Lamarckism

Lamarckism, also known as Lamarckian inheritance or neo-Lamarckism,[2] is the notion that an organism can pass on to its offspring physical characteristics that the parent organism acquired through use or disuse during its lifetime. It is also called the inheritance of acquired characteristics or more recently soft inheritance. The idea is named after the French zoologist Jean-Baptiste Lamarck (1744–1829), who incorporated the classical era theory of soft inheritance into his theory of evolution as a supplement to his concept of orthogenesis, a drive towards complexity.

Lamarck argued, as part of his theory of heredity, that a blacksmith's sons inherit the strong muscles he acquires from his work.[1]

Introductory textbooks contrast Lamarckism with Charles Darwin's theory of evolution by natural selection. However, Darwin's book On the Origin of Species gave credence to the idea of heritable effects of use and disuse, as Lamarck had done, and his own concept of pangenesis similarly implied soft inheritance.[2][3]

Many researchers from the 1860s onwards attempted to find evidence for Lamarckian inheritance, but these have all been explained away,[4][5] either by other mechanisms such as genetic contamination or as fraud. August Weismann's experiment, considered definitive in its time, is now considered to have failed to disprove Lamarckism, as it did not address use and disuse. Later, Mendelian genetics supplanted the notion of inheritance of acquired traits, eventually leading to the development of the modern synthesis, and the general abandonment of Lamarckism in biology. Despite this, interest in Lamarckism has continued.

Studies in the fields of epigenetics, genetics, and somatic hypermutation have highlighted the possible inheritance of traits acquired by the previous generation. The characterization of these findings as Lamarckism has been disputed. The inheritance of the hologenome, consisting of the genomes of all an organism's symbiotic microbes as well as its own genome, is also somewhat Lamarckian in effect, though entirely Darwinian in its mechanisms.

Early history

Origins
Jean-Baptiste Lamarck repeated the ancient folk wisdom of the inheritance of acquired characteristics.

The inheritance of acquired characteristics was proposed in ancient times, and remained a current idea for many centuries. The historian of science Conway Zirkle wrote in 1935 that:[6]

Lamarck was neither the first nor the most distinguished biologist to believe in the inheritance of acquired characters. He merely endorsed a belief which had been generally accepted for at least 2,200 years before his time and used it to explain how evolution could have taken place. The inheritance of acquired characters had been accepted previously by Hippocrates, Aristotle, Galen, Roger Bacon, Jerome Cardan, Levinus Lemnius, John Ray, Michael Adanson, Jo. Fried. Blumenbach and Erasmus Darwin among others.[6]

Zirkle noted that Hippocrates described pangenesis, the theory that what is inherited derives from the whole body of the parent, whereas Aristotle thought it impossible; but that all the same, Aristotle implicitly agreed to the inheritance of acquired characteristics, giving the example of the inheritance of a scar, or of blindness, though noting that children do not always resemble their parents. Zirkle recorded that Pliny the Elder thought much the same. Zirkle pointed out that stories involving the idea of inheritance of acquired characteristics appear numerous times in ancient mythology and the Bible, and persisted through to Rudyard Kipling's Just So Stories.[7] The idea is mentioned in 18th century sources such as Diderots D'Alembert's Dream.[8] Erasmus Darwin's Zoonomia (c. 1795) suggested that warm-blooded animals develop from "one living filament... with the power of acquiring new parts" in response to stimuli, with each round of "improvements" being inherited by successive generations.[9]

Darwin's pangenesis
Charles Darwin's pangenesis theory. Every part of the body emits tiny gemmules which migrate to the gonads and contribute to the next generation via the fertilised egg. Changes to the body during an organism's life would be inherited, as in Lamarckism.

