Editing Orthogenesis

{{short description|Hypothesis that organisms have an innate tendency to evolve towards some goal}}
{{good article}}
[[File:Haeckel arbol bn.png|thumb|Evolutionary progress as a [[tree of life (biology)|tree of life]]. [[Ernst Haeckel]], 1866]]
[[File:Lamarck's Two-Factor Theory.svg|thumb|upright=1.4|Lamarck's two-factor theory involves 1) a complexifying force that drives animal [[body plan]]s towards higher levels (orthogenesis) creating a ladder of [[phylum|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.<ref name="Gould 2001">{{cite book | last=Gould | first=Stephen J. |author-link=Stephen Jay Gould | title=The lying stones of Marrakech : penultimate reflections in natural history | publisher=Vintage | year=2001 | isbn=978-0-09-928583-0 | pages=119–121}}</ref>]]
'''Orthogenesis''', also known as '''orthogenetic evolution''', '''progressive evolution''', '''evolutionary progress''', or '''progressionism''', is an [[Superseded theories in science|<!--Please do not attempt to remove this, it is well attested (see following citations) and you can be blocked for inappropriate editing-->obsolete]] biological [[hypothesis]] that [[organism]]s have an innate tendency to [[evolution|evolve]] in a definite direction [[teleology|towards some goal (teleology)]] due to some internal mechanism or "driving force".{{sfn|Bowler|1989|pages=268–270}}<ref>{{cite book |last=Mayr |first=Ernst |author-link=Ernst Mayr |date=1988 |title=Toward a New Philosophy of Biology: Observations of an Evolutionist |publisher=Harvard University Press |page=499 |isbn=978-0-674-89666-6}}</ref>{{sfn|Ruse|1996|pages=526–539}} According to the theory, the [[largest-scale trends in evolution]] have an absolute goal such as [[Evolution of biological complexity|increasing biological complexity]]. Prominent historical figures who have championed some form of evolutionary progress include [[Jean-Baptiste Lamarck]], [[Teilhard de Chardin|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 (straight-line) model of directed evolution.<ref>{{cite journal |last=Ulett |first=Mark A. |date=2014 |title=Making the case for orthogenesis: The popularization of definitely directed evolution (1890–1926) |journal=Studies in History and Philosophy of Biological and Biomedical Sciences |volume=45 |pages=124–132 |doi=10.1016/j.shpsc.2013.11.009 |pmid=24368232 }}</ref> With the emergence of the [[Modern synthesis (20th century)|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; modern supporters include [[E. O. Wilson]] and [[Simon Conway Morris]]. The evolutionary biologist [[Ernst Mayr]] made the term effectively taboo in the journal ''[[Nature (journal)|Nature]]'' in 1948, by stating that it implied "some supernatural force".{{sfn|Ruse|1996|page=447}}<ref name=Mayr1948>Letter from [[Ernst Mayr]] to R. H. Flower, ''Evolution'' papers, 23 January 1948</ref> The American paleontologist [[George Gaylord Simpson]] (1953) attacked orthogenesis, linking it with [[vitalism]] by describing it as "the mysterious inner force".<ref>{{cite book |last=Simpson |first=George Gaylord |author-link=George Gaylord Simpson |date=1953 |title=Life of the Past: An Introduction to Paleontology |url=https://archive.org/details/lifeofpastintrod00simp |url-access=registration |publisher=Yale University Press |page=[https://archive.org/details/lifeofpastintrod00simp/page/125 125]}}</ref> Despite this, many museum displays and textbook illustrations continue to give the impression that evolution is directed. 

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|thumb|upright|[[Theodor Eimer]]]]
{{further|Teleology in biology}}

The term orthogenesis (from Ancient {{langx|el|ὀρθός}} orthós, "straight", and Ancient {{langx|el|γένεσις génesis}}, "origin") was first used by the biologist [[Wilhelm Haacke]] in 1893.<ref name=LevitOlsson2006/><ref name="Gould 2002">{{cite book |last=Gould |first=Stephen Jay |author-link=Stephen Jay Gould |date=2002 |title=The Structure of Evolutionary Theory |publisher=Harvard University Press |pages=[https://archive.org/details/structureofevolu00goul/page/351 351–352] |isbn=978-0-674-00613-3 |title-link=The Structure of Evolutionary Theory }}</ref> [[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".<ref name="Lane 1996">{{cite book |last=Lane |first=David H. |date=1996 |title=The Phenomenon of Teilhard: Prophet for a New Age |publisher=Mercer University Press |pages=60–64 |isbn=978-0-86554-498-7}}</ref>

In 1922, the zoologist [[Michael F. Guyer]] wrote:

{{quote|[Orthogenesis] has meant many different things to many different people, ranging from a mystical [[inner perfecting principle]], to merely a general trend in development due to the natural constitutional restrictions of the germinal materials, or to the physical limitations imposed by a narrow environment. In most modern statements of the theory, the idea of continuous and progressive change in one or more characters, due according to some to internal factors, according to others to external causes-evolution in a "straight line" seems to be the central idea.<ref>{{cite journal |last=Guyer |first=Michael F. |date=1922 |jstor=2456504 |title=Orthogenesis and Serological Phenomena |journal=The American Naturalist |volume=56 |issue=643 |pages=116–133 |doi=10.1086/279852|doi-access= }}</ref>}}

According to [[Susan R. Schrepfer]] in 1983:

{{quote|Orthogenesis meant literally "straight origins", or "straight line evolution". The term varied in meaning from the overtly vitalistic and theological to the mechanical. It ranged from theories of mystical forces to mere descriptions of a general trend in development due to natural limitations of either the germinal material or the environment&nbsp;... By 1910, however most who subscribed to orthogenesis hypothesized some physical rather than metaphysical determinant of orderly change.<ref>{{cite book |last=Schrepfer |first=Susan R. |author-link=Susan Schrepfer |date=1983 |title=Fight to Save the Redwoods: A History of the Environmental Reform, 1917–1978 |publisher=University of Wisconsin Press |pages=81–82 |isbn=978-0-299-08854-5}}</ref>}}

