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Orthogenesis
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{{good article}}File:Haeckel arbol bn.png|thumb|right|250px|Evolutionary progress as a tree of life. Ernst HaeckelErnst HaeckelFile:Lamarck's Two-Factor Theory.svg|thumb|upright=2|Lamarck's two-factor theory involves 1) a complexifying force that drives animal body plans towards higher levels (orthogenesis) creating a ladder of phyla, and 2) an adaptive force that causes animals with a given body plan to adapt to circumstances (use and disuse, inheritance of acquired characteristics), creating a diversity of species and (genus|genera]]. Popular views of Lamarckism only consider an aspect of the adaptive force.BOOK, Gould, Stephen, The lying stones of Marrakech : penultimate reflections in natural history, Vintage, 2001, 978-0-09-928583-0, 119–121, )Orthogenesis, also known as orthogenetic evolution, progressive evolution, evolutionary progress, or progressionism, is the biological hypothesis that organisms have an innate tendency to evolve in a definite direction towards some goal (teleology) due to some internal mechanism or "driving force".{{sfn|Bowler|1989|pages=268–270}}BOOK, Ernst Mayr, Mayr, Ernst, 1988, Toward a New Philosophy of Biology: Observations of an Evolutionist, Harvard University Press, 499, 0-674-89666-1, {{sfn|Ruse|1996|pages=526–539}} According to the theory, the largest-scale trends in evolution have an absolute goal such as increasing biological complexity. Prominent historical figures who have championed some form of evolutionary progress include Jean-Baptiste Lamarck, Pierre Teilhard de Chardin, and Henri Bergson.The term orthogenesis was introduced by Wilhelm Haacke in 1893 and popularized by Theodor Eimer five years later. Proponents of orthogenesis had rejected the theory of natural selection as the organizing mechanism in evolution for a rectilinear model of directed evolution.JOURNAL, Ulett, Mark A., 2014, Making the case for orthogenesis: The popularization of definitely directed evolution (1890–1926), Studies in History and Philosophy of Biological and Biomedical Sciences, 45, 124–132, 10.1016/j.shpsc.2013.11.009, With the emergence of the modern synthesis, in which genetics was integrated with evolution, orthogenesis and other alternatives to Darwinism were largely abandoned by biologists, but the notion that evolution represents progress is still widely shared. The evolutionary biologist Ernst Mayr made the term effectively taboo in the journal Nature in 1948, by stating that it implied "some supernatural force".{{sfn|Ruse|1996|page=447}}Letter from Ernst Mayr to R. H. Flower, Evolution papers, 23 January 1948 The American paleontologist George Gaylord Simpson (1953) attacked orthogenesis, linking it with vitalism by describing it as "the mysterious inner force".BOOK, George Gaylord Simpson, Simpson, George Gaylord, 1953, Life of the Past: An Introduction to Paleontology, Yale University Press, 125, Modern supporters include E. O. Wilson and Simon Conway Morris, while museum displays and textbook illustrations continue to give the impression of progress in evolution.The philosopher of biology Michael Ruse notes that in popular culture evolution and progress are synonyms, while the unintentionally misleading image of the March of Progress, from apes to modern humans, has been widely imitated.

Definition

File:Theodor Eimer (Professorengalerie Universität Tübingen).jpg|200x200px|thumb|Theodor EimerTheodor Eimer{{further|Teleology in biology}}The term orthogenesis (from Ancient Greek ὀρθός orthós, "straight", and γένεσις génesis, "origin") was first used by the biologist Wilhelm Haacke in 1893.BOOK, Stephen Jay Gould, Gould, Stephen Jay, 2002, The Structure of Evolutionary Theory, Harvard University Press, 351–352, 978-0-674-00613-3, Theodor Eimer was the first to give the word a definition; he defined orthogenesis as "the general law according to which evolutionary development takes place in a noticeable direction, above all in specialized groups".BOOK, Lane, David H., 1996, The Phenomenon of Teilhard: Prophet for a New Age, Mercer University Press, 60–64, 0-86554-498-0, In 1922, the zoologist Michael F. Guyer wrote:}}According to Susan R. Schrepfer in 1983:}}In 1988, Francisco J. Ayala defined progress as "systematic change in a feature belonging to all the members of a sequence in such a way that posterior members of the sequence exhibit an improvement of that feature". He argued that there are two elements in this definition, directional change and improvement according to some standard. Whether a directional change constitutes an improvement is not a scientific question; therefore Ayala suggested that science should focus on the question of whether there is directional change, without regard to whether the change is "improvement".BOOK, Ayala, Francisco J., Francisco J. Ayala, 1988, Can progress be defined as a biological concept?, Evolutionary Progress, Nitecki, M., 75–96, University of Chicago Press, 0-226-58693-6, This may be compared to Stephen Jay Gould's suggestion of "replacing the idea of progress with an operational notion of directionality".BOOK, Gould, Stephen Jay, Stephen Jay Gould, 1997, Full House: The Spread of Excellence from Plato to Darwin, Harmony, 0-609-80140-6, In 1989, Peter J. Bowler defined orthogenesis as:}}In 1996, Michael Ruse defined orthogenesis as "the view that evolution has a kind of momentum of its own that carries organisms along certain tracks".{{sfn|Ruse|1996|page=261}}