Charles Darwin's On the Origin of Species proposed natural selection as the main mechanism for development of species, but (like Lamarck) gave credence to the idea of heritable effects of use and disuse as a supplementary mechanism.[10] Darwin subsequently set out his concept of pangenesis in the final chapter of his book The Variation of Animals and Plants Under Domestication (1868), which gave numerous examples to demonstrate what he thought was the inheritance of acquired characteristics. Pangenesis, which he emphasised was a hypothesis, was based on the idea that somatic cells would, in response to environmental stimulation (use and disuse), throw off 'gemmules' or 'pangenes' which travelled around the body, though not necessarily in the bloodstream. These pangenes were microscopic particles that supposedly contained information about the characteristics of their parent cell, and Darwin believed that they eventually accumulated in the germ cells where they could pass on to the next generation the newly acquired characteristics of the parents.[11][12]

Darwin's half-cousin, Francis Galton, carried out experiments on rabbits, with Darwin's cooperation, in which he transfused the blood of one variety of rabbit into another variety in the expectation that its offspring would show some characteristics of the first. They did not, and Galton declared that he had disproved Darwin's hypothesis of pangenesis, but Darwin objected, in a letter to the scientific journal Nature, that he had done nothing of the sort, since he had never mentioned blood in his writings. He pointed out that he regarded pangenesis as occurring in protozoa and plants, which have no blood, as well as in animals.[13]

Lamarck's evolutionary framework
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. Lamarckism is the name now widely used for the adaptive force.
Further information: Jean-Baptiste Lamarck § Lamarckian evolution

Between 1800 and 1830, Lamarck proposed a systematic theoretical framework for understanding evolution. He saw evolution as comprising four laws:[14][15]

  1. "Life by its own force, tends to increase the volume of all organs which possess the force of life, and the force of life extends the dimensions of those parts up to an extent that those parts bring to themselves;"
  2. "The production of a new organ in an animal body, results from a new requirement arising. and which continues to make itself felt, and a new movement which that requirement gives birth to, and its upkeep/maintenance;"
  3. "The development of the organs, and their ability, are constantly a result of the use of those organs."
  4. "All that has been acquired, traced, or changed, in the physiology of individuals, during their life, is conserved through the genesis, reproduction, and transmitted to new individuals who are related to those who have undergone those changes."

Lamarck's discussion of heredity

In 1830, in an aside from his evolutionary framework, Lamarck briefly mentioned two traditional ideas in his discussion of heredity, in his day considered to be generally true. The first was the idea of use versus disuse; he theorized that individuals lose characteristics they do not require, or use, and develop characteristics that are useful. The second was to argue that the acquired traits were heritable. He gave as an imagined illustration the idea that when giraffes stretch their necks to reach leaves high in trees, they would strengthen and gradually lengthen their necks. These giraffes would then have offspring with slightly longer necks. In the same way, he argued, a blacksmith, through his work, strengthens the muscles in his arms, and thus his sons would have similar muscular development when they mature. Lamarck stated the following two laws:[1]

  1. Première Loi: Dans tout animal qui n' a point dépassé le terme de ses développemens, l' emploi plus fréquent et soutenu d' un organe quelconque, fortifie peu à peu cet organe, le développe, l' agrandit, et lui donne une puissance proportionnée à la durée de cet emploi; tandis que le défaut constant d' usage de tel organe, l'affoiblit insensiblement, le détériore, diminue progressivement ses facultés, et finit par le faire disparoître.[1]
  2. Deuxième Loi: Tout ce que la nature a fait acquérir ou perdre aux individus par l' influence des circonstances où leur race se trouve depuis long-temps exposée, et, par conséquent, par l' influence de l' emploi prédominant de tel organe, ou par celle d' un défaut constant d' usage de telle partie; elle le conserve par la génération aux nouveaux individus qui en proviennent, pourvu que les changemens acquis soient communs aux deux sexes, ou à ceux qui ont produit ces nouveaux individus.[1]

English translation:

  1. First Law [Use and Disuse]: In every animal which has not passed the limit of its development, a more frequent and continuous use of any organ gradually strengthens, develops and enlarges that organ, and gives it a power proportional to the length of time it has been so used; while the permanent disuse of any organ imperceptibly weakens and deteriorates it, and progressively diminishes its functional capacity, until it finally disappears.
  2. Second Law [Soft Inheritance]: All the acquisitions or losses wrought by nature on individuals, through the influence of the environment in which their race has long been placed, and hence through the influence of the predominant use or permanent disuse of any organ; all these are preserved by reproduction to the new individuals which arise, provided that the acquired modifications are common to both sexes, or at least to the individuals which produce the young.[16]