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".<ref name=":1">{{cite book |last=Ayala |first=Francisco J. |author-link=Francisco J. Ayala |year=1988 |chapter=Can progress be defined as a biological concept? |title=Evolutionary Progress |editor=Nitecki, M. |pages=75–96 |publisher=University of Chicago Press |isbn=978-0-226-58693-9}}</ref> This may be compared to [[Stephen Jay Gould]]'s suggestion of "replacing the idea of progress with an operational notion of directionality".<ref name=":2">{{cite book |last=Gould |first=Stephen Jay |author-link=Stephen Jay Gould |date=1997 |title=Full House: The Spread of Excellence from Plato to Darwin |publisher=Harmony |isbn=978-0-609-80140-6 |url=https://archive.org/details/fullhouse00step }}</ref>

In 1989, [[Peter J. Bowler]] defined orthogenesis as:

{{quote|Literally, the term means evolution in a straight line, generally assumed to be evolution that is held to a regular course by forces internal to the organism. Orthogenesis assumes that variation is not random but is directed [[Teleology in biology|towards fixed goals]]. Selection is thus powerless, and the species is carried automatically in the direction marked out by internal factors controlling variation.{{sfn|Bowler|1989|pages=268–270}}}}

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 Lull]]'s ''Ladder of Ascent and Descent of the Mind'', 1305]]

===Medieval===
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 medieval 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.<ref name="Gould 2001"/> 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 [[teleology|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.<ref name=Gould1977>{{cite book |author-link=Stephen Jay Gould |last=Gould |first=Stephen Jay |date=1977 |url=https://archive.org/details/eversincedarwinr00goul |title=Darwin's Dilemma: The Odyssey of Evolution |work=Ever Since Darwin: Reflections in Natural History |publisher=[[W. W. Norton]] |isbn=978-0-393-06425-4 |url-status=dead |access-date=2019-08-01 |archive-url=https://web.archive.org/web/20191216205014/https://archive.org/details/eversincedarwinr00goul |archive-date=2019-12-16 }}</ref> His grandfather, the physician and polymath [[Erasmus Darwin]], was both progressionist and [[Vitalism|vitalist]], seeing "the whole cosmos [as] a living thing propelled by an internal vital force" towards "greater perfection".<ref name="Daly 2018">{{cite journal |last=Daly |first=J. P. |date=4 March 2018 |title=The Botanic Universe: Generative Nature and Erasmus Darwin's Cosmic Transformism |url=https://arcade.stanford.edu/rofl/botanic-universe-generative-nature-and-erasmus-darwins-cosmic-transformism |journal=Republics of Letters |volume=6 |pages=1–57 |access-date=7 December 2021 |archive-date=1 July 2022 |archive-url=https://web.archive.org/web/20220701140038/https://arcade.stanford.edu/rofl/botanic-universe-generative-nature-and-erasmus-darwins-cosmic-transformism |url-status=dead }}</ref> [[Robert Chambers (publisher, born 1802)|Robert Chambers]], in his popular anonymously published 1844 book ''[[Vestiges of the Natural History of Creation]]'' presented a sweeping narrative account of cosmic transmutation, culminating in the evolution of humanity. Chambers included detailed analysis of the fossil record.{{sfn|Bowler|1989|p=134}}

===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 [[evolution]].<ref name=Brown2001/>]]

Ruse 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 Baltic German 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."<ref name=Brown2001>{{cite book |last1=Brown |first1=Keven |last2=Von Kitzing |first2=Eberhard |title=Evolution and Bahá'í Belief: ʻAbduʼl-Bahá's Response to Nineteenth-century Darwinism |url=https://books.google.com/books?id=egDAfpkONRsC&pg=PA159 |year=2001 |publisher=Kalimat Press |isbn=978-1-890688-08-0 |page=159}}</ref><ref>{{cite book |last=Barbieri |first=Marcello |date=2013 |title=Biosemiotics: Information, Codes and Signs in Living Systems |publisher=Nova Science Publishers |page=7 |isbn=978-1-60021-612-1}}</ref><ref>{{cite book |last=Jacobsen |first=Eric Paul |date=2005 |title=From Cosmology to Ecology: The Monist World-view in Germany from 1770 to 1930 |page=100 |publisher=Peter Lang |isbn=978-0-8204-7231-7}}</ref> 
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.<ref name=Vucinich1988>{{cite book |last=Vucinich |first=Alexander |date=1988 |title=Darwin in Russian Thought |publisher=University of California Press |page=137 |isbn=978-0-520-06283-2}}</ref>
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.<ref name=Mawer2006>{{cite book |last=Mawer |first=Simon |author-link=Simon Mawer |title=Gregor Mendel: planting the seeds of genetics |date=2006 |publisher=Harry N. Abrams |isbn=978-0-8109-5748-0 |url=https://archive.org/details/gregormendelplan00simo }}</ref> 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 [[barnacle]]s he studied.<ref name=WatsonAngle>{{cite book |last1=Watson |first1=Marc |last2=Angle |first2=Barbara |title=Man's Selection: Charles Darwin's Theory of Creation, Evolution, And Intelligent Design |url=https://books.google.com/books?id=4lVGDgAAQBAJ&pg=PT149 |year=2017 |publisher=BookBaby |isbn=978-1-936883-14-1 |pages=146–150}}</ref>{{sfn|Ruse|1996|pp=154–155, 162}}
Darwin indeed wrote in his 1859 ''[[Origin of Species]]'':<ref name=Darwin1859>{{cite book |last=Darwin |first=Charles |author-link=Charles Darwin |year=1859 |url=https://www.gutenberg.org/files/1228/1228-h/1228-h.htm#chap10 |title=On the Origin of Species By Means of Natural Selection, or, the Preservation of Favoured Races in the Struggle for Life| at=Chapters 10, 14}}</ref>