History

File:Die Leiter des Auf- und Abstiegs.jpg|thumb|upright|The mediaeval great chain of being as a staircase, implying the possibility of progress:{{sfn|Ruse|1996|pp=21–23}} Ramon LullRamon Lull

Mediaeval

The possibility of progress is embedded in the mediaeval great chain of being, with a linear sequence of forms from lowest to highest. The concept, indeed, had its roots in Aristotle's biology, from insects that produced only a grub, to fish that laid eggs, and on up to animals with blood and live birth. The mediaeval chain, as in Ramon Lull's Ladder of Ascent and Descent of the Mind, 1305, added steps or levels above humans, with orders of angels reaching up to God at the top.{{sfn|Ruse|1996|pp=21–23}}

Pre-Darwinian

The orthogenesis hypothesis had a significant following in the 19th century when evolutionary mechanisms such as Lamarckism were being proposed. The French zoologist Jean-Baptiste Lamarck (1744–1829) himself accepted the idea, and it had a central role in his theory of inheritance of acquired characteristics, the hypothesized mechanism of which resembled the "mysterious inner force" of orthogenesis. Orthogenesis was particularly accepted by paleontologists who saw in their fossils a directional change, and in invertebrate paleontology thought there was a gradual and constant directional change. Those who accepted orthogenesis in this way, however, did not necessarily accept that the mechanism that drove orthogenesis was teleological (had a definite goal). Charles Darwin himself rarely used the term "evolution" now so commonly used to describe his theory, because the term was strongly associated with orthogenesis, as had been common usage since at least 1647.BOOK, Stephen Jay Gould, Gould, Stephen Jay, 1977,weblink Darwin's Dilemma: The Odyssey of Evolution, Ever Since Darwin: Reflections in Natural History, W. W. Norton, 0-393-06425-5, bot: unknown,weblink" title="web.archive.org/web/20051216025526weblink">weblink 2005-12-16,

With Darwin

File:Voyages_de_la_Commission_scientifique_du_Nord,_en_Scandinavie,_en_Laponie,_au_Spitzberg_et_aux_Feröe_-_no-nb_digibok_2009040211001-118.jpg|thumb|upright|Reviewing Darwin's Origin of Species, Karl Ernst von Baer argued for a directed force guiding evolutionevolutionRuse observed that "Progress (sic, his capitalisation) became essentially a nineteenth-century belief. It gave meaning to life—it offered inspiration—after the collapse [with Malthus's pessimism and the shock of the French revolution] of the foundations of the past."{{sfn|Ruse|1996|p=29}}The Russian biologist Karl Ernst von Baer (1792–1876) argued for an orthogenetic force in nature, reasoning in a review of Darwin's 1859 On the Origin of Species that "Forces which are not directed—so-called blind forces—can never produce order."BOOK, Brown, Keven, Von Kitzing, Eberhard, Evolution and Bahá'í Belief: ʻAbduʼl-Bahá's Response to Nineteenth-century Darwinism,weblink 2001, Kalimat Press, 978-1-890688-08-0, 159, BOOK, Barbieri, Marcello, 2013, Biosemiotics: Information, Codes and Signs in Living Systems, Nova Science Publishers, 7, 978-1-60021-612-1, BOOK, Jacobsen, Eric Paul, 2005, From Cosmology to Ecology: The Monist World-view in Germany from 1770 to 1930, 100, Peter Lang, 978-0-8204-7231-7, In 1864, the Swiss anatomist Albert von Kölliker (1817–1905) presented his orthogenetic theory, heterogenesis, arguing for wholly separate lines of descent with no common ancestor.BOOK, Vucinich, Alexander, 1988, Darwin in Russian Thought, University of California Press, 137, 978-0-520-06283-2, In 1884, the Swiss botanist Carl Nägeli (1817–1891) proposed a version of orthogenesis involving an "inner perfecting principle". Gregor Mendel died that same year; Nägeli, who proposed that an "idioplasm" transmitted inherited characteristics, dissuaded Mendel from continuing to work on plant genetics.BOOK, Mawer, Simon, Simon Mawer, Gregor Mendel: planting the seeds of genetics, 2006, Harry N. Abrams, 978-0-8109-5748-0, According to Nägeli many evolutionary developments were nonadaptive and variation was internally programmed. {{sfn|Bowler|1989|pages=268–270}} Charles Darwin saw this as a serious challenge, replying that "There must be some efficient cause for each slight individual difference", but was unable to provide a specific answer without knowledge of genetics. Further, Darwin was himself somewhat progressionist, believing for example that "Man" was "higher" than the barnacles he studied.BOOK, Watson, Marc, Angle, Barbara, Man's Selection: Charles Darwin's Theory of Creation, Evolution, And Intelligent Design,weblink 2017, BookBaby, 978-1-936883-14-1, 146–150, {{sfn|Ruse|1996|pp=154–155, 162}}Darwin indeed wrote in his 1859 Origin of Species:Darwin, C. (1859). On the Origin of Species By Means of Natural Selection, or, the Preservation of Favoured Races in the Struggle for Life. Chapters 10, 14File:Titanothere Osborn.jpg|thumb|Henry Fairfield Osborn's 1934 version of orthogenesis, aristogenesis, argued that aristogenes, not mutation or natural selection, created all novelty. Osborn supposed that the horns of Titanotheres evolved into a baroque form, way beyond the Ruse|1996|pages=266–267}}In 1898, after studying butterfly coloration, Theodor Eimer (1843–1898) introduced the term orthogenesis with a widely read book, On Orthogenesis: And the Impotence of Natural Selection in Species Formation. Eimer claimed there were trends in evolution with no adaptive significance that would be difficult to explain by natural selection.BOOK, Shanahan, Timothy, 2004, The Evolution of Darwinism: Selection, Adaptation, and Progress in Evolutionary Biology, Cambridge University Press, 121, 978-0-521-54198-5, To supporters of orthogenesis, in some cases species could be led by such trends to extinction.BOOK, Jan Sapp, Sapp, Jan, 2003, Genesis: The Evolution of Biology, 69–70, Oxford University Press, 978-0-19-515619-5, Eimer linked orthogenesis to neo-Lamarckism in his 1890 book Organic Evolution as the Result of the Inheritance of Acquired Characteristics According to the Laws of Organic Growth. He used examples such as the evolution of the horse to argue that evolution had proceeded in a regular single direction that was difficult to explain by random variation. Gould described Eimer as a materialist who rejected any vitalist or teleological approach to orthogenesis, arguing that Eimer's criticism of natural selection was common amongst many evolutionists of his generation; they were searching for alternative mechanisms, as they had come to believe that natural selection could not create new species.BOOK, Stephen Jay Gould, Gould, Stephen Jay, 2002, The Structure of Evolutionary Theory, Harvard University Press, 355–364, 978-0-674-00613-3,