In essence, a change in the environment brings about change in "needs" (besoins), resulting in change in behaviour, causing change in organ usage and development, bringing change in form over time—and thus the gradual transmutation of the species. The evolutionary biologists and historians of science Conway Zirkle, Michael Ghiselin, and Stephen Jay Gould have pointed out, these ideas were not original to Lamarck.[6][2][17]

Weismann's experiment
August Weismann's germ plasm theory. The hereditary material, the germ plasm, is confined to the gonads and the gametes. Somatic cells (of the body) develop afresh in each generation from the germ plasm, creating an invisible "Weismann barrier" to Lamarckian influence from the soma to the next generation.

August Weismann's germ plasm theory held that germline cells in the gonads contain information that passes from one generation to the next, unaffected by experience, and independent of the somatic (body) cells. This implied what came to be known as the Weismann barrier, as it would make Lamarckian inheritance from changes to the body difficult or impossible.[18]

Weismann conducted the experiment of removing the tails of 68 white mice, and those of their offspring over five generations, and reporting that no mice were born in consequence without a tail or even with a shorter tail. In 1889, he stated that "901 young were produced by five generations of artificially mutilated parents, and yet there was not a single example of a rudimentary tail or of any other abnormality in this organ."[19] The experiment, and the theory behind it, were thought at the time to be a refutation of Lamarckism.[18]

The experiment's effectiveness in refuting Lamarck's hypothesis is doubtful, as it did not address the use and disuse of characteristics in response to the environment. The biologist Peter Gauthier noted in 1990 that:[20]

Can Weismann's experiment be considered a case of disuse? Lamarck proposed that when an organ was not used, it slowly, and very gradually atrophied. In time, over the course of many generations, it would gradually disappear as it was inherited in its modified form in each successive generation. Cutting the tails off mice does not seem to meet the qualifications of disuse, but rather falls in a category of accidental misuse... Lamarck's hypothesis has never been proven experimentally and there is no known mechanism to support the idea that somatic change, however acquired, can in some way induce a change in the germplasm. On the other hand it is difficult to disprove Lamarck's idea experimentally, and it seems that Weismann's experiment fails to provide the evidence to deny the Lamarckian hypothesis, since it lacks a key factor, namely the willful exertion of the animal in overcoming environmental obstacles.[20]

Ghiselin also considered the Weismann tail-chopping experiment to have no bearing on the Lamarckian hypothesis, writing in 1994 that:[2]

The acquired characteristics that figured in Lamarck's thinking were changes that resulted from an individual's own drives and actions, not from the actions of external agents. Lamarck was not concerned with wounds, injuries or mutilations, and nothing that Lamarck had set forth was tested or "disproven" by the Weismann tail-chopping experiment.[2]

The historian of science Rasmus Winther stated that Weismann had nuanced views about the role of the environment on the germ plasm. Indeed, like Darwin, he consistently insisted that a variable environment was necessary to cause variation in the hereditary material.[21]

Textbook Lamarckism
The long neck of the giraffe is often used as an example in popular explanations of Lamarckism. However, this was only a small part of his theory of evolution towards "perfection"; it was a hypothetical illustration; and he used it to discuss his theory of heredity, not evolution.[2]

The identification of Lamarckism with the inheritance of acquired characteristics is regarded by evolutionary biologists including Ghiselin as a falsified artifact of the subsequent history of evolutionary thought, repeated in textbooks without analysis, and wrongly contrasted with a falsified picture of Darwin's thinking. Ghiselin notes that "Darwin accepted the inheritance of acquired characteristics, just as Lamarck did, and Darwin even thought that there was some experimental evidence to support it."[2] Gould wrote that in the late 19th century, evolutionists "re-read Lamarck, cast aside the guts of it ... and elevated one aspect of the mechanics—inheritance of acquired characters—to a central focus it never had for Lamarck himself."[22] He argued that "the restriction of 'Lamarckism' to this relatively small and non-distinctive corner of Lamarck's thought must be labelled as more than a misnomer, and truly a discredit to the memory of a man and his much more comprehensive system."[3][23]

