Orthogenesis

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Evolutionary progress as a tree of life. Ernst Haeckel, 1866 Haeckel arbol bn.png
Evolutionary progress as a tree of life. Ernst Haeckel, 1866
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 genera. Popular views of Lamarckism only consider an aspect of the adaptive force. Lamarck's Two-Factor Theory.svg
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 genera. Popular views of Lamarckism only consider an aspect of the adaptive force.

Orthogenesis, also known as orthogenetic evolution, progressive evolution, evolutionary progress, or progressionism, is an obsolete 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". [2] [3] [4] 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.

Contents

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. [5] 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; modern supporters include E. O. Wilson and Simon Conway Morris. The evolutionary biologist Ernst Mayr made the term effectively taboo in the journal Nature in 1948, by stating that it implied "some supernatural force". [6] [7] The American paleontologist George Gaylord Simpson (1953) attacked orthogenesis, linking it with vitalism by describing it as "the mysterious inner force". [8] 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

Theodor Eimer Theodor Eimer (Professorengalerie Universitat Tubingen).jpg
Theodor Eimer

The term orthogenesis (from Ancient Greek : ὀρθός orthós, "straight", and Ancient Greek : γένεσις génesis, "origin") was first used by the biologist Wilhelm Haacke in 1893. [9] [10] 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". [11]

In 1922, the zoologist Michael F. Guyer wrote:

[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. [12]

According to Susan R. Schrepfer in 1983:

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 ... By 1910, however most who subscribed to orthogenesis hypothesized some physical rather than metaphysical determinant of orderly change. [13]

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". [14] This may be compared to Stephen Jay Gould's suggestion of "replacing the idea of progress with an operational notion of directionality". [15]

In 1989, Peter J. Bowler defined orthogenesis as:

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 towards fixed goals. Selection is thus powerless, and the species is carried automatically in the direction marked out by internal factors controlling variation. [2]

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". [16]

History

The mediaeval great chain of being as a staircase, implying the possibility of progress: Ramon Lull's Ladder of Ascent and Descent of the Mind, 1305 Die Leiter des Auf- und Abstiegs.jpg
The mediaeval great chain of being as a staircase, implying the possibility of progress: 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. [17]

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. [1] 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. [18] His grandfather, the physician and polymath Erasmus Darwin, was both progressionist and vitalist, seeing "the whole cosmos [as] a living thing propelled by an internal vital force" towards "greater perfection". [19] 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. [20]

With Darwin

Reviewing Darwin's Origin of Species, Karl Ernst von Baer argued for a directed force guiding evolution. Voyages de la Commission scientifique du Nord, en Scandinavie, en Laponie, au Spitzberg et aux Feroe - no-nb digibok 2009040211001-118.jpg
Reviewing Darwin's Origin of Species , Karl Ernst von Baer argued for a directed force guiding evolution.

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." [22] 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." [21] [23] [24] 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. [25] 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. [26] According to Nägeli many evolutionary developments were nonadaptive and variation was internally programmed. [2] 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. [27] [28] Darwin indeed wrote in his 1859 Origin of Species : [29]

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] [29]

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] [29]

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 adaptive optimum. Titanothere Osborn.jpg
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 adaptive optimum.

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. [32] To supporters of orthogenesis, in some cases species could be led by such trends to extinction. [33] 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. [34]

Nineteenth and twentieth centuries

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. [35] 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. [36] [37] [38] [39] 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. [40] [41] In 1964 George Gaylord Simpson argued that orthogenetic theories such as those promulgated by Du Noüy and Sinnott were essentially theology rather than biology. [35]

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. [42] 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. [43] [44] The term orthoselection was first used by Ludwig Hermann Plate, and was incorporated into the modern synthesis by Julian Huxley and Bernard Rensch. [9]

Recent work has supported the mechanism and existence of mutation biased adaptation, meaning that limited local orthogenesis is now seen as possible. [45] [46] [47]

Theories

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.

