Structuralism (biology)

Last updated

In his 1917 book On Growth and Form, D'Arcy Thompson illustrates the geometric transformation of one fish's body form into another with a 20deg shear mapping. He does not discuss the evolutionary causes of such a structural change, and has accordingly been suspected of vitalism. Transformation of Argyropelecus olfersi into Sternoptyx diaphana.jpg
In his 1917 book On Growth and Form , D'Arcy Thompson illustrates the geometric transformation of one fish's body form into another with a 20° shear mapping. He does not discuss the evolutionary causes of such a structural change, and has accordingly been suspected of vitalism.

Biological or process structuralism is a school of biological thought that objects to an exclusively Darwinian or adaptationist explanation of natural selection such as is described in the 20th century's modern synthesis. It proposes instead that evolution is guided differently, basically by more or less physical forces which shape the development of an animal's body, and sometimes implies that these forces supersede selection altogether.

Contents

Structuralists have proposed different mechanisms that might have guided the formation of body plans. Before Darwin, Étienne Geoffroy Saint-Hilaire argued that animals shared homologous parts, and that if one was enlarged, the others would be reduced in compensation. After Darwin, D'Arcy Thompson hinted at vitalism and offered geometric explanations in his classic 1917 book On Growth and Form . Adolf Seilacher suggested mechanical inflation for "pneu" structures in Ediacaran biota fossils such as Dickinsonia . Günter P. Wagner argued for developmental bias, structural constraints on embryonic development. Stuart Kauffman favoured self-organisation, the idea that complex structure emerges holistically and spontaneously from the dynamic interaction of all parts of an organism. Michael Denton argued for laws of form by which Platonic universals or "Types" are self-organised. Stephen J. Gould and Richard Lewontin proposed biological "spandrels", features created as a byproduct of the adaptation of nearby structures. Gerd B. Müller and Stuart A. Newman argued that the appearance in the fossil record of most of the current phyla in the Cambrian explosion was "pre-Mendelian" [lower-alpha 1] evolution caused by physical factors. Brian Goodwin, described by Wagner as part of "a fringe movement in evolutionary biology", [2] denies that biological complexity can be reduced to natural selection, and argues that pattern formation is driven by morphogenetic fields.

Darwinian biologists have criticised structuralism, emphasising that there is plentiful evidence both that natural selection is effective and, from deep homology, that genes have been involved in shaping organisms throughout evolutionary history. They accept that some structures such as the cell membrane self-assemble, but deny the ability of self-organisation to drive large-scale evolution.

History

Multiple alternatives to Darwinism have been offered since the 19th century to explain how evolution took place, given that many scientists initially objected to natural selection. Many of these theories, including structural or developmental constraints, led (solid blue arrows) to some form of directed evolution (orthogenesis), with or without invoking divine control (dotted blue arrows). These theories were largely swept aside by the modern synthesis of genetics and natural selection in the early 20th century (dashed orange arrows). Alternatives to Darwinism.svg
Multiple alternatives to Darwinism have been offered since the 19th century to explain how evolution took place, given that many scientists initially objected to natural selection. Many of these theories, including structural or developmental constraints, led (solid blue arrows) to some form of directed evolution (orthogenesis), with or without invoking divine control (dotted blue arrows). These theories were largely swept aside by the modern synthesis of genetics and natural selection in the early 20th century (dashed orange arrows).

Geoffroy's law of compensation

In 1830, Étienne Geoffroy Saint-Hilaire argued a structuralist case against the functionalist (teleological) position of Georges Cuvier. Geoffroy believed that homologies of structure between animals indicated that they shared an ideal pattern; these did not imply evolution but a unity of plan, a law of nature. [lower-alpha 2] He further believed that if one part was more developed within a structure, the other parts would necessarily be reduced in compensation, as nature always used the same materials: if more of them were used for one feature, less was available for the others. [4]