Nineteenth and twentieth century

{{see|Eclipse of Darwinism}}Numerous versions of orthogenesis (see table) have been proposed. Debate centred on whether such theories were scientific, or whether orthogenesis was inherently vitalistic or essentially theological. For example, biologists such as Maynard M. Metcalf (1914), John Merle Coulter (1915), David Starr Jordan (1920) and Charles B. Lipman (1922) claimed evidence for orthogenesis in bacteria, fish populations and plants.JOURNAL, Metcalf Maynard M, 1913, Adaptation Through Natural Selection and Orthogenesis, The American Naturalist, 47, 554, 65–71, 2455865, 10.1086/279329, John Merle Coulter. (1915). A Suggested Explanation of 'Orthogenesis' in Plants Science, Vol. 42, No. 1094. pp. 859–863.JOURNAL, Starr Jordan David, David Starr Jordan, 1920, Orthogenesis among Fishes, 1646251, Science, 52, 1331, 13–14, 10.1126/science.52.1331.13-a, JOURNAL, Lipman Charles B, 1922, Orthogenesis in Bacteria, The American Naturalist, 56, 643, 105–115, 2456503, 10.1086/279851, In 1950, the German paleontologist Otto Schindewolf argued that variation tends to move in a predetermined direction. He believed this was purely mechanistic, denying any kind of vitalism, but that evolution occurs due to a periodic cycle of evolutionary processes dictated by factors internal to the organism.BOOK, Kwa, Chunglin, 2011, Styles of Knowing: A New History of Science from Ancient Times to the Present, University of Pittsburgh Press, 237, 978-0-8229-6151-2, JOURNAL, Dimichele William A, 1995, Basic Questions in Paleontology: Geologic Time, Organic Evolution, and Biological Systematics, by Otto H. Schindewolf,weblink PDF, Review of Palaeobotany and Palynology, 84, 481–483, 10.1016/0034-6667(95)90007-1, In 1964 George Gaylord Simpson argued that orthogenetic theories such as those promulgated by Du Noüy and Sinnott were essentially theology rather than biology.BOOK, Simpson, George Gaylord, George Gaylord Simpson, 1964, "Evolutionary Theology: The New Mysticism", This View of Life: The World of an Evolutionist, Harcourt, Brace & World, 213–233, Though evolution is not progressive, it does sometimes proceed in a linear way, reinforcing characteristics in certain lineages, but such examples are entirely consistent with the modern neo-Darwinian theory of evolution.JOURNAL, Jepsen, Glenn L., 1949, Selection. Orthogenesis, and the Fossil Record, Proceedings of the American Philosophical Society, 93, 6, 479–500, These examples have sometimes been referred to as orthoselection but are not strictly orthogenetic, and simply appear as linear and constant changes because of environmental and molecular constraints on the direction of change.JOURNAL, Jacobs, Susan C., Larson, Allan, Cheverud, James M., 1995, Phylogenetic Relationships and Orthogenetic Evolution of Coat Color Among Tamarins (Genus Saguinus), Syst. Biol., 44, 4, 515–532, 10.1093/sysbio/44.4.515, JOURNAL, Ranganath, H. A., Hägel, K., 1981, Karyotypic orthoselection in Drosophila, Naturwissenschaften, 68, 10, 527–528, 10.1007/bf00365385, 1981NW.....68..527R, The term orthoselection was first used by Ludwig Hermann Plate, and was incorporated into the modern synthesis by Julian Huxley and Bernard Rensch.JOURNAL, Levit, Georgy S., Olsson, Lennart, 'Evolution on Rails' : Mechanisms and Levels of Orthogenesis, Annals of the History and Philosophy of Biology, 11, 2006, 99–138,weblink Recent work has supported the mechanism and existence of mutation-biased adaptation, meaning that limited local orthogenesis is now seen as possible.JOURNAL, Yampolsky, L. Y., Stoltzfus, A., 2001, Bias in the introduction of variation as an orienting factor in evolution, Evol Dev, 3, 73–83, 10.1046/j.1525-142x.2001.003002073.x, 11341676, 2, BOOK, Stoltzfus, A., Mutation-Biased Adaptation in a Protein NK Model, Mol Biol Evol, 23, 1852–1862, 2006, JOURNAL, Stoltzfus, A, Yampolsky, L. Y., 2009, Climbing Mount Probable: Mutation as a Cause of Nonrandomness in Evolution, J Hered, 100, 637–647, 10.1093/jhered/esp048, 19625453, 5, {{-}}