Neo-Lamarckism

Context
Edward Drinker Cope

The period of the history of evolutionary thought between Darwin's death in the 1880s, and the foundation of population genetics in the 1920s and the beginnings of the modern evolutionary synthesis in the 1930s, is called the eclipse of Darwinism by some historians of science. During that time many scientists and philosophers accepted the reality of evolution but doubted whether natural selection was the main evolutionary mechanism.[24]

Among the most popular alternatives were theories involving the inheritance of characteristics acquired during an organism's lifetime. Scientists who felt that such Lamarckian mechanisms were the key to evolution were called neo-Lamarckians. They included the British botanist George Henslow (1835–1925), who studied the effects of environmental stress on the growth of plants, in the belief that such environmentally-induced variation might explain much of plant evolution, and the American entomologist Alpheus Spring Packard, Jr., who studied blind animals living in caves and wrote a book in 1901 about Lamarck and his work.[25][26] Also included were paleontologists like Edward Drinker Cope and Alpheus Hyatt, who observed that the fossil record showed orderly, almost linear, patterns of development that they felt were better explained by Lamarckian mechanisms than by natural selection. Some people, including Cope and the Darwin critic Samuel Butler, felt that inheritance of acquired characteristics would let organisms shape their own evolution, since organisms that acquired new habits would change the use patterns of their organs, which would kick-start Lamarckian evolution. They considered this philosophically superior to Darwin's mechanism of random variation acted on by selective pressures. Lamarckism also appealed to those, like the philosopher Herbert Spencer and the German anatomist Ernst Haeckel, who saw evolution as an inherently progressive process.[25] The German zoologist Theodor Eimer combined Larmarckism with ideas about orthogenesis, the idea that evolution is directed towards a goal.[27]

With the development of the modern synthesis of the theory of evolution, and a lack of evidence for a mechanism for acquiring and passing on new characteristics, or even their heritability, Lamarckism largely fell from favour. Unlike neo-Darwinism, neo-Lamarckism is a loose grouping of largely heterodox theories and mechanisms that emerged after Lamarck's time, rather than a coherent body of theoretical work.[28]

19th century
Charles-Édouard Brown-Séquard tried to demonstrate Lamarckism by mutilating guinea pigs.

Neo-Lamarckian versions of evolution were widespread in the late 19th century. The idea that living things could to some degree choose the characteristics that would be inherited allowed them to be in charge of their own destiny as opposed to the Darwinian view, which placed them at the mercy of the environment. Such ideas were more popular than natural selection in the late 19th century as it made it possible for biological evolution to fit into a framework of a divine or naturally willed plan, thus the neo-Lamarckian view of evolution was often advocated by proponents of orthogenesis.[29] According to the historian of science Peter J. Bowler, writing in 2003:

One of the most emotionally compelling arguments used by the neo-Lamarckians of the late nineteenth century was the claim that Darwinism was a mechanistic theory which reduced living things to puppets driven by heredity. The selection theory made life into a game of Russian roulette, where life or death was predetermined by the genes one inherited. The individual could do nothing to mitigate bad heredity. Lamarckism, in contrast, allowed the individual to choose a new habit when faced with an environmental challenge and shape the whole future course of evolution.[30]

Scientists from the 1860s onwards conducted numerous experiments that purported to show Lamarckian inheritance. Some examples are described in the table.

19th century experiments attempting to demonstrate Lamarckian inheritance
ScientistDateExperimentClaimed resultRebuttal
Charles-Édouard Brown-Séquard1869 to 1891Cut sciatic nerve and dorsal spinal cord of guinea pigs, causing abnormal nervous condition resembling epilepsyEpileptic offspringNot Lamarckism, as no use and disuse in response to environment; results could not be replicated; cause possibly a transmitted disease.[31][32][33][34][35][36]
Gaston Bonnier1884, 1886Transplant plants at different altitudes in Alps, PyreneesAcquired adaptationsNot controlled from weeds; likely cause genetic contamination[37]
Joseph Thomas Cunningham1891, 1893, 1895Shine light on underside of flatfishInherited production of pigmentDisputed cause[38][39][40][41][42][43]
Max Standfuss1892 to 1917Raise butterflies at low temperatureVariations in offspring even without low temperatureRichard Goldschmidt agreed; Ernst Mayr "difficult to interpret".[44][45][46][47]

Early 20th century
Paul Kammerer claimed in the 1920s to have found evidence for Lamarckian inheritance in midwife toads, in a case celebrated by the journalist Arthur Koestler, but the results are thought to be either fraudulent or at best misinterpreted.