D'Arcy Thompson's morphology

In his "eccentric, beautiful" [5] 1917 book On Growth and Form , D'Arcy Wentworth Thompson revisited the old idea of "universal laws of form" to explain the observed forms of living organisms. [1] The science writer Philip Ball states that Thompson "presents mathematical principles as a shaping agency that may supersede natural selection, showing how the structures of the living world often echo those in inorganic nature", and notes his "frustration at the 'Just So' explanations of morphology offered by Darwinians." Instead, Ball writes, Thompson elaborates on how not heredity but physical forces govern biological form. [6] The philosopher of biology Michael Ruse similarly wrote that Thompson "had little time for natural selection", certainly preferring "mechanical explanations" and possibly straying into vitalism. [1]

Seilacher's pneu structures

Dickinsonia fossil described as a "pneu" structure with chambers inflated like a quilted air mattress. In Adolf Seilacher's structuralist view, the structure is determined mechanically by the need to distribute the tension across the surface, rather than having been guided by natural selection. DickinsoniaCostata.jpg
Dickinsonia fossil described as a "pneu" structure with chambers inflated like a quilted air mattress. In Adolf Seilacher's structuralist view, the structure is determined mechanically by the need to distribute the tension across the surface, rather than having been guided by natural selection.

Like Thompson, the palaeontologist Adolf Seilacher emphasised fabricational constraints on form. He interpreted fossils such as Dickinsonia in the Ediacaran biota as "pneu" structures determined by mechanical inflation like a quilted air mattress, rather than having been driven by natural selection. [7] [8]

Wagner's constraints on development

In his 2014 book Homology, Genes, and Evolutionary Innovation, the evolutionary biologist Günter P. Wagner argues for "the study of novelty as distinct from adaptation." He defines novelty as occurring when some part of the body develops an individual and quasi-independent existence, in other words as a distinct and recognisable structure, which he implies might occur before natural selection begins to adapt the structure for some function. [2] [9] He forms a structuralist picture of evolutionary developmental biology, using empirical evidence, arguing that homology and biological novelty are key aspects requiring explanation, and that developmental bias (i.e. structural constraints on embryonic development) is a key explanation for these. [10] [11]

Kauffman's self-organisation

Darwinists and structuralists agree that cell structures like the cell membrane spontaneously self-organize. They disagree on how important self-organization is in other areas of biology. Fluid Mosaic.svg
Darwinists and structuralists agree that cell structures like the cell membrane spontaneously self-organize. They disagree on how important self-organization is in other areas of biology.

The mathematical biologist Stuart Kauffman suggested in 1993 that self-organization may play a role alongside natural selection in three areas of evolutionary biology, namely population dynamics, molecular evolution, and morphogenesis. With respect to molecular biology, Kauffman has been criticised for ignoring the role of energy in driving biochemical reactions in cells, which can fairly be called self-catalysing but which do not simply self-organise. [12]

Denton's 'Types'

The biochemist Michael Denton has argued a structuralist case for self-organization. In a 2013 paper, he claimed that "the basic forms of the natural world—the Types—are immanent in nature, and determined by a set of special natural biological laws, the so called 'laws of form'." He asserts that these "recurring patterns and forms" are "genuine universals". [lower-alpha 3] Form is in this view not shaped by natural selection, but by "self-organizing properties of particular categories of matter" and by "cosmic fine-tuning of the laws of nature". [14] Denton has been criticised by the biochemist Laurence A. Moran as anti-Darwinian and favouring creationism. [15]

Gould and Lewontin's spandrels

An ornamented bridge spandrel. Steven J. Gould and Richard Lewontin argued that the triangular area is a byproduct of the adaptation of structures around it. Central Park New York City New York 33.jpg
An ornamented bridge spandrel. Steven J. Gould and Richard Lewontin argued that the triangular area is a byproduct of the adaptation of structures around it.