Theories of orthogenesis in evolutionary biology [48]
AuthorTitleFieldDate Lamarck. Mutat. Nat. Sel. Vital. Features
Lamarck Inherent progressive tendency Zoology1809yesIn his Philosophie Zoologique , inherent progressive tendency drives organisms continuously towards greater complexity, in separate lineages (phyla), no extinction. [18] ("Lamarckism", use and disuse, and inheritance of acquired characteristics, was a secondary aspect of this, an adaptive force creating species within a phylum. [1] )
Baer Purposeful creation Embryology1859"Forces which are not directed—so-called blind forces—can never produce order." [21]
Kölliker Heterogenesis Anatomy1864yesWholly separate lines of descent with no common ancestor [25]
Cope Law of acceleration Palaeontology1868yesCombined orthogenetic constraints with Lamarckian use and disuse. "On the Origin of Genera"; [49] [50] [9] See also Cope's rule (linear increase in size of species)
Nägeli Inner perfecting principle Botany1884yesnoAn "idioplasm" transmitted inherited characteristics; many evolutionary developments nonadaptive; variation internally programmed. [2] [9]
Spencer Progressionism
'The Development Hypothesis'		Social theory1852Yes [51] Cultural value of progress; "Spencer has no rivals when it comes to open, flagrant connections of social Progress with evolutionary progress."—Michael Ruse [52]
Darwin (concept of higher and lower species), Pangenesis Evolution1859yesyes Origin of Species is somewhat progressionist, e.g. man higher than animals, alongside natural selection [29] [27] Pangenesis theory of inheritance by gemmules from all over body was Lamarckian: parents could pass on traits acquired in lifetime. [53] [54]
Haacke OrthogenesisZoology1893yesAccompanied by epimorphism , a tendency to increasing perfection [49] [9]
Eimer OrthogenesisZoology1898noOn Orthogenesis: And the Impotence of Natural Selection in Species Formation: trends in evolution with no adaptive significance, claimed hard to explain by natural selection. [32] [9]
Bergson Elan vital Philosophy1907yes Creative Evolution [55]
Przibram Apogenesis Embryology1910s [49]
Plate Orthoselection or Old-DarwinismZoology1913yesyesyesCombined theory [9]
Rosa Hologenesis Zoology1918yesHologenesis: a New Theory of Evolution and the Geographical Distribution of Living Beings [56] [9]
Whitman OrthogenesisZoology1919nononoOrthogenetic Evolution in Pigeons posthumous [57] [58]
Berg Nomogenesis Zoology1926noyesnoChemical forces direct evolution, leading to humans [59] [9] [60]
Abel Trägheitsgesetz (the law of inertia)Palaeontology1928based 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) [9]
Lwoff Physiological degradation Physiology1930s–1940syesDirected loss of functions in microorganisms [49] [61] [62]
Beurlen OrthogenesisPalaeontology1930nonoStart is random metakinesis, generating variety; then palingenesis (in Beurlen's sense, repeating developmental pathway of ancestors) as mechanism for orthogenesis [9]
Victor Jollos  [ pl ]Directed mutationProtozoology, Zoology1931yesCombined orthogenesis with Lamarckism (inheriting acquired characteristics after heat shock as dauermodifications, passed on by plasmatic inheritance in the cytoplasm) [9]
Osborn Aristogenesis Palaeontology1934yesnono [30] [63]
Willis Differentiation (orthogenesis) Botany1942yesa 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 [64]
Noüy Telefinalism Biophysics1947yesIn book Human Destiny, [65] essentially religious [65]
Vandel  [ fr ] Organicism Zoology1949NoL'Homme et L'Evolution [49]
Sinnott Telism Botany1950yesIn book Cell and Psyche, [65] essentially religious [35]
Schindewolf Typostrophism Palaeontology1950yesBasic Questions in Paleontology: Geologic Time, Organic Evolution and Biological Systematics; evolution due to periodic cycle of processes dictated by factors internal to organism. [40] [9]
Teilhard de Chardin Directed additivity
Omega Point 		Palaeontology
Mysticism 		1959yes 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. [9] Censured by Gaylord Simpson for nonscientific spiritualistic "doubletalk". [11] [66] [67]
Croizat Biological synthesis
Panbiogeography 		Botany1964mechanistic, caused by developmental constraints or phylogenetic constraints [49] [68]
Lima-de-Faria Autoevolutionism Physics, Chemistry1988NoNoNoNoNatural selection is immaterial so cannot work. [69]
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 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, represented by a sequence of arrows on the left of the diagram. The development of modern Darwinism is indicated by dashed orange arrows. Alternatives to Darwinism.svg
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 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, 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 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. [70] Other palaeontologists and field naturalists continued to hold beliefs combining orthogenesis and Lamarckism until the modern synthesis in the 1930s. [71]