A century after Lamarck, scientists and philosophers continued to seek mechanisms and evidence for the inheritance of acquired characteristics. Experiments were sometimes reported as successful, but from the beginning these were either criticised on scientific grounds or shown to be fakes.[48][49][50][4][5] For instance, in 1906, the philosopher Eugenio Rignano argued for a version that he called "centro-epigenesis",[51][52][53][54][55][56] but it was rejected by most scientists.[57] Some of the experimental approaches are described in the table.

Early 20th century experiments attempting to demonstrate Lamarckian inheritance
ScientistDateExperimentClaimed resultRebuttal
William Lawrence Tower1907 to 1910Colorado potato beetles in extreme humidity, temperatureHeritable changes in size, colourCriticised by William Bateson; Tower claimed all results lost in fire; William E. Castle visited laboratory, found fire suspicious, doubted claim that steam leak had killed all beetles, concluded faked data.[58][59][60][49][50]
Gustav Tornier1907 to 1918Goldfish, embryos of frogs, newtsAbnormalities inheritedDisputed; possibly an osmotic effect[61][62][63][64]
Charles Rupert Stockard1910Repeated alcohol intoxication of pregnant guinea pigsInherited malformationsRaymond Pearl unable to reproduce findings in chickens; Darwinian explanation[65][48]
Francis Bertody Sumner1921Reared mice at different temperatures, humiditiesInherited longer bodies, tails, hind feetInconsistent results[66][67]
Michael F. Guyer, Elizabeth A. Smith1918 to 1924Injected fowl serum antibodies for rabbit lens-protein into pregnant rabbitsEye defects inherited for 8 generationsDisputed, results not replicated[68][69]
Paul Kammerer1920sMidwife toadBlack foot-pads inheritedFraud, ink injected; or, results misinterpreted; case celebrated by Arthur Koestler arguing that opposition was political[4][70]
William McDougall1920sRats solving mazesOffspring learnt mazes quicker (20 vs 165 trials)Poor experimental controls[71][72][73][74][75][76][5]
John William Heslop-Harrison1920sPeppered moths exposed to sootInherited mutations caused by sootFailure to replicate results; implausible mutation rate[77][78]
Ivan Pavlov1926Conditioned reflex in mice to food and bellOffspring easier to conditionPavlov retracted claim; results not replicable[79][80]
Coleman Griffith, John Detlefson1920 to 1925Reared rats on rotating table for 3 monthsInherited balance disorderResults not replicable; likely cause ear infection[81][82][83][84][85][86]
Victor Jollos1930sHeat treatment in Drosophila melanogasterDirected mutagenesis, a form of orthogenesisResults not replicable[87][88]

Late 20th century

The British anthropologist Frederic Wood Jones and the South African paleontologist Robert Broom supported a neo-Lamarckian view of human evolution. The German anthropologist Hermann Klaatsch relied on a neo-Lamarckian model of evolution to try and explain the origin of bipedalism. Neo-Lamarckism remained influential in biology until the 1940s when the role of natural selection was reasserted in evolution as part of the modern evolutionary synthesis.[89] Herbert Graham Cannon, a British zoologist, defended Lamarckism in his 1959 book Lamarck and Modern Genetics.[90] In the 1960s, "biochemical Lamarckism" was advocated by the embryologist Paul Wintrebert.[91]

Neo-Lamarckism was dominant in French biology for more than a century. French scientists who supported neo-Lamarckism included Edmond Perrier (1844–1921), Alfred Giard (1846–1908), Gaston Bonnier (1853–1922) and Pierre-Paul Grassé (1895–1985). They followed two traditions, one mechanistic, one vitalistic after Henri Bergson's philosophy of evolution.[92]