In 1979, influenced by Seilacher among others, the paleontologist Stephen J. Gould and the population geneticist Richard Lewontin wrote what Wagner called "the most influential structuralist manifesto", "The Spandrels of San Marco and the Panglossian Paradigm". [16] [2] They pointed out that biological features (like architectural spandrels) did not necessarily have adaptation as their direct cause. Instead, architects couldn't help creating small triangular areas between arches and pillars, as arches need (evolve) to be curved, and pillars need to be vertical. The resulting spandrels are exaptations, consequences of other evolutionary changes. Evolution, they argued, did not select for a protruding human chin: instead, reducing the length of the tooth row left the jaw protruding. [2]

Müller and Newman's pre-Mendelian evolution

Extreme structuralists like Gerd B. Müller and Stuart A. Newman, inheriting the viewpoint of D'Arcy Thompson, have proposed that physical laws of structure, not genetics, govern major diversifications such as the Cambrian explosion, followed later by co-opted genetic mechanisms. [17] [18] They argued further that there was a "pre-Mendelian" phase of the evolution of animals, involving physical forces, before genes took over. [17] [19] Darwinian biologists freely admit that physical factors such as surface tension can cause self-assembly, but insist that genes play a crucial role. They note for example that deep homologies between widely separated groups of organisms, such as the signalling pathways and transcription factors of choanoflagellates and metazoans, demonstrate that genes have been involved throughout evolutionary history. [20]

Goodwin's morphogenetic fields

Natural patterns like those on the skin of the Giant pufferfish can be created by spatial oscillations of chemical signals. Giant Puffer fish skin pattern.JPG
Natural patterns like those on the skin of the Giant pufferfish can be created by spatial oscillations of chemical signals.

What Wagner calls "a fringe movement in evolutionary biology", [2] the form of structuralism exemplified by Brian Goodwin, [2] [21] effectively denies that natural selection is important, [2] [22] or at least that biological complexity could be reduced to natural selection. [22] [23] This led to conflict with Darwinists such as Richard Dawkins. [24] Goodwin related the old concept of a morphogenetic field to the spatial distribution of chemical signals in a developing embryo. [25] He demonstrated with a mathematical model that a variety of patterns could be formed by choosing parameter values to set up either static geometric patterns or dynamic oscillations, [22] [23] implying that the signalling system involved was somehow an alternative to natural selection. [15] Dawkins commented "He thinks he's anti-Darwinian, although he can't be, because he has no alternative explanation." [26]

Criticism

Laurence Moran notes that structuralism focuses mainly on animals, but that animals (emphasized) form only a small portion of the tree of life. CollapsedtreeLabels-simplified, Animals emphasized.svg
Laurence Moran notes that structuralism focuses mainly on animals, but that animals (emphasized) form only a small portion of the tree of life.

While agreeing that pattern formation mechanisms such as those described by Goodwin exist, the biologists Richard Dawkins, Stephen J. Gould, Lynn Margulis, and Steve Jones have criticised Goodwin for suggesting that chemical signalling forms an alternative to natural selection. [15]

Moran, a "skeptical biochemist", comments that 'structuralism' is a "new buzzword ... guaranteed to impress the creationist crowd because nobody understands what it means but it sounds very 'sciency' and philosophical." [15] The philosopher of science Paul E. Griffiths writes that structuralists "view this structuring of the space of biological possibility as part of the fundamental physical structure of nature. But the phenomena of phylogenetic inertia and developmental constraint do not support this interpretation. These phenomena show that the evolutionary pathways available to an organism are a function of the developmental structure of the organism." [27]

Moran summarizes: "There's nothing in science that supports the views of the structuralists. We have perfectly good explanations for why bumblebees are different than mushrooms and why all vertebrates have vertebrae and not exoskeletons. There's no evidence to support the idea that if you replay the tape of life it will come out looking anything like what we see today. You can be confident that when you visit another planet you will not find vertebrates." [15]

The evolutionary developmental biologist Lewis Held wrote that "The notion that aspects of anatomy can be explained by physical forces (like expansion cracking) was advocated ~ 100 years earlier in D'Arcy Thompson's 1917 On Growth and Form and in Theodore Cook's 1914 book The Curves of Life. [lower-alpha 4] Over the intervening century, various traits have been proposed to arise mechanically rather than genetically: brain convolutions, cartilage condensations, flower corrugations, tooth cusps, and fish otoliths. To this kooky list we can now add the crooked smile of the crocodile, or at least the cracked skin that surrounds it." [lower-alpha 5] [28]

