Editing Orthogenesis (section)

==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>