{{quote|The inhabitants of each successive period in the world's history have beaten their predecessors in the race for life, and are, insofar, higher in the scale of nature; and this may account for that vague yet ill-defined sentiment, felt by many palaeontologists, that organisation on the whole has progressed. [Chapter 10]<ref name=Darwin1859/>}}

{{quote|As all the living forms of life are the lineal descendants of those which lived long before the Silurian epoch, we may feel certain that the ordinary succession by generation has never once been broken, and that no cataclysm has desolated the whole world. Hence we may look with some confidence to a secure future of equally inappreciable length. And as natural selection works solely by and for the good of each being, all corporeal and mental endowments will tend to progress towards perfection. [Chapter 14]<ref name=Darwin1859/>}}

[[File:Titanothere Osborn.jpg|thumb|upright|[[Henry Fairfield Osborn]]'s 1934 version of orthogenesis,  ''[[aristogenesis]]'', argued that aristogenes, not mutation or natural selection, created all novelty.<ref name=Wallace2005/> Osborn supposed that the horns of [[Titanothere]]s evolved into a [[baroque]] form, way beyond the [[Adaptation|adaptive]] optimum.{{sfn|Ruse|1996|pages=266–267}}]]

In 1898, after studying [[butterfly]] coloration, [[Theodor Eimer<!--mild overlink, seems helpful here-->]] (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 [[adaptation|adaptive]] significance that would be difficult to explain by natural selection.<ref name=Shanahan2004>{{cite book |last=Shanahan |first=Timothy |date=2004 |title=The Evolution of Darwinism: Selection, Adaptation, and Progress in Evolutionary Biology |publisher=Cambridge University Press |page=121 |isbn=978-0-521-54198-5}}</ref> To supporters of orthogenesis, in some cases [[species]] could be led by such trends to [[extinction]].<ref name=Sapp2003>{{cite book |author-link=Jan Sapp |last=Sapp |first=Jan |date=2003 |title=Genesis: The Evolution of Biology |pages=[https://archive.org/details/genesisevolution00sapp/page/69 69–70] |publisher=Oxford University Press |isbn=978-0-19-515619-5 |url=https://archive.org/details/genesisevolution00sapp/page/69 }}</ref> Eimer linked orthogenesis to [[Lamarckism|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]].<ref name=Gould2002>{{cite book |author-link=Stephen Jay Gould |last=Gould |first=Stephen Jay |date=2002 |title=The Structure of Evolutionary Theory |publisher=Harvard University Press |pages=[https://archive.org/details/structureofevolu00goul/page/355 355–364] |isbn=978-0-674-00613-3 |url=https://archive.org/details/structureofevolu00goul/page/355 }}</ref>

===Nineteenth and twentieth centuries===
{{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.<ref name=Simpson1964/> 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 [[plant]]s.<ref>{{cite journal | last=Metcalf |first=Maynard M. | year=1913 | title=Adaptation Through Natural Selection and Orthogenesis | journal=The American Naturalist | volume=47 | issue=554| pages=65–71 | jstor=2455865 | doi=10.1086/279329| doi-access= }}</ref><ref>[[John Merle Coulter]]. (1915). [https://www.jstor.org/stable/1638957 A Suggested Explanation of 'Orthogenesis' in Plants] Science, Vol. 42, No. 1094. pp. 859–863.</ref><ref>{{cite journal | last=Starr |first=Jordan David | author-link=David Starr Jordan | year=1920 | title=Orthogenesis among Fishes | jstor=1646251 | journal=Science | volume=52 | issue=1331| pages=13–14 | doi=10.1126/science.52.1331.13-a| pmid=17793787 | bibcode=1920Sci....52...13S | url=https://zenodo.org/record/1448255 }}</ref><ref>{{cite journal | last=Lipman |first=Charles B. | year=1922 | title=Orthogenesis in Bacteria | journal=The American Naturalist | volume=56 | issue=643| pages=105–115 | jstor=2456503 | doi=10.1086/279851|s2cid=85365933 }}</ref> 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.<ref name=Kwa2011>{{cite book |last=Kwa |first=Chunglin |date=2011 |title=Styles of Knowing: A New History of Science from Ancient Times to the Present |publisher=University of Pittsburgh Press |page=237 |isbn=978-0-8229-6151-2}}</ref><ref>{{cite journal | last=Dimichele | first=William A. | year=1995 | title=Basic Questions in Paleontology: Geologic Time, Organic Evolution, and Biological Systematics, by Otto H. Schindewolf | url=http://si-pddr.si.edu/jspui/bitstream/10088/7124/1/paleo_1995_DiMichele_Schindewolf_book_review_RPP.pdf | journal=Review of Palaeobotany and Palynology | volume=84 | issue=3–4 | pages=481–483 | doi=10.1016/0034-6667(95)90007-1 }}{{Dead link|date=August 2024 |bot=InternetArchiveBot |fix-attempted=yes }}</ref> In 1964 [[George Gaylord Simpson]] argued that orthogenetic theories such as those promulgated by Du Noüy and Sinnott were essentially theology rather than biology.<ref name=Simpson1964>{{cite book |last=Simpson |first=George Gaylord |author-link=George Gaylord Simpson |date=1964 |title=Evolutionary Theology: The New Mysticism |work=This View of Life: The World of an Evolutionist |publisher=Harcourt, Brace & World |pages=213–233}}</ref>