See also

Notes

  1. ↑ Gregor Mendel pioneered the study of genetics.
  2. ↑ In this, Geoffroy's homologies were like Aristotle's forms.
  3. ↑ Universals are central to the ancient Greek theory, Platonic realism. [13]
  4. ↑ The artist Theodore Cook's The Curves of Life , Constable, 1914 to some extent anticipated D'Arcy Thompson, exploring spirals in art and nature.
  5. ↑ Held's final point was that the cracks in the crocodile's skin are genuinely explained by cracking, unlike all the other examples he lists. [28]

Related Research Articles

Evolutionary linguistics or Darwinian linguistics is a sociobiological approach to the study of language. Evolutionary linguists consider linguistics as a subfield of sociobiology and evolutionary psychology. The approach is also closely linked with evolutionary anthropology, cognitive linguistics and biolinguistics. Studying languages as the products of nature, it is interested in the biological origin and development of language. Evolutionary linguistics is contrasted with humanistic approaches, especially structural linguistics.

<span class="mw-page-title-main">Evolutionary developmental biology</span> Comparison of organism developmental processes

Evolutionary developmental biology is a field of biological research that compares the developmental processes of different organisms to infer how developmental processes evolved.

<span class="mw-page-title-main">Modern synthesis (20th century)</span> Fusion of natural selection with Mendelian inheritance

The modern synthesis was the early 20th-century synthesis of Charles Darwin's theory of evolution and Gregor Mendel's ideas on heredity into a joint mathematical framework. Julian Huxley coined the term in his 1942 book, Evolution: The Modern Synthesis.

<span class="mw-page-title-main">Homology (biology)</span> Shared ancestry between a pair of structures or genes in different taxa

In biology, homology is similarity due to shared ancestry between a pair of structures or genes in different taxa. A common example of homologous structures is the forelimbs of vertebrates, where the wings of bats and birds, the arms of primates, the front flippers of whales and the forelegs of four-legged vertebrates like dogs and crocodiles are all derived from the same ancestral tetrapod structure. Evolutionary biology explains homologous structures adapted to different purposes as the result of descent with modification from a common ancestor. The term was first applied to biology in a non-evolutionary context by the anatomist Richard Owen in 1843. Homology was later explained by Charles Darwin's theory of evolution in 1859, but had been observed before this, from Aristotle onwards, and it was explicitly analysed by Pierre Belon in 1555.

In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the population during that process. Thirdly, it is a phenotypic trait or adaptive trait, with a functional role in each individual organism, that is maintained and has evolved through natural selection.

<span class="mw-page-title-main">Brian Goodwin</span> Canadian mathematician and biologist

Brian Carey Goodwin was a Canadian mathematician and biologist, a Professor Emeritus at the Open University and a founder of theoretical biology and biomathematics. He introduced the use of complex systems and generative models in developmental biology. He suggested that a reductionist view of nature fails to explain complex features, controversially proposing the structuralist theory that morphogenetic fields might substitute for natural selection in driving evolution. He was also a visible member of the Third Culture movement.

Adaptationism is the Darwinian view that many physical and psychological traits of organisms are evolved adaptations. Pan-adaptationism is the strong form of this, deriving from the early 20th century modern synthesis, that all traits are adaptations, a view now shared by only a few biologists.

<span class="mw-page-title-main">Mutationism</span> One of several alternatives to evolution by natural selection.

Mutationism is one of several alternatives to evolution by natural selection that have existed both before and after the publication of Charles Darwin's 1859 book On the Origin of Species. In the theory, mutation was the source of novelty, creating new forms and new species, potentially instantaneously, in sudden jumps. This was envisaged as driving evolution, which was thought to be limited by the supply of mutations.

<i>Origination of Organismal Form</i>

Origination of Organismal Form: Beyond the Gene in Developmental and Evolutionary Biology is an anthology published in 2003 edited by Gerd B. Müller and Stuart A. Newman. The book is the outcome of the 4th Altenberg Workshop in Theoretical Biology on "Origins of Organismal Form: Beyond the Gene Paradigm", hosted in 1999 at the Konrad Lorenz Institute for Evolution and Cognition Research. It has been cited over 200 times and has a major influence on extended evolutionary synthesis research.