Status

In science

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 Human pedigree.jpg
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 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. [72] The historian of biology Edward J. Larson commented that

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. [73]

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. [74] [75] [76]

Ernst Mayr considered orthogenesis effectively taboo in 1948. Ernst Mayr PLoS.jpg
Ernst Mayr considered orthogenesis effectively taboo in 1948.

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." [6] [7] For these and other reasons, belief in evolutionary progress has remained "a persistent heresy", [49] among evolutionary biologists including E. O. Wilson [77] 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." [78] He argued that progressionism has harmed the status of evolutionary biology as a mature, professional science. [79] 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. [4]

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. [80]

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. Huxley - Mans Place in Nature.jpg
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.

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." [81] 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 . [81]

Man is But a Worm by Edward Linley Sambourne, Punch's Almanack for 1882 Man is But a Worm.jpg
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 "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. [81]

One of many versions of the progressionist meme: Astronomy Evolution 2 artwork by Giuseppe Donatiello, 2016 Astronomy Evolution 2 (27458655072).jpg
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. [4] Dawkins can speak of "progressive rather than random ... trends in evolution". [82] [83] Dawkins and John Krebs deny the "earlier [Darwinian] prejudice" [84] that there is anything "inherently progressive about evolution", [85] [84] but, Ruse argues, 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". [86] [84]

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", [4] until mathematicians such as Fisher [87] 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. [4]

Facilitated variation

Heliconius erato Richard Bartz.jpg
Heliconius erato
Heliconius melpomene 2b Richard Bartz.jpg
Heliconius melpomene
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.

Biology has largely rejected the idea that evolution is guided in any way, [88] [73] 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. [89]

See also

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"Survival of the fittest" is a phrase that originated from Darwinian evolutionary theory as a way of describing the mechanism of natural selection. The biological concept of fitness is defined as reproductive success. In Darwinian terms, the phrase is best understood as "[s]urvival of the form that in successive generations will leave most copies of itself."

<span class="mw-page-title-main">Michael Ruse</span> Canadian philosopher of science (born 1940)

Michael Ruse is a British-born Canadian philosopher of science who specializes in the philosophy of biology and works on the relationship between science and religion, the creation–evolution controversy, and the demarcation problem within science. Ruse currently teaches at Florida State University.

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

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

Devolution, de-evolution, or backward evolution is the notion that species can revert to supposedly more primitive forms over time. The concept relates to the idea that evolution has a purpose (teleology) and is progressive (orthogenesis), for example that feet might be better than hooves or lungs than gills. However, evolutionary biology makes no such assumptions, and natural selection shapes adaptations with no foreknowledge of any kind. It is possible for small changes to be reversed by chance or selection, but this is no different from the normal course of evolution and as such de-evolution is not compatible with a proper understanding of evolution due to natural selection.

In biology, saltation is a sudden and large mutational change from one generation to the next, potentially causing single-step speciation. This was historically offered as an alternative to Darwinism. Some forms of mutationism were effectively saltationist, implying large discontinuous jumps.

<span class="mw-page-title-main">Structuralism (biology)</span> Attempt to explain evolution by forces other than natural selection

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

Julian Huxley used the phrase "the eclipse of Darwinism" to describe the state of affairs prior to what he called the "modern synthesis". During the "eclipse", evolution was widely accepted in scientific circles but relatively few biologists believed that natural selection was its primary mechanism. Historians of science such as Peter J. Bowler have used the same phrase as a label for the period within the history of evolutionary thought from the 1880s to around 1920, when alternatives to natural selection were developed and explored—as many biologists considered natural selection to have been a wrong guess on Charles Darwin's part, or at least to be of relatively minor importance.