Theories

{{further|Alternatives to evolution by natural selection}}For the columns for other philosophies of evolution (i.e., combined theories including any of Lamarckism, Mutationism, Natural selection, and Vitalism), "yes" means that person definitely supports the theory; "no" means explicit opposition to the theory; a blank means the matter is apparently not discussed, not part of the theory.{| class="wikitable sortable"evolutionary biologyCIGNA, ARRIGO A., Durante, Marco, Radiation Risk Estimates in Normal and Emergency Situations,weblink 2007, Springer Science & Business Media, 978-1-4020-4956-9, 213, ! Author !! Title !! Field !! Date !! Lamarck. !! Mutat. !! Nat. Sel. !! Vital. !! Features
Jean-Baptiste Lamarck>Lamarck Inherent progressive tendency Zoology 1809 yes In his Philosophie Zoologique, inherent progressive tendency drives organisms continuously towards greater Evolution of biological complexity, in separate lineages (phylum>phyla), no extinction. ("Lamarckism", use and disuse, and inheritance of acquired characteristics, was a secondary aspect of this, an adaptive force creating species within a phylum.)
Karl Ernst von Baer>Baer Purposeful creation Embryology 1859 "Forces which are not directed—so-called blind forces—can never produce order."
Albert von Kölliker>Kölliker Heterogenesis Anatomy 1864 yes Wholly separate lines of descent with no common ancestor
Edward Drinker Cope>Cope Law of acceleration Palaeontology 1868 yes Combined orthogenetic constraints with Lamarckian use and disuse. "On the Origin of Genera";BARNES >FIRST1=M. ELIZABETH URL=HTTPS://EMBRYO.ASU.EDU/PAGES/EDWARD-DRINKER-COPES-LAW-ACCELERATION-GROWTH, 24 July 2014, See also Cope's rule (linear increase in size of species)
Carl Nägeli>Nägeli Inner perfecting principle Botany 1884 yes no An "idioplasm" transmitted inherited characteristics; many evolutionary developments nonadaptive; variation internally programmed.{{sfn1989|pages=268–270}}
Herbert Spencer>Spencer Progressionism'The Development Hypothesis' Social theory 1852 Yes{{sfn1996Michael Ruse{{sfn>Rusepages=181–191}}
Charles Darwin>Darwin (concept of higher and lower species), Pangenesis Evolution 1859 yes yes Origin of Species is somewhat progressionist, e.g. man higher than animals, alongside natural selection Pangenesis theory of inheritance by gemmules from all over body was Lamarckism: parents could inheritance of acquired characters>pass on traits acquired in lifetime.GHISELIN>FIRST = MICHAEL T.PUBLICATION-DATE = SEPTEMBER–OCTOBER 1994CONTRIBUTION-URL =HTTP://WWW.TEXTBOOKLEAGUE.ORG/54MARCK.HTMPUBLISHER =THE TEXTBOOK LEAGUEACCESSDATE=2008-01-23, MAGNER >FIRST = LOIS N. EDITION = THIRD MARCEL DEKKER, CRC PRESS >YEAR = 2002 URL = HTTPS://BOOKS.GOOGLE.COM/?ID=YKJ6GVYBRGWC&PRINTSEC=FRONTCOVER#V=ONEPAGE&Q,
Wilhelm Haacke>Haacke Orthogenesis Zoology 1893 yes Accompanied by epimorphism, a tendency to increasing perfection
Theodor Eimer>Eimer Orthogenesis Zoology 1898 no On Orthogenesis: And the Impotence of Natural Selection in Species Formation: trends in evolution with no adaptive significance, claimed hard to explain by natural selection.
Henri Bergson>Bergson ''Élan vital >Creative Evolution (book)>Creative Evolution''{{sfn1989|pages=116–117}}
Hans Przibram>Przibram Apogenesis Embryology 1910s
Ludwig Hermann Plate>Plate Orthoselection or Old-Darwinism Zoology 1913 yes yes yes Combined theory
Daniele Rosa>Rosa Hologenesis Zoology 1918 yes Hologenesis: a New Theory of Evolution and the Geographical Distribution of Living BeingsLUZZATTO, MICHELE>AUTHOR2=PALESTRINI, CLAUDIADATE=2000 JOURNAL=ITALIAN JOURNAL OF ZOOLOGY PAGES=129–138, 10.1080/11250000009356303,
Charles Otis Whitman>Whitman Orthogenesis Zoology 1919 no no no Orthogenetic Evolution in Pigeons posthumousCASTLE >FIRST1=W.E. TITLE=REVIEW OF ORTHOGENETIC EVOLUTION IN PIGEONS VOLUME=54 PAGES=188–192 DATE=2002 PUBLISHER=HARVARD UNIVERSITY PRESS ISBN=978-0-674-00613-3,
Lev Berg>Berg Nomogenesis Zoology 1926 no yes no Chemical forces direct evolution, leading to humans{{sfn1996Bowlerp=157}}
Othenio Abel>Abel Trägheitsgesetz (the law of inertia) Palaeontology 1928 based on Dollo's law of irreversibility of evolution (which can be explained without orthogenesis as a statistical improbability that a path should be exactly reversed)
André Michel Lwoff>Lwoff Physiological degradation Physiology 1930s–1940s yes Directed loss of functions in microorganismsLWOFF, A. >AUTHORLINK=ANDRé MICHEL LWOFF LOCATION=PARIS DATE=1944 QUOTE=L'IDéE S'IMPOSA QUE LES MICROORGANISMES AVAIENT SUBI DES PERTES DE FONCTION. CELLES-CI APPARURENT COMME LA MANIFESTATION D'UNE éVOLUTION PHYSIOLOGIQUE, DEFINIE COMME UNE DEGRADATION, UNE ORTHOGENESE REGRESSIVE., LOISON >FIRST1=LAURENT FIRST2=JEAN FIRST3=RICHARD M. JOURNAL=JOURNAL OF THE HISTORY OF BIOLOGY VOLUME=50 PAGES=5–52 URL=HTTPS://LINK.SPRINGER.COM/ARTICLE/10.1007/S10739-015-9434-3,