Modularity refers to the ability of a system to organize discrete, individual units that can overall increase the efficiency of network activity and, in a biological sense, facilitates selective forces upon the network. Modularity is observed in all model systems, and can be studied at nearly every scale of biological organization, from molecular interactions all the way up to the whole organism.

Cultural selection theory is the study of cultural change modelled on theories of evolutionary biology. Cultural selection theory has so far never been a separate discipline. However it has been proposed that human culture exhibits key Darwinian evolutionary properties, and "the structure of a science of cultural evolution should share fundamental features with the structure of the science of biological evolution". In addition to Darwin's work the term historically covers a diverse range of theories from both the sciences and the humanities including those of Lamark, politics and economics e.g. Bagehot, anthropology e.g. Edward B. Tylor, literature e.g. Ferdinand Brunetière, evolutionary ethics e.g. Leslie Stephen, sociology e.g. Albert Keller, anthropology e.g. Bronislaw Malinowski, Biosciences e.g. Alex Mesoudi, geography e.g. Richard Ormrod, sociobiology and biodiversity e.g. E.O. Wilson, computer programming e.g. Richard Brodie, and other fields e.g. Neoevolutionism, and Evolutionary archaeology.

<span class="mw-page-title-main">Spandrel (biology)</span> Evolutionary byproduct of some other characteristic

In evolutionary biology, a spandrel is a phenotypic trait that is a byproduct of the evolution of some other characteristic, rather than a direct product of adaptive selection. Stephen Jay Gould and Richard Lewontin brought the term into biology in their 1979 paper "The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme". Adaptationism is a point of view that sees most organismal traits as adaptive products of natural selection. Gould and Lewontin sought to temper what they saw as adaptationist bias by promoting a more structuralist view of evolution.

Developmental systems theory (DST) is an overarching theoretical perspective on biological development, heredity, and evolution. It emphasizes the shared contributions of genes, environment, and epigenetic factors on developmental processes. DST, unlike conventional scientific theories, is not directly used to help make predictions for testing experimental results; instead, it is seen as a collection of philosophical, psychological, and scientific models of development and evolution. As a whole, these models argue the inadequacy of the modern evolutionary synthesis on the roles of genes and natural selection as the principal explanation of living structures. Developmental systems theory embraces a large range of positions that expand biological explanations of organismal development and hold modern evolutionary theory as a misconception of the nature of living processes.

In evolutionary biology, function is the reason some object or process occurred in a system that evolved through natural selection. That reason is typically that it achieves some result, such as that chlorophyll helps to capture the energy of sunlight in photosynthesis. Hence, the organism that contains it is more likely to survive and reproduce, in other words the function increases the organism's fitness. A characteristic that assists in evolution is called an adaptation; other characteristics may be non-functional spandrels, though these in turn may later be co-opted by evolution to serve new functions.

Alessandro Minelli is an Italian biologist, formerly professor of zoology in the Faculty of Mathematical, Physical and Natural Sciences of the University of Padova mainly working on evo-devo subjects.

<span class="mw-page-title-main">Outline of evolution</span>

The following outline is provided as an overview of and topical guide to evolution:

The extended evolutionary synthesis consists of a set of theoretical concepts argued to be more comprehensive than the earlier modern synthesis of evolutionary biology that took place between 1918 and 1942. The extended evolutionary synthesis was called for in the 1950s by C. H. Waddington, argued for on the basis of punctuated equilibrium by Stephen Jay Gould and Niles Eldredge in the 1980s, and was reconceptualized in 2007 by Massimo Pigliucci and Gerd B. Müller. Notably, Dr. Müller concluded from this research that Natural Selection has no way of explaining speciation, saying: “selection has no innovative capacity...the generative and the ordering aspects of morphological evolution are thus absent from evolutionary theory.”