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.<ref>{{cite journal |last=Jepsen |first=Glenn L. |date=1949 |title=Selection. Orthogenesis, and the Fossil Record |journal=Proceedings of the American Philosophical Society |volume=93 |issue=6  |pages=479–500|pmid=15408469 }}</ref> 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.<ref>{{cite journal |author1=Jacobs, Susan C.|author2=Larson, Allan|author3=Cheverud, James M. |date=1995 |title=Phylogenetic Relationships and Orthogenetic Evolution of Coat Color Among Tamarins (Genus Saguinus) |journal=Systematic Biology |volume=44 |issue=4 |pages=515–532 |doi=10.1093/sysbio/44.4.515}}</ref><ref>{{cite journal |last1=Ranganath |first1=H. A. |last2=Hägel |first2=K. |date=1981 |title=Karyotypic orthoselection in Drosophila |journal=Naturwissenschaften |volume=68 |issue=10 |pages=527–528 |doi=10.1007/bf00365385 |bibcode=1981NW.....68..527R |s2cid=29736048 |url=https://zenodo.org/records/10764278/files/Ranganath%20%26%20Haegele,%201981.pdf }}</ref> The term orthoselection was first used by [[Ludwig Hermann Plate]], and was incorporated into the [[Modern synthesis (20th century)|modern synthesis]] by [[Julian Huxley]] and [[Bernard Rensch]].<ref name=LevitOlsson2006>{{cite journal |last1=Levit |first1=Georgy S. |last2=Olsson |first2=Lennart |title='Evolution on Rails': Mechanisms and Levels of Orthogenesis |journal=Annals of the History and Philosophy of Biology | issue=11 |year=2006 |pages=99–138 |url=https://www.univerlag.uni-goettingen.de/bitstream/handle/3/isbn-978-3-938616-85-7/annals%2011_DGGBT.pdf?sequence=1&isAllowed=y}}</ref>

Recent work has supported the mechanism and existence of [[mutation bias]]ed adaptation, meaning that limited local orthogenesis is now seen as possible.<ref name=Sto01>{{cite journal |last1=Yampolsky |first1=L. Y. |last2=Stoltzfus |first2=A. |year=2001 |title=Bias in the introduction of variation as an orienting factor in evolution |journal=Evolution & Development |volume=3 | pages=73–83 |doi=10.1046/j.1525-142x.2001.003002073.x |pmid=11341676 |issue=2|s2cid=26956345 }}</ref><ref name=Stoltzfus_2006_NK>{{cite journal | last=Stoltzfus |first=A. | title=Mutation-Biased Adaptation in a Protein NK Model | journal=Molecular Biology and Evolution | volume=23 | issue=10 | pages=1852–1862 | year=2006 | doi=10.1093/molbev/msl064 | pmid=16857856 | doi-access= }}</ref><ref name=Sto09>{{cite journal |last1=Stoltzfus |first1=A. |last2=Yampolsky |first2=L. Y. |year=2009 |title=Climbing Mount Probable: Mutation as a Cause of Nonrandomness in Evolution |journal=Journal of Heredity |volume=100 | pages=637–647 |doi=10.1093/jhered/esp048 |pmid=19625453 |issue=5 |doi-access=free }}</ref>

==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"
|+ Theories of orthogenesis in [[evolutionary biology]]<ref>{{cite book |editor1=Cigna, Arrigo A.|editor2=Durante, Marco |title=Radiation Risk Estimates in Normal and Emergency Situations |url=https://books.google.com/books?id=lgrABAAAQBAJ&pg=PA213 |year=2007 |publisher=Springer Science & Business Media |isbn=978-1-4020-4956-9 |page=213}}</ref>
! Author !! Title !! Field !! Date !! [[Lamarckism|Lamarck.]] !! [[Mutationism|Mutat.]] !! [[Natural selection|Nat.&nbsp;Sel.]] !! [[Vitalism|Vital.]] !! Features
|-
| [[Jean-Baptiste Lamarck|Lamarck]] || ''[[Inherent progressive tendency]]'' || Zoology || 1809 || yes ||<!--Mutat.--> ||<!--Nat. Sel.--> ||<!--Vital.--> || In his ''[[Philosophie Zoologique]]'', inherent progressive tendency drives organisms continuously towards greater [[Evolution of biological complexity|complexity]], in separate lineages ([[phylum|phyla]]), no [[extinction]].<ref name=Gould1977/> ("[[Lamarckism]]", use and disuse, and [[inheritance of acquired characteristics]], was a secondary aspect of this, an adaptive force creating species within a phylum.<ref name="Gould 2001"/>)

|-
| [[Karl Ernst von Baer|Baer]] || ''[[Purposeful creation]]'' || Embryology || 1859 ||<!--Lamarck.--> ||<!--Mutat.--> ||<!--Nat. Sel.--> ||<!--Vital.--> ||  "Forces which are not directed—so-called blind forces—can never produce order."<ref name=Brown2001/>

|-
| [[Albert von Kölliker|Kölliker]] || ''[[Heterogenesis]]'' || Anatomy || 1864 || yes ||<!--Mutat.--> ||<!--Nat. Sel.--> ||<!--Vital.--> || Wholly separate lines of descent with no common ancestor<ref name=Vucinich1988/>

|-
| [[Edward Drinker Cope|Cope]] || ''[[Law of acceleration]]'' || Palaeontology || 1868 || yes ||<!--Mutat.--> ||<!--Nat. Sel.--> ||<!--Vital.--> || Combined orthogenetic constraints with [[Lamarckian]] use and disuse. "On the Origin of Genera";<ref name=Popov2005/><ref>{{cite web |last1=Barnes |first1=M. Elizabeth |title=Edward Drinker Cope's Law of Acceleration of Growth |url=https://embryo.asu.edu/pages/edward-drinker-copes-law-acceleration-growth  |date=24 July 2014}}</ref><ref name=LevitOlsson2006/> See also [[Cope's rule]] (linear increase in size of species)

|-
| [[Carl Nägeli|Nägeli]] || ''[[Inner perfecting principle]]'' || Botany || 1884 || yes ||<!--Mutat.--> || no ||<!--Vital.--> || An "[[idioplasm]]" transmitted inherited characteristics; many evolutionary developments nonadaptive; variation internally programmed.{{sfn|Bowler|1989|pages=268–270}}<ref name=LevitOlsson2006/>