Karl Beurlen>Beurlen Orthogenesis Palaeontology 1930 no no Start is random metakinesis, generating variety; then palingenesis (in Beurlen's sense, repeating developmental pathway of ancestors) as mechanism for orthogenesis
Victor Jollos| Combined orthogenesis with Lamarckism (inheriting acquired characteristics after heat shock as dauermodifications, passed on by plasmatic inheritance in the cytoplasm)
Henry Fairfield Osborn>Osborn Aristogenesis Palaeontology 1934 yes no no WALLACE, DAVID RAINS >DATE=2005 PUBLISHER=UNIVERSITY OF CALIFORNIA PRESS ISBN=978-0-520-24684-3, REGAL, BRIAN >DATE=2002 PUBLISHER=ASHGATE ISBN=978-0-7546-0587-4,
John Christopher Willis>Willis Differentiation (orthogenesis) Botany 1942 yes a force "working upon some definite law that we do not yet comprehend", compromise between special creation and natural selection, driven by large mutations involving chromosome alterationsThe Course of Evolution by J. C. Willis. Review by: Carl L. Hubbs. The American Naturalist, Vol. 76, No. 762 (Jan. Feb., 1942), pp. 96–101.
Pierre Lecomte du Noüy>Noüy Telefinalism Biophysics 1947 yes In book Human Destiny, essentially religiousKOCH, LEO FRANCIS >DATE=1957 JOURNAL=THE SCIENTIFIC MONTHLY ISSUE=5 BIBCODE=1957SCIMO..85..245K,
Albert Vandellt=Vandel}} Organicism Zoology 1949 L'Homme et L'Evolution
Edmund Ware Sinnott>Sinnott Telism Botany 1950 yes In book Cell and Psyche, essentially religious
Otto Schindewolf>Schindewolf Typostrophism Palaeontology 1950 yes Basic Questions in Paleontology: Geologic Time, Organic Evolution and Biological Systematics; evolution due to periodic cycle of processes dictated by factors internal to organism.
Pierre Teilhard de Chardin>Teilhard de Chardin Directed additivity,NoogenesisOmega principle PalaeontologyMysticism 1959 yes The Phenomenon of Man posthumous; combined orthogenesis with non-material vitalist directive force aiming for a supposed "Omega Point" with creation of consciousness. Noosphere concept from Vladimir Vernadsky. Censured by Gaylord Simpson for nonscientific spiritualistic "doubletalk".Chardin, Pierre Teilhard de. (2003) [1959] The Human Phenomenon. Sussex Academic Press. p. 65. {{ISBNTITLE=MARXIST WRITINGS ON HISTORY & PHILOSOPHY YEAR=2002 ISBN=978-1-876646-23-3, 207,
Léon Croizat>Croizat ''Biological synthesis (philosophy)Panbiogeography'' >developmental constraints or phylogenetic constraintsGRAY TITLE=OPPOSITIONS IN PANBIOGEOGRAPHY: CAN THE CONFLICTS BETWEEN SELECTION, CONSTRAINT, ECOLOGY, AND HISTORY BE RESOLVED? DATE=1989 PAGES=787–806 DOI=10.1080/03014223.1989.10422935,
File:Alternatives to Darwinism.svg|thumb|upright=1.5|Multiple explanations have been offered since the 19th century for how evolution took place, given that many scientists initially had objections to natural selection. Many of these theories led to some form of orthogenesis, with or without invoking divine control directly or indirectly. For example, evolutionists like Bowler|1989|pp=261-262}} represented by a sequence of arrows on the left of the diagram.The various alternatives to Darwinian evolution by natural selection were not necessarily mutually exclusive. The evolutionary philosophy of the American palaeontologist Edward Drinker Cope is a case in point. Cope, a religious man, began his career denying the possibility of evolution. In the 1860s, he accepted that evolution could occur, but, influenced by Agassiz, rejected natural selection. Cope accepted instead the theory of recapitulation of evolutionary history during the growth of the embryo - that ontogeny recapitulates phylogeny, which Agassiz believed showed a divine plan leading straight up to man, in a pattern revealed both in embryology and palaeontology. Cope did not go so far, seeing that evolution created a branching tree of forms, as Darwin had suggested. Each evolutionary step was however non-random: the direction was determined in advance and had a regular pattern (orthogenesis), and steps were not adaptive but part of a divine plan (theistic evolution). This left unanswered the question of why each step should occur, and Cope switched his theory to accommodate functional adaptation for each change. Still rejecting natural selection as the cause of adaptation, Cope turned to Lamarckism to provide the force guiding evolution. Finally, Cope supposed that Lamarckian use and disuse operated by causing a vitalist growth-force substance, "bathmism", to be concentrated in the areas of the body being most intensively used; in turn, it made these areas develop at the expense of the rest. Cope's complex set of beliefs thus assembled five evolutionary philosophies: recapitulationism, orthogenesis, theistic evolution, Lamarckism, and vitalism.{{sfn|Bowler|1989|pp=261-262}} Other palaeontologists and field naturalists continued to hold beliefs combining orthogenesis and Lamarckism until the modern synthesis in the 1930s.{{sfn|Bowler|1989|p=264}}