<span class="mw-page-title-main">Teleology in biology</span> Use of language of goal-directedness in the context of evolutionary adaptation

Teleology in biology is the use of the language of goal-directedness in accounts of evolutionary adaptation, which some biologists and philosophers of science find problematic. The term teleonomy has also been proposed. Before Darwin, organisms were seen as existing because God had designed and created them; their features such as eyes were taken by natural theology to have been made to enable them to carry out their functions, such as seeing. Evolutionary biologists often use similar teleological formulations that invoke purpose, but these imply natural selection rather than actual goals, whether conscious or not. Biologists and religious thinkers held that evolution itself was somehow goal-directed (orthogenesis), and in vitalist versions, driven by a purposeful life force. With evolution working by natural selection acting on inherited variation, the use of teleology in biology has attracted criticism, and attempts have been made to teach students to avoid teleological language.

<span class="mw-page-title-main">Alternatives to Darwinian evolution</span> List of alternatives to Darwinian natural selection

Alternatives to Darwinian evolution have been proposed by scholars investigating biology to explain signs of evolution and the relatedness of different groups of living things. The alternatives in question do not deny that evolutionary changes over time are the origin of the diversity of life, nor that the organisms alive today share a common ancestor from the distant past ; rather, they propose alternative mechanisms of evolutionary change over time, arguing against mutations acted on by natural selection as the most important driver of evolutionary change.

In evolutionary biology, developmental bias refers to the production against or towards certain ontogenetic trajectories which ultimately influence the direction and outcome of evolutionary change by affecting the rates, magnitudes, directions and limits of trait evolution. Historically, the term was synonymous with developmental constraint, however, the latter has been more recently interpreted as referring solely to the negative role of development in evolution.