|-
| [[Herbert Spencer|Spencer]] || Progressionism<br />'The Development Hypothesis' || Social theory || 1852
|| Yes{{sfn|Ruse|1996|page=189}} ||<!--Mutat.--> || <!--Nat. Sel.--> ||<!--Vital.--> || Cultural value of progress; "Spencer has no rivals when it comes to open, flagrant connections of social Progress with evolutionary progress."—[[Michael Ruse]]{{sfn|Ruse|1996|pages=181–191}}

|-
| [[Charles Darwin|Darwin]] || (concept of higher and lower species), [[Pangenesis]] || Evolution || 1859 || yes ||<!--Mutat.--> || yes ||<!--Vital.--> || ''[[Origin of Species]]'' is somewhat progressionist, e.g. man higher than animals, alongside [[natural selection]]<ref name=Darwin1859/><ref name=WatsonAngle/> Pangenesis theory of inheritance by gemmules from all over body was [[Lamarckism|Lamarckian]]: parents could [[inheritance of acquired characters|pass on traits acquired]] in lifetime.<ref name= ImaginaryLamarck>{{Cite book |last =Ghiselin|first=Michael T.|author-link=Michael Ghiselin|date=September–October 1994| contribution =Nonsense in schoolbooks: 'The Imaginary Lamarck'|contribution-url =http://www.textbookleague.org/54marck.htm| title =The Textbook Letter|publisher =The Textbook League|url =http://www.textbookleague.org/|archive-url =https://web.archive.org/web/20000115220615/http://www.textbookleague.org/|url-status =usurped|archive-date =January 15, 2000|access-date=2008-01-23}}</ref><ref>{{cite book |last=Magner |first=Lois N. |title=A History of the Life Sciences |edition=Third |publisher=[[Marcel Dekker]], [[CRC Press]] |year=2002 |isbn=978-0-203-91100-6 |url=https://books.google.com/books?id=YKJ6gVYbrGwC}}</ref>

|-
| [[Wilhelm Haacke|Haacke]] || Orthogenesis || Zoology || 1893 || yes ||<!--Mutat.--> ||<!--Nat. Sel.--> ||<!--Vital.--> || Accompanied by ''[[Epimorphism (biology)|epimorphism]]'', a tendency to increasing perfection<ref name=Popov2005/><ref name=LevitOlsson2006/>

|-
| [[Theodor Eimer|Eimer]] || Orthogenesis|| Zoology  || 1898 ||<!--Lamarck.--> ||<!--Mutat.--> || no ||<!--Vital.--> || ''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.<ref name=Shanahan2004/><ref name=LevitOlsson2006/>
|-
| [[Henri Bergson|Bergson]] || ''[[Élan vital|Elan vital<!--to permit sorting-->]]'' || Philosophy || 1907 ||<!--Lamarck.--> ||<!--Mutat.--> ||<!--Nat. Sel.--> || yes || ''[[Creative Evolution (book)|Creative Evolution]]''{{sfn|Bowler|1989|pages=116–117}}

|-
| [[Hans Przibram|Przibram]] || ''[[Apogenesis]]'' || Embryology || 1910s ||<!--Lamarck.--> ||<!--Mutat.--> ||<!--Nat. Sel.--> ||<!--Vital.--> || <ref name=Popov2005/>

|-
| [[Ludwig Hermann Plate|Plate]] || ''[[Orthoselection]]'' or ''Old-Darwinism'' || Zoology || 1913 || yes || yes || yes ||<!--Vital.--> || Combined theory<ref name=LevitOlsson2006/>

|-
| [[Daniele Rosa|Rosa]] || ''[[Hologenesis]]'' || Zoology  || 1918 || yes ||<!--Mutat.--> ||<!--Nat. Sel.--> ||<!--Vital.--> || ''Hologenesis: a New Theory of Evolution and the Geographical Distribution of Living Beings''<ref name=Luzzatto2000>{{cite journal |author1=Luzzatto, Michele|author2=Palestrini, Claudia|author3=D'entrèves, Passerin Pietro |date=2000 |title=Hologenesis: The Last and Lost Theory of Evolutionary Change |journal=Italian Journal of Zoology |volume=67 |pages=129–138 |doi=10.1080/11250000009356303|s2cid=85796293 }}</ref><ref name=LevitOlsson2006/>

|-
| [[Charles Otis Whitman|Whitman]] || Orthogenesis || Zoology  || 1919 || no || no || no ||<!--Vital.--> || ''Orthogenetic Evolution in Pigeons'' posthumous<ref name=Castle1920>{{cite journal |last1=Castle |first1=W.E. |year=1920 |title=Review of Orthogenetic Evolution in Pigeons |journal=The American Naturalist |volume=54 |issue=631 |pages=188–192 |doi=10.1086/279751|doi-access=free }}</ref><ref>{{cite book |author-link=Stephen Jay Gould |last=Gould |first=Stephen Jay |date=2002 |title=The Structure of Evolutionary Theory |publisher=Harvard University Press |page=[https://archive.org/details/structureofevolu00goul/page/283 283] |isbn=978-0-674-00613-3 |url=https://archive.org/details/structureofevolu00goul/page/283 }}</ref>

|-
| [[Lev Berg|Berg]] || ''[[Nomogenesis]]''|| Zoology || 1926 || no || yes || no ||<!--Vital.--> || Chemical forces direct evolution, leading to humans{{sfn|Ruse|1996|p=395}}<ref name=LevitOlsson2006/>{{sfn|Bowler|1983|p=157}}

|-
| [[Othenio Abel|Abel]] || ''Trägheitsgesetz'' (the law of inertia) || Palaeontology || 1928 ||<!--Lamarck.--> ||<!--Mutat.--> ||<!--Nat. Sel.--> ||<!--Vital.--> || 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)<ref name=LevitOlsson2006/>