Status

In science

File:Human pidegree.jpg|thumb|left|upright=0.9|A satirical opinion of Ernst Haeckel's 1874 The modern theory of the descent of man, showing a linear sequence of forms leading up to 'Man'. Illustration by G. Avery for Scientific AmericanScientific AmericanThe stronger versions of the orthogenetic hypothesis began to lose popularity when it became clear that they were inconsistent with the patterns found by paleontologists in the fossil record, which were non-rectilinear (richly branching) with many complications. The hypothesis was abandoned by the mainstream of evolutionists when no mechanism could be found that would account for the process, and the theory of evolution by natural selection came to prevail.BOOK, Ernst Mayr, Mayr, Ernst, 1982, The Growth of Biological Thought: Diversity, Evolution, and Inheritance, Harvard University Press, 530–531, 0-674-36446-5, The historian of biology Edward J. Larson commented that}}The modern synthesis of the 1930s and 1940s, in which the genetic mechanisms of evolution were incorporated, appeared to refute the hypothesis for good. As more was understood about these mechanisms it came to be held that there was no naturalistic way in which the newly discovered mechanism of heredity could be far-sighted or have a memory of past trends. Orthogenesis was seen to lie outside the methodological naturalism of the sciences.BOOK, Stephen Jay Gould, Gould, Stephen Jay, The Structure of Evolutionary Theory, 2002, Chapter 7, section "Synthesis as Restriction", 0-674-00613-5, BOOK, Levinton, Jeffrey S., 2001, Genetics, Paleontology, and Macroevolution, Cambridge University Press, 14–16, 0-521-80317-9, BOOK, Montgomery, Georgina M., Largent, Mark A., 2015, A Companion to the History of American Science, Wiley, 218, 978-1-4051-5625-7, With the integration of Mendelian genetics and population genetics into evolutionary theory in the 1930s a new generation of biologists applied mathematical techniques to investigate how changes in the frequency of genes in populations combined with natural selection could produce species change. This demonstrated that Darwinian natural selection was the primary mechanism for evolution and that other models of evolution, such as neo-Lamarckism and orthogenesis, were invalid., File:Ernst Mayr PLoS.jpg|thumb|right|Ruse|1996|page=447}}By 1948, the evolutionary biologist Ernst Mayr, as editor of the journal Evolution, made the use of the term orthogenesis taboo: "It might be well to abstain from use of the word 'orthogenesis' .. since so many of the geneticists seem to be of the opinion that the use of the term implies some supernatural force."{{sfn|Ruse|1996|page=447}} With the rise of evolutionary developmental biology in the late 20th-early 21st centuries, however, which is open to an expanded concept of heredity that incorporates the physics of self-organization, ideas of constraint and preferred directions of morphological change have made a reappearance in evolutionary theory.see, for example, BOOK, Gerd Müller (theoretical biologist), Müller, Gerd B., Stuart Newman, Newman, Stuart A., Origination of Organismal Form, 2003, Bradford, 0-262-13419-5, For these and other reasons, belief in evolutionary progress has remained "a persistent heresy",WEB, Popov, Igor, The Persistence of Heresy: The Concepts of Directed Evolution (Orthogenesis),weblink 15 April 2017, 7 April 2005, among evolutionary biologists including E. O. WilsonWEB, Ruse, Michael, Michael Ruse, Edward O. Wilson on Sociobiology,weblink The Chronicle of Higher Education, 4 April 2017, 31 March 2010, and Simon Conway Morris, although often denied or veiled. The philosopher of biology Michael Ruse wrote that "some of the most significant of today's evolutionists are progressionists, and that because of this we find (absolute) progressionism alive and well in their work."