Form and Function: A Contribution to the History of Animal Morphology Informative Summary


This book delves into the historical development of animal morphology, exploring the diverse perspectives and key figures who shaped this field. The author, E.S. Russell, focuses on three major currents of morphological thought: the functional, the formal, and the materialistic.

The functional approach, championed by Aristotle, Cuvier, and von Baer, emphasizes the importance of function in determining form. This perspective sees living organisms as active, purposeful agents whose structures are adapted to their needs and environments. The formal approach, most prominently represented by E. Geoffroy St. Hilaire, prioritizes the unity of plan and composition, seeking underlying structural principles that govern all forms of life. Lastly, the materialistic attitude, prevalent across various fields of thought, seeks to explain biological phenomena entirely in terms of physical and chemical processes.

Key Findings:

  • The book highlights the ongoing tension between the functional and formal approaches to animal morphology, emphasizing the importance of understanding the intricate interplay of structure and function.
  • The rise of embryology, particularly with the work of von Baer, revolutionized morphology by introducing a new criterion for understanding homologies and unveiling the significance of developmental processes.
  • The cell theory, while valuable in its contributions to histology, also influenced morphology to focus on cellular processes, sometimes leading to a neglect of the broader organization and functional aspects of living organisms.
  • The book examines the limitations of purely speculative approaches to evolution, emphasizing the importance of empirical evidence and the need to integrate data from various disciplines, such as palæontology.


  • The concept of Unity of Plan: This concept, initially recognized by Aristotle, emphasizes the shared structural features within major animal groups, suggesting a common underlying plan. It has been interpreted both functionally (as adaptation to similar needs) and formally (as a manifestation of a universal blueprint).
  • The importance of Embryological Evidence: The work of von Baer and his successors demonstrated that studying embryonic development is crucial for understanding homologies and the evolution of complex structures. This approach proved instrumental in challenging the oversimplification of the Meckel-Serres law, which emphasized recapitulation of adult forms in the embryo.
  • The Complexity of the Organism: The organism is not simply an aggregation of cells, as suggested by the cell theory and its proponents. The functional approach, championed by Cuvier and later by Roux, recognizes the intricate interplay of parts and the importance of considering the organism as a whole, emphasizing the interdependence of function and structure.
  • The Influence of Function: The book stresses that function is a primary driver of form. While some structures are inherited, others arise and evolve through active adaptation and response to environmental pressures. This concept of functional adaptation is a core principle of the book and emphasizes the dynamism of living organisms.

Historical Context:

The text provides insights into the intellectual climate of the early 20th century. The author emphasizes the prevailing influence of Darwinism and the rise of mechanistic explanations for biological phenomena. However, he also points to a growing dissatisfaction with simplistic materialistic interpretations of life and form, suggesting a potential shift toward a more nuanced and integrated approach.