<span class="mw-page-title-main">History of evolutionary thought</span>

Evolutionary thought, the recognition that species change over time and the perceived understanding of how such processes work, has roots in antiquity—in the ideas of the ancient Greeks, Romans, Chinese, Church Fathers as well as in medieval Islamic science. With the beginnings of modern biological taxonomy in the late 17th century, two opposed ideas influenced Western biological thinking: essentialism, the belief that every species has essential characteristics that are unalterable, a concept which had developed from medieval Aristotelian metaphysics, and that fit well with natural theology; and the development of the new anti-Aristotelian approach to modern science: as the Enlightenment progressed, evolutionary cosmology and the mechanical philosophy spread from the physical sciences to natural history. Naturalists began to focus on the variability of species; the emergence of palaeontology with the concept of extinction further undermined static views of nature. In the early 19th century prior to Darwinism, Jean-Baptiste Lamarck (1744–1829) proposed his theory of the transmutation of species, the first fully formed theory of evolution.

<span class="mw-page-title-main">Daniele Rosa</span>

Daniele Rosa (1857–1944) was an Italian invertebrate zoologist.

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

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

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

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

<i>Monad to Man</i> 1996 book by Michael Ruse

Monad to Man: the concept of progress in evolutionary biology is a 1996 book about the longstanding idea that evolution is progressive by the philosopher of biology Michael Ruse. It analyses the connection between ideas of progress in culture generally and its application in evolutionary biology.

<span class="mw-page-title-main">Evolution in fiction</span> Role of evolution as a theme in fiction

Evolution has been an important theme in fiction, including speculative evolution in science fiction, since the late 19th century, though it began before Charles Darwin's time, and reflects progressionist and Lamarckist views as well as Darwin's. Darwinian evolution is pervasive in literature, whether taken optimistically in terms of how humanity may evolve towards perfection, or pessimistically in terms of the dire consequences of the interaction of human nature and the struggle for survival. Other themes include the replacement of humanity, either by other species or by intelligent machines.