{{sfn|Ruse|1996|p=536}} He argued that progressionism has harmed the status of evolutionary biology as a mature, professional science.{{sfn|Ruse|1996|p=530}} Presentations of evolution remain characteristically progressionist, with humans at the top of the "Tower of Time" in the Smithsonian Institution in Washington D.C., while Scientific American magazine could illustrate the history of life leading progressively from mammals to dinosaurs to primates and finally man. Ruse noted that at the popular level, progress and evolution are simply synonyms, as they were in the nineteenth century, though confidence in the value of cultural and technological progress has declined.{{sfn|Ruse|1996|pages=526–539}}

In popular culture

File:Huxley - Mans Place in Nature.jpg|thumb|left|upright=1.3|The frontispiece to Thomas Henry Huxley's 1863 Evidence as to Man's Place in Nature was intended to compare the skeletons of apes and humans, but unintentionally created a durable meme of supposed "monkey-to-man" progress.NEWS, Tucker, Jennifer, What our most famous evolutionary cartoon gets wrong,weblink 29 December 2017, The Boston GlobeBoston Globe{{further|March of Progress}}In popular culture, progressionist images of evolution are widespread. The historian Jennifer Tucker, writing in The Boston Globe, notes that Thomas Henry Huxley's 1863 illustration comparing the skeletons of apes and humans "has become an iconic and instantly recognizable visual shorthand for evolution." She calls its history extraordinary, saying that it is "one of the most intriguing, and most misleading, drawings in the modern history of science." Nobody, Tucker observes, supposes that the "monkey-to-man" sequence accurately depicts Darwinian evolution. The Origin of Species had only one illustration, a diagrams showing that random events create a process of branching evolution, a view that Tucker notes is broadly acceptable to modern biologists. But Huxley's image recalled the great chain of being, implying with the force of a visual image a "logical, evenly paced progression" leading up to Homo sapiens, a view denounced by Stephen Jay Gould in Wonderful Life.File:Man is But a Worm.jpg|thumb|upright|Man is But a Worm by Edward Linley SambourneEdward Linley SambournePopular perception, however, had seized upon the idea of linear progress. Edward Linley Sambourne's Man is But a Worm, drawn for Punch's Almanack, mocked the idea of any evolutionary link between humans and animals, with a sequence from chaos to earthworm to apes, primitive men, a Victorian beau, and Darwin in a pose that according to Tucker recalls Michelangelo's figure of Adam in his fresco adorning the ceiling of the Sistine Chapel. This was followed by a flood of variations on the evolution-as-progress theme, including The New Yorker's 1925 "The Rise and Fall of Man", the sequence running from a chimpanzee to Neanderthal man, Socrates, and finally the lawyer William Jennings Bryan who argued for the anti-evolutionist prosecution in the Scopes Trial on the State of Tennessee law limiting the teaching of evolution. Tucker noted that Rudolph Franz Zallinger's 1965 "The Road to Homo Sapiens" fold-out illustration in F. Clark Howell's Early Man, showing a sequence of 14 walking figures ending with modern man, fitted the palaeoanthropological discoveries "not into a branching Darwinian scheme, but into the framework of the original Huxley diagram." Howell ruefully commented that the "powerful and emotional" graphic had overwhelmed his Darwinian text.File:Astronomy Evolution 2 (27458655072).jpg|thumb|right|upright=1.3|One of many versions of the progressionist (meme]]: Astronomy Evolution 2 artwork by Giuseppe Donatiello, 2016)