|-
| [[André Michel Lwoff|Lwoff]] || ''[[Physiological degradation]]''|| Physiology || 1930s–1940s || yes ||<!--Mutat.--> ||<!--Nat. Sel.--> ||<!--Vital.--> || Directed loss of functions in microorganisms<ref name=Popov2005/><ref>{{cite book |last=Lwoff |first=A. |author-link=André Michel Lwoff |title=L'evolution physiologique. Etude des pertes de fonctions chez les microorganismes |location=Paris |publisher=Hermann |date=1944 |pages=1–308 |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.}}</ref><ref>{{cite journal |last1=Loison |first1=Laurent |last2=Gayon |first2=Jean |last3=Burian |first3=Richard M. |title=The Contributions – and Collapse – of Lamarckian Heredity in Pasteurian Molecular Biology: 1. Lysogeny, 1900–1960 |journal=Journal of the History of Biology |date=2017 |volume=50 |issue=5 |pages=5–52 |doi=10.1007/s10739-015-9434-3 |pmid=26732271 |s2cid=30286465 }}</ref>

|-
| [[Karl Beurlen|Beurlen]] || Orthogenesis  || Palaeontology || 1930 || no ||<!--Mutat.--> || no ||<!--Vital.--> || Start is random ''metakinesis'', generating variety; then ''palingenesis'' (in Beurlen's sense, repeating developmental pathway of ancestors) as mechanism for orthogenesis<ref name=LevitOlsson2006/>

|-
| {{ill|Victor Jollos|pl}} || Directed mutation || Protozoology, Zoology || 1931 || yes ||<!--Mutat.--> ||<!--Nat. Sel.--> ||<!--Vital.--> || Combined orthogenesis with [[Lamarckism]] (inheriting acquired characteristics after heat shock as ''dauermodifications'', passed on by ''plasmatic inheritance'' in the [[cytoplasm]])<ref name=LevitOlsson2006/>

|-
| [[Henry Fairfield Osborn|Osborn]] || ''[[Aristogenesis]]'' || Palaeontology || 1934 || yes || no || no ||<!--Vital.--> || <ref name=Wallace2005>{{cite book |last=Wallace |first=David Rains |date=2005 |title=Beasts of Eden: Walking Whales, Dawn Horses, And Other Enigmas of Mammal Evolution |publisher=University of California Press |page=96 |isbn=978-0-520-24684-3}}</ref><ref>{{cite book |last=Regal |first=Brian |date=2002 |title=Henry Fairfield Osborn: Race, and the Search for the Origins of Man |publisher=Ashgate |pages=184–192 |isbn=978-0-7546-0587-4}}</ref>

|-
| [[John Christopher Willis|Willis]] ||''[[Differentiation (orthogenesis)]]'' || Botany || 1942 ||<!--Lamarck.--> ||<!--Mutat.--> ||<!--Nat. Sel.--> || 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 alterations<ref>{{cite journal |last=Hubbs |first=Carl L. |title=[Review:] The Course of Evolution by J. C. Willis |journal=The American Naturalist |volume=76 |issue=762 (Jan. Feb., 1942) |pages=96–101|doi=10.1086/281018 }}</ref>
|-
| [[Pierre Lecomte du Noüy|Noüy]] || ''[[Telefinalism]]'' || Biophysics || 1947 ||<!--Lamarck.--> ||<!--Mutat.--> ||<!--Nat. Sel.--> || yes || In book ''Human Destiny'',<ref name=Koch1957/> essentially religious<ref name=Koch1957>{{cite journal |last=Koch |first=Leo Francis |date=1957 |title=Vitalistic-Mechanistic Controversy |journal=[[The Scientific Monthly]] |volume=85 |issue=5 |pages=245–255|bibcode=1957SciMo..85..245K }}</ref>

|-
| {{ill|Albert Vandel|fr|lt=Vandel}} || ''[[Organicism]]'' || Zoology || 1949 ||<!--Lamarck.--> ||<!--Mutat.--> ||<!--Nat. Sel.-->No ||<!--Vital.--> || ''L'Homme et L'Evolution''<ref name=Popov2005/>
|-
| [[Edmund Ware Sinnott|Sinnott]] || ''[[Telism]]'' || Botany || 1950 ||<!--Lamarck.--> ||<!--Mutat.--> ||<!--Nat. Sel.--> || yes ||  In book ''Cell and Psyche'',<ref name=Koch1957/> essentially religious<ref name=Simpson1964/>

|-
|  [[Otto Schindewolf|Schindewolf]] || ''[[Typostrophism]]'' || Palaeontology  || 1950 ||<!--Lamarck.--> || yes ||<!--Nat. Sel.--> ||<!--Vital.--> || ''Basic Questions in Paleontology: Geologic Time, Organic Evolution and Biological Systematics''; evolution due to periodic cycle of processes dictated by factors internal to organism.<ref name=Kwa2011/><ref name=LevitOlsson2006/>

|-
| [[Pierre Teilhard de Chardin|Teilhard de Chardin]] || ''[[Directed additivity]]''<br />''[[Omega Point]]'' || Palaeontology<br />[[Mysticism]] || 1959 ||<!--Lamarck.--> ||<!--Mutat.--> ||<!--Nat. Sel.--> || 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]].<ref name=LevitOlsson2006/> Censured by Gaylord Simpson for nonscientific spiritualistic "doubletalk".<ref name="Lane 1996"/><ref name=Teilhard2003>{{cite book |last=Chardin |first=Pierre Teilhard de |year=2003 |orig-year=1959 |title=The Human Phenomenon |publisher=Sussex Academic Press |page=65 |isbn=1-902210-30-1}}</ref><ref>{{cite book |last=Novack |first=George |title=Marxist Writings on History & Philosophy |url=https://books.google.com/books?id=do0aStFKEK0C&pg=PA207 |year=2002 |publisher=Resistance Books |isbn=978-1-876646-23-3 |page=207}}</ref>