  • Aristotle’s extensive knowledge of animals: He described over 500 species, including various invertebrates like cuttlefish, snails, and insects.
  • Aristotle’s first scientific classification of animals: He divided them into Sanguinea (with red blood) and Exsanguinea (without red blood), establishing fundamental groups.
  • Aristotle’s recognition of unity of plan: He recognized that animals within each major group were built upon one structural plan, despite variations.
  • Aristotle’s emphasis on function: He saw form as a consequence of function, emphasizing the adaptedness of structure.
  • Aristotle’s recognition of correlation: He noted relationships between organs that were not directly linked by function, such as the absence of horns in animals with upper jaw front teeth.
  • Aristotle’s distinction between homogeneous and heterogeneous parts: He distinguished between tissues (homogeneous) and organs (heterogeneous) based on their material composition and function.
  • Aristotle’s foreshadowing of von Baer’s law: He recognized that general characters develop before specific ones in embryos.
  • Aristotle’s introduction of the Échelle des êtres: He envisioned a graded series of organisms, from simple to complex, highlighting the gradual transition from inanimate to animate things.
  • Belon’s comparison of bird and mammal skeletons: He was one of the first to show the homology of bones by placing bird and mammal skeletons side by side, highlighting their shared structural plan.
  • Fabricius’s observations on chick development: He made early contributions to comparative embryology by studying the development of the chick.
  • Willis’s emphasis on comparative brain structure: He recognized the importance of studying the brain of different animals to understand its structure and function.
  • Claude Perrault’s recognition of unity of type: He saw the same basic plan underlying the organization of animals, even suggesting an arterial system in plants.
  • Malpighi’s pioneering histological studies: He investigated the microscopic structure of various tissues, revealing the complexity of the liver and discovering Malpighian bodies in the kidney.
  • Swammerdam’s meticulous dissections: He performed incredibly detailed anatomical studies of insects and other small animals.
  • John Hunter’s vast anatomical experience: He dissected a wide range of animals, highlighting his interest in functional anatomy.
  • Bonnet’s complete expression of the Échelle des êtres: He extended the concept of gradation to all of nature, creating a hierarchical scheme for all organic and inorganic beings.
  • Buffon’s tentative evolutionism: He suggested that species within a family might have descended from a single type species, foreshadowing Darwin’s ideas.
  • Bichat’s distinction between animal and organic life: He differentiated the functions and organs related to the individual (organic life, centered on the heart) from those related to the species (animal life, centered on the brain).
  • Cuvier’s recognition of functional unity: He argued that organs are not merely assembled, but are interdependent and function together for the good of the whole organism.
  • Cuvier’s principle of correlation: This principle states that from the structure of one organ, one can deduce the structure of other organs due to their interconnected function.
  • Cuvier’s classification of animal types: He established four primary types: Vertebrates, Molluscs, Articulates, and Radiates, based on their neuromuscular systems.
  • Cuvier’s opposition to evolution: He argued that the absence of transitional forms in the fossil record and the limited variability of species contradicted the idea of gradual evolution.
  • E. Geoffroy’s “principle of connections”: He argued that the position and relationships of parts were more important than form in determining homology.
  • E. Geoffroy’s “theory of analogues”: He proposed that all animal organs are variations of a common scheme, emphasizing unity of composition.
  • E. Geoffroy’s “law of balancement”: This law suggests that an increase in the development of one organ can be balanced by a decrease in another.
  • Serres’s extension of Meckel-Serres law: He proposed that the development of higher animals recapitulates the adult forms of lower animals.
  • Savigny’s study of Arthropod appendages: He demonstrated the unity of plan in the appendages of Arthropods, showing their homology despite functional modifications.
  • Oken’s vertebral theory of the skull: He proposed that the skull is composed of modified vertebrae, a concept adopted by many subsequent morphologists.
  • Meckel’s concept of “homologues”: He drew parallels between organs, sometimes with dubious validity, suggesting a repetition of the whole organism in its parts.
  • Rathke’s discovery of gill-slits in mammalian embryos: This finding was a crucial piece of evidence for the evolutionary relationship of vertebrates and provided significant insight into embryonic development.
  • Rathke’s detailed account of skull development: He demonstrated that the basilar plate and trabeculae in the developing skull provided a framework for the later ossification of specific bones.
  • Reichert’s work on visceral arches and ear ossicles: He established the homologies of the ear ossicles with structures in the first and second visceral arches, contributing significantly to our understanding of the development of the head.
  • Huxley’s defence of the embryological method: He argued that studying development is the ultimate criterion for determining homologies.
  • Agassiz’s classification of skull bones in fish: He categorized skull bones based on their development from cartilage or membrane, providing a foundation for subsequent studies.
  • Kölliker’s distinction between membrane and cartilage bones: He argued that membrane bones, arising from fibrous tissue, differed significantly in their development and morphological significance from cartilage bones.
  • Dollo’s theory of arboreal Marsupial ancestry: He inferred that the characteristics of the marsupial foot provided evidence of an arboreal past.
  • Roux’s concept of functional adaptation: He argued that many organs develop and are maintained in direct response to the functions they perform.
  • Roux’s experiments on isolated blastomeres: He demonstrated the role of cell division in early development, contributing to the understanding of the “mosaic theory.”
  • Driesch’s challenge to the mosaic theory: His experiments with sea urchin eggs showed that complete embryos could develop from isolated blastomeres, indicating a degree of regulatory capacity in development.
  • Wilson’s discovery of cytoplasmic organ-forming stuffs: He showed that specific regions of the cytoplasm contribute to the development of specific organs in the embryo.
  • Hering’s theory of memory as a general function of organized matter: He proposed that memory, a function of the brain, could be viewed as a broader property of all organized matter, potentially linking it to heredity.
  • Semon’s mnemic theory: He proposed that stimuli leave permanent material traces (engrams) in the organism, affecting its subsequent behavior and contributing to heredity.
  • Rignano’s theory of “specific nervous accumulators”: He suggested that functional stimuli lead to the deposition of specific substances in the nucleus, contributing to heredity and explaining the development of acquired characters.


