physiological Embryology: A Preliminary Skirmish
Roux’s Entwicklungsmechanik and the Biogenetic Low
Recapitulation and Substantive Issues in Experimental Embryology: The New Preformationism
Mendel’s Resurrection, Haeckel’s Fall, and the Generalization of Recapitulation
Part II: Heterocrony and Paedomorphosis
7. Heterochrony and the Parallel of Ontogeny and Phylogeny
Acceleration and Retardation
Confusion in and after Haeckel’s Wake
Guidelines for a Resolution
The Reduction of de Beer’s Categories of Heterochrony to Acceleration and Retardation
A Historical Paradox: The Supposed Dominance of Recapitulation
Dissociability and Heterochrony
Correlation and Disociability
Dissociation of the Three Processes
A Metric for Dissociation
Temporal Shift as a Mechanism of Dissociation
A Clock Model of Heterochrony
Appendix: A Note on the Multivariate Representation of Dissociation
8. The Ecological and Evolutionary Significance of Heterochrony
The Argument from Frequency
The Importance of Recapitulation
The Importance of Heterochronic Change: Selected Cases
Frequency of Paedomorphosis in the Origin of Higher Taxa
A Critique of the Classical Significance of Heterochrony
The Classical Arguments
Retrospective and Immediate Significance
Heterochrony, Ecology, and Life-History Strategies
The Potential Ease and Rapidity of Heterochronic Change
The Control of Metamorphosis in Insects
Amphibian Paedomorphosis and the Thyroid Gland
9. Progenesis and Neoteny Insect Progenesis
Prothetely and Metathetely
Paedogenesis (Parthenogenetic Progenesis) in Gall Midges and Beetles
Progenesis in Wingless, Parthenogenetic Aphids
Additional Cases of Progenesis with a Similar Ecological Basis
Neotenic Solitary Locusts: Are They an Exception to the Rule?
Amphibian Neoteny
The Ecological Determinants of Progenesis
Unstable Environments
Colonization
Parasites
Male Dispersal
Progenesis as an Adaptive Response to Pressures for Small Size
The Role of Heterochrony in Macroevolution: Contrasting Flexibilities for Progenesis and Neoteny
Progenesis
Neoteny
The Social Correlates of Neoteny in Higher Vertebrates
10. Retardation and Neoteny in Human Evolution
The Seeds of Neoteny
The Fetalization Theory of Louis Bolk
Bolk’s Data
Bolk’s Interpretation
Bolk’s Evolutionary Theory
A Tradition of Argument
Retardation in Human Evolution
Morphology in the Matrix of Retardation
Of Enumeration
Of Prototypes
Of Correlation
The Adaptive Significance of Retarded Development
11. Epilogue
Notes
Bibliography
Glossary
Index
Stephen Jay Gould was the Alexander Agassiz Professor of Zoology at Harvard University and Vincent Astor Visiting Professor of Biology at New York University. A MacArthur Prize Fellow, he received innumerable honors and awards and wrote many books, including Ontogeny and Phylogeny and Time's Arrow, Time's Cycle (both from Harvard).
"In Gould's [...] new book [...] Ontogeny and Phylogeny, a scholarly study of the theory of recapitulation, he not only explains scientific theory but comments on science itself, with clarity and wit, simultaneously entertaining and teaching [...] [This] is a rich book."
– James Gorman, The New York Times Book Review
"Steven Jay Gould has given us a superb analysis of the use of ontogenetic analogy, the controversies over ontogeny and phylogeny, and the classification of the different processes observable in comparing different ontogenies. His massive book (in each chapter of which there is as much material as in whole books by other writers) is both a historical exposition of the whole subject of ontogeny and phylogeny, and [...] a fascinating attempt at a functional interpretation of those phylogenetic alterations that involve changes of timing developmental processes in related organisms."
– A.J. Cain, Nature
"This [is a] fat, handsome book crammed with provocative ideas [...] Ontogeny and Phylogeny is an important and thoughtful book which will be a valuable source of ideas and controversies for anyone interested in evolutionary or developmental biology."
– Matt Cartmill, Science
"It is rare indeed to read a new book and recognize it for a classic [...] Gould has given biologists a new way to see the organisms they study. The result is a major achievement."
– S. Rachootin, American Scientist
"Gould's book – pervaded, I should say, with an erudition and felicity of style that make it a delight to read – is a radical work in every sense [...] It returns one's attention to the roots of our science – the questions about the great pageant of evolution, the marvelous diversity of form that our theory is meant to explain."
– D. Futuyma, Quarterly Review of Biology
"A distinguished and pioneering work."
– Ernst Mayr