References

  1. 1 2 3 Gould, Stephen J. (2001). The lying stones of Marrakech : penultimate reflections in natural history. Vintage. pp. 119–121. ISBN   978-0-09-928583-0.
  2. 1 2 3 4 Bowler 1989, pp. 268–270.
  3. ↑ Mayr, Ernst (1988). Toward a New Philosophy of Biology: Observations of an Evolutionist. Harvard University Press. p. 499. ISBN   978-0-674-89666-6.
  4. 1 2 3 4 5 Ruse 1996, pp. 526–539.
  5. ↑ 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. doi:10.1016/j.shpsc.2013.11.009. PMID   24368232.
  6. 1 2 3 Ruse 1996, p. 447.
  7. 1 2 Letter from Ernst Mayr to R. H. Flower, Evolution papers, 23 January 1948
  8. ↑ Simpson, George Gaylord (1953). Life of the Past: An Introduction to Paleontology . Yale University Press. p.  125.
  9. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 Levit, Georgy S.; Olsson, Lennart (2006). "'Evolution on Rails': Mechanisms and Levels of Orthogenesis" (PDF). Annals of the History and Philosophy of Biology (11): 99–138.
  10. ↑ Gould, Stephen Jay (2002). The Structure of Evolutionary Theory. Harvard University Press. pp.  351–352. ISBN   978-0-674-00613-3.
  11. 1 2 Lane, David H. (1996). The Phenomenon of Teilhard: Prophet for a New Age. Mercer University Press. pp. 60–64. ISBN   978-0-86554-498-7.
  12. ↑ Guyer, Michael F. (1922). "Orthogenesis and Serological Phenomena". The American Naturalist. 56 (643): 116–133. doi:10.1086/279852. JSTOR   2456504.
  13. ↑ Schrepfer, Susan R. (1983). Fight to Save the Redwoods: A History of the Environmental Reform, 1917–1978. University of Wisconsin Press. pp. 81–82. ISBN   978-0-299-08854-5.
  14. ↑ Ayala, Francisco J. (1988). "Can progress be defined as a biological concept?". In Nitecki, M. (ed.). Evolutionary Progress. University of Chicago Press. pp. 75–96. ISBN   978-0-226-58693-9.
  15. ↑ Gould, Stephen Jay (1997). Full House: The Spread of Excellence from Plato to Darwin. Harmony. ISBN   978-0-609-80140-6.
  16. ↑ Ruse 1996, p. 261.
  17. 1 2 Ruse 1996, pp. 21–23.
  18. 1 2 Gould, Stephen Jay (1977). Darwin's Dilemma: The Odyssey of Evolution. W. W. Norton. ISBN   978-0-393-06425-4. Archived from the original on 2019-12-16. Retrieved 2019-08-01.{{cite book}}: |work= ignored (help)
  19. ↑ Daly, J. P. (4 March 2018). "The Botanic Universe: Generative Nature and Erasmus Darwin's Cosmic Transformism". Republics of Letters. 6: 1–57.
  20. ↑ Bowler 1989, p. 134.
  21. 1 2 3 Brown, Keven; Von Kitzing, Eberhard (2001). Evolution and Bahá'í Belief: ʻAbduʼl-Bahá's Response to Nineteenth-century Darwinism. Kalimat Press. p. 159. ISBN   978-1-890688-08-0.
  22. ↑ Ruse 1996, p. 29.
  23. ↑ Barbieri, Marcello (2013). Biosemiotics: Information, Codes and Signs in Living Systems. Nova Science Publishers. p. 7. ISBN   978-1-60021-612-1.
  24. ↑ Jacobsen, Eric Paul (2005). From Cosmology to Ecology: The Monist World-view in Germany from 1770 to 1930. Peter Lang. p. 100. ISBN   978-0-8204-7231-7.
  25. 1 2 Vucinich, Alexander (1988). Darwin in Russian Thought. University of California Press. p. 137. ISBN   978-0-520-06283-2.
  26. ↑ Mawer, Simon (2006). Gregor Mendel: planting the seeds of genetics. Harry N. Abrams. ISBN   978-0-8109-5748-0.
  27. 1 2 Watson, Marc; Angle, Barbara (2017). Man's Selection: Charles Darwin's Theory of Creation, Evolution, And Intelligent Design. BookBaby. pp. 146–150. ISBN   978-1-936883-14-1.
  28. ↑ Ruse 1996, pp. 154–155, 162.
  29. 1 2 3 4 Darwin, Charles (1859). On the Origin of Species By Means of Natural Selection, or, the Preservation of Favoured Races in the Struggle for Life. Chapters 10, 14.
  30. 1 2 Wallace, David Rains (2005). Beasts of Eden: Walking Whales, Dawn Horses, And Other Enigmas of Mammal Evolution. University of California Press. p. 96. ISBN   978-0-520-24684-3.
  31. ↑ Ruse 1996, pp. 266–267.