References

  1. 1 2 3 Ruse, Michael (2013). "17. From Organicism to Mechanism-and Halfway Back?". In Henning, Brian G.; Scarfe, Adam (eds.). Beyond Mechanism: Putting Life Back Into Biology. Lexington Books. p. 419. ISBN   9780739174371.
  2. 1 2 3 4 5 6 7 Wagner, Günter P., Homology, Genes, and Evolutionary Innovation. Princeton University Press. 2014. ISBN   978-0691156460. Pages 7–38, 125
  3. ↑ Bowler, Peter J. (1989) [1983]. The Eclipse of Darwinism: anti-Darwinian evolutionary theories in the decades around 1900. Johns Hopkins University Press. pp. 261–262, 280–281. ISBN   978-0-8018-4391-4.
  4. ↑ Racine, Valerie (7 October 2013). "Essay: The Cuvier-Geoffroy Debate". The Embryo Project Encyclopedia, Arizona State University. Retrieved 10 December 2016.
  5. ↑ Leroi, Armand Marie (2014). The Lagoon: How Aristotle Invented Science. Bloomsbury. p. 13. ISBN   978-1-4088-3622-4.
  6. ↑ Ball, Philip (7 February 2013). "In retrospect: On Growth and Form". Nature. 494 (7435): 32–33. Bibcode:2013Natur.494...32B. doi: 10.1038/494032a . S2CID   205076253.
  7. ↑ Seilacher, Adolf (1991). "Self-Organizing Mechanisms in Morphogenesis and Evolution". In Schmidt-Kittler, Norbert; Vogel, Klaus (eds.). Constructional Morphology and Evolution. Springer. pp. 251–271. doi:10.1007/978-3-642-76156-0_17. ISBN   978-3-642-76158-4.
  8. ↑ Seilacher, Adolf (July 1989). "Vendozoa: Organismic construction in the Proterozoic biosphere". Lethaia. 22 (3): 229–239. doi:10.1111/j.1502-3931.1989.tb01332.x.
  9. ↑ Simpson, Carl; Erwin, Douglas H. (quoted) (13 April 2014). Homology, Genes, and Evolutionary Innovation Günter P. Wagner. Princeton University Press. ISBN   9780691156460 . Retrieved 9 December 2016.
  10. ↑ Brown, Rachael L. (November 2015). "Why development matters". Biology & Philosophy. 30 (6): 889–899. doi:10.1007/s10539-015-9488-9. S2CID   82602032.
  11. ↑ Muller, G. B.; Wagner, G. P. (1991). "Novelty in Evolution: Restructuring the Concept" (PDF). Annual Review of Ecology and Systematics. 22 (1): 229–256. doi:10.1146/annurev.es.22.110191.001305. ISSN   0066-4162. Archived (PDF) from the original on 26 September 2017.
  12. ↑ Fox, Ronald F. (December 1993). "Review of Stuart Kauffman, The Origins of Order: Self-Organization and Selection in Evolution". Biophys. J. 65 (6): 2698–2699. Bibcode:1993BpJ....65.2698F. doi:10.1016/s0006-3495(93)81321-3. PMC   1226010 .
  13. ↑ Silverman, Allan. "Plato's Middle Period Metaphysics and Epistemology". In Zalta, Edward N. (ed.). Stanford Encyclopedia of Philosophy .
  14. ↑ Denton, Michael J. (August 2013). "The Types: A Persistent Structuralist Challenge to Darwinian Pan-Selectionism". BIO-Complexity. 2013 (3). doi:10.5048/BIO-C.2013.3.
  15. 1 2 3 4 5 6 Moran, Laurence A. (2016-02-02). "What is "Structuralism"?". Sandwalk (blog of a recognised expert). Retrieved 9 December 2016.
  16. ↑ Stephen Jay Gould; Richard Lewontin (1979). "The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme". Proc. R. Soc. Lond. B. 205 (1161): 581–598. Bibcode:1979RSPSB.205..581G. doi:10.1098/rspb.1979.0086. PMID   42062. S2CID   2129408.
  17. 1 2 Erwin, Douglas H. (September 2011). "Evolutionary uniformitarianism". Developmental Biology. 357 (1): 27–34. doi: 10.1016/j.ydbio.2011.01.020 . PMID   21276788.
  18. ↑ Müller, Gerd B.; Newman, Stuart A. (2005). "The innovation triad: an EvoDevo agenda". J. Exp. Zool. (Mol. Dev Evol). 304B (6): 487–503. CiteSeerX   10.1.1.501.1440 . doi:10.1002/jez.b.21081. PMID   16299770.
  19. ↑ Newman, Stuart A.; Forgacs, Gabor; Müller, Gerd B. (2006). "Before programs: The physical origination of multicellular forms". The International Journal of Developmental Biology. 50 (2–3): 289–299. doi: 10.1387/ijdb.052049sn . PMID   16479496.
  20. ↑ King, Nicole (2004). "The Unicellular Ancestry of Animal Development". Developmental Cell. 7 (3): 313–325. doi: 10.1016/j.devcel.2004.08.010 . PMID   15363407.
  21. ↑ Goodwin, Brian (2009). Ruse, Michael; Travis, Joseph (eds.). Beyond the Darwinian Paradigm: Understanding Biological Forms. ISBN   978-0674062214.{{cite book}}: |work= ignored (help)
  22. 1 2 3 Price, Catherine S. C.; Goodwin, Brian (1995). "Structurally Unsound". Evolution. 49 (6): 1298. doi:10.2307/2410461. JSTOR   2410461.
  23. 1 2 Wake, David B. (1996). "How the Leopard Changed Its Spots: The Evolution of Complexity by Brian Goodwin". American Scientist . 84 (3): 300–301. JSTOR   29775684.
  24. ↑ Brian Goodwin obituary - The Guardian, 9 August 2009
  25. ↑ Dickinson, W. Joseph (1998). "Form and Transformation: Generative and Relational Principles in Biology. by Gerry Webster; Brian Goodwin". The Quarterly Review of Biology . 73 (1): 62–63. doi:10.1086/420070.
  26. ↑ Dawkins, Richard (1 May 1996). "Chapter 3 "A Survival Machine"". Edge. Retrieved 11 February 2018.
  27. ↑ Griffiths, Paul E. (1996). "Darwinism, Process Structuralism, and Natural Kinds". Philosophy of Science. 63: S1–S9. doi:10.1086/289930. JSTOR   188505. S2CID   146674266.
  28. 1 2 Held, Lewis Irving (2014). How the snake lost its legs: curious tales from the frontier of evo-devo. Cambridge, United Kingdom New York: Cambridge University Press. p. 121. ISBN   978-1-107-62139-8.
This page is based on this Wikipedia article
Text is available under the CC BY-SA 4.0 license; additional terms may apply.
Images, videos and audio are available under their respective licenses.