|-
| [[Léon Croizat|Croizat]] || ''[[Biological synthesis (philosophy)|Biological synthesis]]''<br />''[[Panbiogeography]]'' || Botany || 1964 ||<!--Lamarck.--> ||<!--Mutat.--> ||<!--Nat. Sel.--> ||<!--Vital.--> || mechanistic, caused by [[developmental constraint]]s or [[phylogenetic constraint]]s<ref name=Popov2005/><ref name="Gray1989">{{cite journal |last=Gray |first=Russell |title=Oppositions in panbiogeography: can the conflicts between selection, constraint, ecology, and history be resolved? |journal=New Zealand Journal of Zoology |date=1989 |volume=16 |issue=4 |pages=787–806 |doi=10.1080/03014223.1989.10422935}}</ref>

|-
| Lima-de-Faria || ''[[Autoevolutionism]]'' || Physics, Chemistry || 1988 || No || No || No || No || Natural selection is immaterial so cannot work.<ref>{{cite book |last=Lima-de-Faria |first=A. |year=1988 |title=Evolution Without Selection: Form and Function by Autoevolution |publisher=Elsevier |isbn=978-0444809636}}</ref>

|}

[[File:Alternatives to Darwinism.svg|thumb|upright=1.5|[[Alternatives to evolution by natural selection|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 (solid blue arrows) to some form of orthogenesis, with or without invoking [[theistic evolution|divine control]] (dotted blue arrows) directly or indirectly. For example, evolutionists like [[Edward Drinker Cope]] believed in a combination of theistic evolution, Lamarckism, vitalism, and orthogenesis,{{sfn|Bowler|1989|pp=261-262}} represented by a sequence of arrows on the left of the diagram. The development of modern Darwinism is indicated by dashed orange arrows.]]

The various [[Alternatives to evolution by natural selection|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 pedigree.jpg|thumb|left|upright=0.9<!--img is almost square, equalizing area-->|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 American]]'', 11 March 1876]]

The stronger versions of the orthogenetic hypothesis began to lose popularity when it became clear that they were inconsistent with the patterns found by [[paleontology|paleontologists]] in the [[fossil record]], which were non-rectilinear (richly branching) with many complications. The hypothesis was abandoned by mainstream biologists when no mechanism could be found that would account for the process, and the theory of evolution by natural selection came to prevail.<ref name=Mayr1982>{{cite book |last=Mayr |first=Ernst |author-link=Ernst Mayr |date=1982 |title=The Growth of Biological Thought: Diversity, Evolution, and Inheritance |publisher=Harvard University Press |pages=530–531 |isbn=978-0-674-36446-2}}</ref> The historian of biology Edward J. Larson commented that

{{quote|At theoretical and philosophical levels, Lamarckism and orthogenesis seemed to solve too many problems to be dismissed out of hand—yet biologists could never reliably document them happening in nature or in the laboratory. Support for both concepts evaporated rapidly once a plausible alternative appeared on the scene.{{sfn|Larson|2004|page=127}}}}

The [[Modern synthesis (20th century)|modern synthesis]] of the 1930s and 1940s, in which the [[genetics|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.<ref>{{cite book |author-link=Stephen Jay Gould |last=Gould |first=Stephen Jay |title=The Structure of Evolutionary Theory |date=2002 |pages=[https://archive.org/details/structureofevolu00goul/page/ Chapter 7, section "Synthesis as Restriction"] |isbn=978-0-674-00613-3 |title-link=The Structure of Evolutionary Theory }}</ref><ref>{{cite book |last=Levinton |first=Jeffrey S. |date=2001 |title=Genetics, Paleontology, and Macroevolution |publisher=Cambridge University Press |pages=14–16 |isbn=978-0-521-80317-5}}</ref><ref>{{cite book |last1=Montgomery |first1=Georgina M. |last2=Largent |first2=Mark A. |date=2015 |title=A Companion to the History of American Science  |publisher=Wiley |page=218 |isbn=978-1-4051-5625-7 |quote=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.}}</ref>

[[File:Ernst Mayr PLoS.jpg|thumb|[[Ernst Mayr]] considered orthogenesis effectively taboo in 1948.{{sfn|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}}<ref name=Mayr1948/> For these and other reasons, belief in evolutionary progress has remained "a persistent [[heresy]]",<ref name=Popov2005>{{cite web |last1=Popov |first1=Igor |title=The Persistence of Heresy: The Concepts of Directed Evolution (Orthogenesis) |url=http://www.kli.ac.at/events/event-detail/1112872500/the-persistence-of-heresy-the-concepts-of-directed-evolution-orthogenesis |access-date=15 April 2017 |date=7 April 2005 |archive-date=15 April 2017 |archive-url=https://web.archive.org/web/20170415202806/http://www.kli.ac.at/events/event-detail/1112872500/the-persistence-of-heresy-the-concepts-of-directed-evolution-orthogenesis |url-status=dead }}</ref> among evolutionary biologists including [[E. O. Wilson]]<ref name=Ruse>{{cite web |last1=Ruse |first1=Michael |author-link1=Michael Ruse |title=Edward O. Wilson on Sociobiology |url=http://www.chronicle.com/blogs/brainstorm/edward-o-wilson-on-sociobiology/22185 |publisher=The Chronicle of Higher Education |access-date=4 April 2017 |date=31 March 2010}}</ref> 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}}

The discipline of [[evolutionary developmental biology]], however, is open to an expanded concept of heredity that incorporates the physics of [[self-organization]]. With its rise in the late 20th-early 21st centuries, ideas of constraint and preferred directions of morphological change have made a reappearance in evolutionary theory.<ref>see, for example, {{cite book |editor1-link=Gerd Müller (theoretical biologist)|editor1=Müller, Gerd B.|editor2-link=Stuart Newman|editor2=Newman, Stuart A.|title=Origination of Organismal Form |date=2003 |publisher=Bradford |isbn=978-0-262-13419-4|title-link=Origination of Organismal Form}}</ref>

===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.<ref name="Tucker2012">{{cite news|last1=Tucker|first1=Jennifer|title=What our most famous evolutionary cartoon gets wrong|url=https://www.bostonglobe.com/ideas/2012/10/27/what-our-most-famous-evolutionary-cartoon-gets-wrong/drKMD5121W6EUxXJ4pF0YL/story.html|access-date=29 December 2017|work=The [[Boston Globe]]|date=28 October 2012}}</ref>]]
{{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."<ref name="Tucker2012"/> 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 diagram 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 (book)|Wonderful Life]]''.<ref name="Tucker2012"/>