Sliding between meanings

Scientists, Ruse argues, continue to slide easily from one notion of progress to another: even committed Darwinians like Richard Dawkins embed the idea of cultural progress in a theory of cultural units, memes, that act much like genes.{{sfn|Ruse|1996|pages=526–539}} Dawkins can speak of "progressive rather than random ... trends in evolution".JOURNAL, Dawkins, Richard, Krebs, J. R., Richard Dawkins, John Krebs, Baron Krebs, Arms races between and within species, 1979, Proceedings of the Royal Society B, 205, 489–511, 10.1098/rspb.1979.0081, 1979RSPSB.205..489D, {{sfn|Ruse|1996|page=466}} Dawkins and John Krebs deny the "earlier [Darwinian] prejudice"{{sfn|Ruse|1996|page=468}} that there is anything "inherently progressive about evolution",{{sfn|Dawkins|1986|page=178}}{{sfn|Ruse|1996|page=468}} but the feeling of progress comes from evolutionary arms races which remain in Dawkins's words "by far the most satisfactory explanation for the existence of the advanced and complex machinery that animals and plants possess".{{sfn|Dawkins|1986|page=181}}{{sfn|Ruse|1996|page=468}}Ruse concludes his detailed analysis of the idea of Progress, meaning a progressionist philosophy, in evolutionary biology by stating that evolutionary thought came out of that philosophy. Before Darwin, Ruse argues, evolution was just a pseudoscience; Darwin made it respectable, but "only as popular science". "There it remained frozen, for nearly another hundred years",{{sfn|Ruse|1996|pages=526–539}} until mathematicians such as Fisher{{sfn|Ruse|1996|pages=292–295}} provided "both models and status", enabling evolutionary biologists to construct the modern synthesis of the 1930s and 1940s. That made biology a professional science, at the price of ejecting the notion of progress. That, Ruse argues, was a significant cost to "people [biologists] still firmly committed to Progress" as a philosophy.{{sfn|Ruse|1996|pages=526–539}}

Facilitated variation

{{multiple image | footer=Different species of Heliconius butterfly have independently evolved similar patterns, apparently both facilitated and constrained by the available developmental-genetic toolkit genes controlling wing pattern formation. | align=right | image1=Heliconius erato Richard Bartz.jpg | width1=125 | caption1=Heliconius erato | image2=Heliconius melpomene 2b Richard Bartz.jpg | width2=175 | caption2=Heliconius melpomene}}Biology has largely rejected the idea that evolution is guided in any way,{{sfn|Bowler|1989|page=270}}{{sfn|Larson|2004|page=127}} but the evolution of some features is indeed facilitated by the genes of the developmental-genetic toolkit studied in evolutionary developmental biology. An example is the development of wing pattern in some species of Heliconius butterfly, which have independently evolved similar patterns. These butterflies are Müllerian mimics of each other, so natural selection is the driving force, but their wing patterns, which arose in separate evolutionary events, are controlled by the same genes.JOURNAL, Baxter, S.W., Papa, R., Chamberlain, N., Humphray, S.J., Joron, M., Morrison, C., ffrench-Constant, R.H., McMillan, W.O., Jiggins, C.D., 2008, Convergent Evolution in the Genetic Basis of Mullerian Mimicry in Heliconius Butterflies, Genetics, 180, 3, 1567–1577, 18791259, 2581958, 10.1534/genetics.107.082982,

See also

References

{{reflist|30em}}

Sources

  • BOOK, Bowler, Peter J., Peter J. Bowler, The Eclipse of Darwinism: anti-Darwinian evolutionary theories in the decades around 1900, Johns Hopkins University Press, 1983, 978-0-8018-4391-4, harv,
  • BOOK, Bowler, Peter J., Peter J. Bowler, 1989, Evolution: The History of an Idea, University of California Press, 978-0-520-06385-3, harv,
  • BOOK, Dawkins, Richard, Richard Dawkins, The Blind Watchmaker, 1986, Longman, 978-0-393-31570-7, harv,
  • BOOK, Larson, Edward J., Evolution, 2004, Modern Library, 978-0-679-64288-6, harv,
  • BOOK, Ruse, Michael, Michael Ruse, 1996, Monad to man: the Concept of Progress in Evolutionary Biology, Harvard University Press, 978-0-674-03248-4, harv,

Further reading

{{Evolution}}

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