  • 500+ species of animals described by Aristotle
  • 9 sternal elements composing the sternum in all Vertebrates (according to Geoffroy)
  • 15 segments in all Arthropods (according to Latreille)
  • 4 cranial vertebrae in the vertebrate skull (according to Owen)
  • 3 cranial vertebrae in the vertebrate skull (according to Müller and Reichert)
  • 6 vertebrate types (according to von Baer)
  • 5 primary stages of ontogeny (according to Haeckel)
  • 4 vertebrate classes (according to Cuvier and von Baer)


  • Homology: Similarity of structure due to common ancestry.
  • Analogy: Similarity of function, not necessarily due to common ancestry.
  • Archetype: A hypothetical, generalized model of structure.
  • Metastasis: A shift in the position of an organ or organ system.
  • Loi de balancement: A law of compensation, suggesting that an increase in one organ can be balanced by a decrease in another.
  • Meckel-Serres law: The theory that the development of higher animals recapitulates the adult forms of lower animals.
  • Biogenetic law: The theory that ontogeny (individual development) recapitulates phylogeny (evolutionary history).
  • Palingenesis: The true recapitulation of ancestral history in development.
  • Cenogenesis: Secondary adaptations or modifications in development that obscure or alter the true course of phylogenetic recapitulation.
  • Heterochrony: Alterations in the timing of developmental events.
  • Heterotopy: Changes in the spatial location of developmental events.
  • Gastræa: A hypothetical, two-layered, ancestral form of all Metazoa.
  • Cœlom: The main body cavity of animals, arising from pouches of the archenteron.
  • Enterocœl: A cœlom arising from pouches of the archenteron.
  • Schizocœl: A cœlom arising from the splitting of the mesoblast.
  • Mneme: The memory principle, suggesting that stimuli leave lasting traces (engrams) in the organism.
  • Entwicklungsmechanik: A mechanistic approach to studying the causes of development.
  • Functional adaptation: Structures arising or evolving in response to the functions they perform.
  • Mosaic theory: The theory that developmental potencies are pre-determined in the germ and partitioned to specific cells.
  • Specific organ-forming stuffs: Specific substances in the cytoplasm of the egg that influence the development of specific organs.
  • Sentiment intérieur: An internal sense of need that drives behavior and influences evolutionary changes (Lamarck).


  • Aristotle’s comparison of the chameleon to other animals: He noted how the chameleon’s structure was similar to, yet different from, other animals, demonstrating his ability to observe and compare.
  • Belon’s skeletal comparison of birds and mammals: He highlighted the homology of bones by placing the skeletons of a bird and a mammal side by side.
  • Rathke’s discovery of gill-slits in a pig embryo: This finding was groundbreaking, providing evidence for the evolutionary connection between vertebrates and demonstrating the presence of ancestral features in development.
  • Reichert’s detailed study of ear ossicles: He traced the development of ear ossicles back to their origin in visceral arches, establishing key homologies.
  • Huxley’s examination of the skull in different vertebrates: He showed that the skull of fish, although cartilaginous, shared a basic plan with the bony skulls of other vertebrates, supporting his argument for an embryological archetype.
  • Kölliker’s observation of membrane bone formation: He demonstrated that specific bones, like the parietals and frontals, develop directly from fibrous tissue, highlighting the distinction between membrane and cartilage bones.
  • Dollo’s analysis of the marsupial foot: He used the structure of the foot in marsupials to infer their arboreal ancestry.
  • Roux’s experiments on the development of isolated blastomeres: His work demonstrated the role of cell division in development and supported the “mosaic theory.”
  • Driesch’s experiment with sea urchin eggs: His findings, showing that complete embryos could develop from isolated blastomeres, challenged the mosaic theory and underscored the organism’s capacity for regulation.
  • Wilson’s investigation of the cytoplasm in Dentalium: His work provided conclusive evidence for the existence of cytoplasmic organ-forming stuffs, emphasizing the role of the cytoplasm in development.


The book “Form and Function” offers a fascinating historical perspective on the evolution of animal morphology. It highlights the crucial role of key thinkers like Aristotle, Cuvier, von Baer, Geoffroy, Owen, and Darwin in shaping this field. It reveals the ongoing struggle between functional and formal interpretations of animal structure, emphasizing the importance of understanding the interplay of form and function. The book also emphasizes the significant contribution of embryology in redefining our understanding of homologies and revealing the historical dimension of living organisms. While acknowledging the valuable insights provided by evolutionary morphology, the book also points to its limitations and the need for a more comprehensive, integrated approach to understanding the complex interplay of form, function, and history in the evolution of life.

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