[[File:Man is But a Worm.jpg|thumb|upright|''Man is But a Worm'' by [[Edward Linley Sambourne]], ''Punch's Almanack'' for 1882]]

Popular 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 "[[March of Progress|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.<ref name="Tucker2012"/>

[[File:Astronomy Evolution 2 (27458655072).jpg|thumb|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, [[meme]]s, that act much like genes.{{sfn|Ruse|1996|pages=526–539}} Dawkins can speak of "progressive rather than random&nbsp;... trends in evolution".<ref>{{cite journal |author1=Dawkins, Richard |author2=Krebs, J. R. |author-link1=Richard Dawkins |author-link2=John Krebs, Baron Krebs |title=Arms races between and within species |date=1979 |journal=Proceedings of the Royal Society B |volume=205 |issue=1161 |pages=489–511 |doi=10.1098/rspb.1979.0081|pmid=42057 |bibcode=1979RSPSB.205..489D |s2cid=9695900 }}</ref>{{sfn|Ruse|1996|page=466}} Dawkins and [[John Krebs, Baron Krebs|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, Ruse argues, the feeling of progress comes from evolutionary [[arms race]]s 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 [[population genetics|models]] and status", enabling evolutionary biologists to construct the [[Modern synthesis (20th century)|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 [[parallel evolution|independently evolved]] similar patterns, apparently both [[facilitated variation|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]]''}}
{{main|Facilitated variation}}

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 variation|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 [[parallel evolution|independently evolved]] similar patterns. These butterflies are [[Müllerian mimicry|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.<ref>{{cite journal |author1=Baxter, S.W. |author2=Papa, R. |author3=Chamberlain, N. |author4=Humphray, S.J. |author5=Joron, M. |author6=Morrison, C. |author7=ffrench-Constant, R.H. |author8=McMillan, W.O. |author9=Jiggins, C.D. |year=2008 |title=Convergent Evolution in the Genetic Basis of Mullerian Mimicry in Heliconius Butterflies |journal=Genetics |volume=180 |issue=3 |pages=1567–1577 | pmid=18791259 |pmc=2581958 | doi=10.1534/genetics.107.082982}}</ref>

==See also==

* [[Adaptive mutation]]
* [[Convergent evolution]] (contrastable with orthogenesis, not involving teleology)
* [[Devolution (biology)|Devolution]]
* [[Directed evolution]] (in protein engineering)
*[[Directed evolution (transhumanism)]]
* [[Evolutionism]]
* [[Evolution of biological complexity]]
* [[History of evolutionary thought]]
* [[Structuralism (biology)|Structuralism]]
* [[Teleonomy]]
*[[Teleological argument]]

== References==
{{reflist|30em}}

==Sources==

* {{cite book |last=Bowler |first=Peter J. |author-link=Peter J. Bowler |title=The Eclipse of Darwinism: anti-Darwinian evolutionary theories in the decades around 1900 |publisher=Johns Hopkins University Press |year=1983 |isbn=978-0-8018-4391-4 }}
* {{cite book |last=Bowler |first=Peter J. |author-link=Peter J. Bowler |date=1989 |title=Evolution: The History of an Idea |publisher=University of California Press |isbn=978-0-520-06385-3 |url-access=registration |url=https://archive.org/details/evolutionhistory0000bowl }}
* {{cite book |last1=Dawkins |first1=Richard |author-link=Richard Dawkins |title=The Blind Watchmaker |date=1986 |publisher=Longman |isbn=978-0-393-31570-7 |url-access=registration |url=https://archive.org/details/blindwatchmaker0000dawk }}
* {{cite book |last1=Larson |first1=Edward J. |title=Evolution |date=2004 |publisher=Modern Library |isbn=978-0-679-64288-6 |url=https://archive.org/details/evolutionremarka00lars }}
* {{cite book |last=Ruse |first=Michael |author-link=Michael Ruse |date=1996 |title=Monad to man: the Concept of Progress in Evolutionary Biology |url=https://archive.org/details/monadtomanconcep0000ruse |url-access=registration |publisher=Harvard University Press |isbn=978-0-674-03248-4 }}

==Further reading==
* [[William Bateson|Bateson, William]] (1909). "[https://gutenberg.org/etext/1909 Heredity and variation in modern lights]", in ''Darwin and Modern Science'' (A.C. Seward ed.)  Cambridge University Press. Chapter V. 
* [[Daniel Dennett|Dennett, Daniel]] (1995). ''[[Darwin's Dangerous Idea]]''.  Simon & Schuster. {{ISBN|978-0140167344}}.
* [[Julian Huxley|Huxley, Julian]] (1942). ''[[Evolution: The Modern Synthesis]]'', London: George Allen and Unwin.
* {{cite book |last=Mayr |first=Ernst |author-link=Ernst Mayr |date=2002 |title=What Evolution Is |publisher=Weidenfeld and Nicolson |isbn=9780297607410 |ref=none}}
* [[George Gaylord Simpson|Simpson, George G.]] (1957). ''Life Of The Past: Introduction to Paleontology''.  Yale University Press, p.&nbsp;119.
* Wilkins, John (1997). [http://www.talkorigins.org/faqs/macroevolution.html "What is macroevolution?"] 13 October 2004.

==External links==
* [https://www.bostonglobe.com/ideas/2012/10/27/what-our-most-famous-evolutionary-cartoon-gets-wrong/drKMD5121W6EUxXJ4pF0YL/story.html What our most famous evolutionary cartoon gets wrong]

{{Evolution}}

[[Category:Orthogenesis| ]]
[[Category:Non-Darwinian evolution]]
[[Category:History of evolutionary biology]]
[[Category:Teleology]]
[[Category:Vitalism]]
[[Category:Obsolete biology theories]]