  32. 1 2 Shanahan, Timothy (2004). The Evolution of Darwinism: Selection, Adaptation, and Progress in Evolutionary Biology. Cambridge University Press. p. 121. ISBN   978-0-521-54198-5.
  33. ↑ Sapp, Jan (2003). Genesis: The Evolution of Biology. Oxford University Press. pp.  69–70. ISBN   978-0-19-515619-5.
  34. ↑ Gould, Stephen Jay (2002). The Structure of Evolutionary Theory. Harvard University Press. pp.  355–364. ISBN   978-0-674-00613-3.
  35. 1 2 3 Simpson, George Gaylord (1964). Evolutionary Theology: The New Mysticism. Harcourt, Brace & World. pp. 213–233.{{cite book}}: |work= ignored (help)
  36. ↑ Metcalf, Maynard M. (1913). "Adaptation Through Natural Selection and Orthogenesis". The American Naturalist. 47 (554): 65–71. doi:10.1086/279329. JSTOR   2455865.
  37. ↑ John Merle Coulter. (1915). A Suggested Explanation of 'Orthogenesis' in Plants Science, Vol. 42, No. 1094. pp. 859–863.
  38. ↑ Starr, Jordan David (1920). "Orthogenesis among Fishes". Science. 52 (1331): 13–14. Bibcode:1920Sci....52...13S. doi:10.1126/science.52.1331.13-a. JSTOR   1646251. PMID   17793787.
  39. ↑ Lipman, Charles B. (1922). "Orthogenesis in Bacteria". The American Naturalist. 56 (643): 105–115. doi:10.1086/279851. JSTOR   2456503. S2CID   85365933.
  40. 1 2 Kwa, Chunglin (2011). Styles of Knowing: A New History of Science from Ancient Times to the Present. University of Pittsburgh Press. p. 237. ISBN   978-0-8229-6151-2.
  41. ↑ Dimichele, William A. (1995). "Basic Questions in Paleontology: Geologic Time, Organic Evolution, and Biological Systematics, by Otto H. Schindewolf" (PDF). Review of Palaeobotany and Palynology. 84 (3–4): 481–483. doi:10.1016/0034-6667(95)90007-1.
  42. ↑ Jepsen, Glenn L. (1949). "Selection. Orthogenesis, and the Fossil Record". Proceedings of the American Philosophical Society. 93 (6): 479–500. PMID   15408469.
  43. ↑ Jacobs, Susan C.; Larson, Allan; Cheverud, James M. (1995). "Phylogenetic Relationships and Orthogenetic Evolution of Coat Color Among Tamarins (Genus Saguinus)". Systematic Biology. 44 (4): 515–532. doi:10.1093/sysbio/44.4.515.
  44. ↑ Ranganath, H. A.; Hägel, K. (1981). "Karyotypic orthoselection in Drosophila". Naturwissenschaften. 68 (10): 527–528. Bibcode:1981NW.....68..527R. doi:10.1007/bf00365385. S2CID   29736048.
  45. ↑ Yampolsky, L. Y.; Stoltzfus, A. (2001). "Bias in the introduction of variation as an orienting factor in evolution". Evolution & Development. 3 (2): 73–83. doi:10.1046/j.1525-142x.2001.003002073.x. PMID   11341676. S2CID   26956345.
  46. ↑ Stoltzfus, A. (2006). "Mutation-Biased Adaptation in a Protein NK Model". Molecular Biology and Evolution. 23 (10): 1852–1862. doi:10.1093/molbev/msl064. PMID   16857856.
  47. ↑ Stoltzfus, A.; Yampolsky, L. Y. (2009). "Climbing Mount Probable: Mutation as a Cause of Nonrandomness in Evolution". Journal of Heredity. 100 (5): 637–647. doi: 10.1093/jhered/esp048 . PMID   19625453.
  48. ↑ Cigna, Arrigo A.; Durante, Marco, eds. (2007). Radiation Risk Estimates in Normal and Emergency Situations. Springer Science & Business Media. p. 213. ISBN   978-1-4020-4956-9.
  49. 1 2 3 4 5 6 7 Popov, Igor (7 April 2005). "The Persistence of Heresy: The Concepts of Directed Evolution (Orthogenesis)" . Retrieved 15 April 2017.
  50. ↑ Barnes, M. Elizabeth (24 July 2014). "Edward Drinker Cope's Law of Acceleration of Growth".
  51. ↑ Ruse 1996, p. 189.
  52. ↑ Ruse 1996, pp. 181–191.
  53. ↑ Ghiselin, Michael T. (September–October 1994). "Nonsense in schoolbooks: 'The Imaginary Lamarck'". The Textbook Letter. The Textbook League. Retrieved 2008-01-23.
  54. ↑ Magner, Lois N. (2002). A History of the Life Sciences (Third ed.). Marcel Dekker, CRC Press. ISBN   978-0-203-91100-6.
  55. ↑ Bowler 1989, pp. 116–117.
  56. ↑ Luzzatto, Michele; Palestrini, Claudia; D'entrèves, Passerin Pietro (2000). "Hologenesis: The Last and Lost Theory of Evolutionary Change". Italian Journal of Zoology. 67: 129–138. doi:10.1080/11250000009356303. S2CID   85796293.
  57. ↑ Castle, W.E. (1920). "Review of Orthogenetic Evolution in Pigeons". The American Naturalist. 54 (631): 188–192. doi: 10.1086/279751 .
  58. ↑ Gould, Stephen Jay (2002). The Structure of Evolutionary Theory. Harvard University Press. p.  283. ISBN   978-0-674-00613-3.
  59. ↑ Ruse 1996, p. 395.
  60. ↑ Bowler 1983, p. 157.
  61. ↑ Lwoff, A. (1944). L'evolution physiologique. Etude des pertes de fonctions chez les microorganismes. Paris: Hermann. pp. 1–308. 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.
  62. ↑ Loison, Laurent; Gayon, Jean; Burian, Richard M. (2017). "The Contributions – and Collapse – of Lamarckian Heredity in Pasteurian Molecular Biology: 1. Lysogeny, 1900–1960". Journal of the History of Biology. 50 (5): 5–52. doi:10.1007/s10739-015-9434-3. PMID   26732271. S2CID   30286465.
  63. ↑ Regal, Brian (2002). Henry Fairfield Osborn: Race, and the Search for the Origins of Man. Ashgate. pp. 184–192. ISBN   978-0-7546-0587-4.
  64. ↑ Hubbs, Carl L. "[Review:] The Course of Evolution by J. C. Willis". The American Naturalist. 76 (762 (Jan. Feb., 1942)): 96–101. doi:10.1086/281018.
  65. 1 2 3 Koch, Leo Francis (1957). "Vitalistic-Mechanistic Controversy". The Scientific Monthly . 85 (5): 245–255. Bibcode:1957SciMo..85..245K.
  66. ↑ Chardin, Pierre Teilhard de (2003) [1959]. The Human Phenomenon. Sussex Academic Press. p. 65. ISBN   1-902210-30-1.
  67. ↑ Novack, George (2002). Marxist Writings on History & Philosophy. Resistance Books. p. 207. ISBN   978-1-876646-23-3.
  68. ↑ Gray, Russell (1989). "Oppositions in panbiogeography: can the conflicts between selection, constraint, ecology, and history be resolved?". New Zealand Journal of Zoology. 16 (4): 787–806. doi:10.1080/03014223.1989.10422935.
  69. ↑ Lima-de-Faria, A. (1988). Evolution Without Selection: Form and Function by Autoevolution. Elsevier. ISBN   978-0444809636.
  70. 1 2 Bowler 1989, pp. 261–262.
  71. ↑ Bowler 1989, p. 264.
  72. ↑ Mayr, Ernst (1982). The Growth of Biological Thought: Diversity, Evolution, and Inheritance. Harvard University Press. pp. 530–531. ISBN   978-0-674-36446-2.
  73. 1 2 Larson 2004, p. 127.
  74. ↑ Gould, Stephen Jay (2002). The Structure of Evolutionary Theory. pp.  Chapter 7, section "Synthesis as Restriction". ISBN   978-0-674-00613-3.
  75. ↑ Levinton, Jeffrey S. (2001). Genetics, Paleontology, and Macroevolution. Cambridge University Press. pp. 14–16. ISBN   978-0-521-80317-5.
  76. ↑ Montgomery, Georgina M.; Largent, Mark A. (2015). A Companion to the History of American Science. Wiley. p. 218. ISBN   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.
  77. ↑ Ruse, Michael (31 March 2010). "Edward O. Wilson on Sociobiology". The Chronicle of Higher Education. Retrieved 4 April 2017.
  78. ↑ Ruse 1996, p. 536.
  79. ↑ Ruse 1996, p. 530.
  80. ↑ see, for example, Müller, Gerd B.; Newman, Stuart A., eds. (2003). Origination of Organismal Form. Bradford. ISBN   978-0-262-13419-4.
  81. 1 2 3 4 Tucker, Jennifer (28 October 2012). "What our most famous evolutionary cartoon gets wrong". The Boston Globe . Retrieved 29 December 2017.
  82. ↑ Dawkins, Richard; Krebs, J. R. (1979). "Arms races between and within species". Proceedings of the Royal Society B. 205 (1161): 489–511. Bibcode:1979RSPSB.205..489D. doi:10.1098/rspb.1979.0081. PMID   42057. S2CID   9695900.
  83. ↑ Ruse 1996, p. 466.
  84. 1 2 3 Ruse 1996, p. 468.
  85. ↑ Dawkins 1986, p. 178.
  86. ↑ Dawkins 1986, p. 181.
  87. ↑ Ruse 1996, pp. 292–295.
  88. ↑ Bowler 1989, p. 270.
  89. ↑ 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. doi:10.1534/genetics.107.082982. PMC   2581958 . PMID   18791259.

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