In 1987, Ryuichi Matsuda coined the term "pan-environmentalism" for his evolutionary theory which he saw as a fusion of Darwinism with neo-Lamarckism. He held that heterochrony is a main mechanism for evolutionary change and that novelty in evolution can be generated by genetic assimilation.[93][94] His views were criticized by Arthur M. Shapiro for providing no solid evidence for his theory. Shapiro noted that "Matsuda himself accepts too much at face value and is prone to wish-fulfilling interpretation."[94]

Ideological neo-Lamarckism
Trofim Lysenko promoted an ideological form of neo-Lamarckism which adversely influenced Soviet agricultural policy in the 1930s.

A form of Lamarckism was revived in the Soviet Union of the 1930s when Trofim Lysenko promoted the ideologically-driven research programme, Lysenkoism; this suited the ideological opposition of Joseph Stalin to genetics. Lysenkoism influenced Soviet agricultural policy which in turn was later blamed for crop failures.[95]

Critique

George Gaylord Simpson in his book Tempo and Mode in Evolution (1944) claimed that experiments in heredity have failed to corroborate any Lamarckian process.[96] Simpson noted that neo-Lamarckism "stresses a factor that Lamarck rejected: inheritance of direct effects of the environment" and neo-Lamarckism is closer to Darwin's pangenesis than Lamarck's views.[97] Simpson wrote, "the inheritance of acquired characters, failed to meet the tests of observation and has been almost universally discarded by biologists."[98]

Botanist Conway Zirkle pointed out that Lamarck did not originate the hypothesis that acquired characteristics could be inherited, so it is incorrect to refer to it as Lamarckism:

What Lamarck really did was to accept the hypothesis that acquired characters were heritable, a notion which had been held almost universally for well over two thousand years and which his contemporaries accepted as a matter of course, and to assume that the results of such inheritance were cumulative from generation to generation, thus producing, in time, new species. His individual contribution to biological theory consisted in his application to the problem of the origin of species of the view that acquired characters were inherited and in showing that evolution could be inferred logically from the accepted biological hypotheses. He would doubtless have been greatly astonished to learn that a belief in the inheritance of acquired characters is now labeled "Lamarckian," although he would almost certainly have felt flattered if evolution itself had been so designated.[7]

Peter Medawar wrote regarding Lamarckism, "very few professional biologists believe that anything of the kind occurs—or can occur—but the notion persists for a variety of nonscientific reasons." Medawar stated there is no known mechanism by which an adaptation acquired in an individual's lifetime can be imprinted on the genome and Lamarckian inheritance is not valid unless it excludes the possibility of natural selection but this has not been demonstrated in any experiment.[99]

Martin Gardner wrote in his book Fads and Fallacies in the Name of Science (1957):

A host of experiments have been designed to test Lamarckianism. All that have been verified have proved negative. On the other hand, tens of thousands of experiments— reported in the journals and carefully checked and rechecked by geneticists throughout the world— have established the correctness of the gene-mutation theory beyond all reasonable doubt... In spite of the rapidly increasing evidence for natural selection, Lamarck has never ceased to have loyal followers.... There is indeed a strong emotional appeal in the thought that every little effort an animal puts forth is somehow transmitted to his progeny.[100]

According to Ernst Mayr, any Lamarckian theory involving the inheritance of acquired characters has been refuted as "DNA does not directly participate in the making of the phenotype and that the phenotype, in turn, does not control the composition of the DNA."[101] Peter J. Bowler has written that although many early scientists took Lamarckism seriously, it was discredited by genetics in the early twentieth century.[102]

Mechanisms resembling Lamarckism

Studies in the field of epigenetics, genetics and somatic hypermutation[103][104] have highlighted the possible inheritance of traits acquired by the previous generation.[105][106][107][108][109] However, the characterization of these findings as Lamarckism has been disputed.[110][111][112][113]

Transgenerational epigenetic inheritance
DNA molecule with epigenetic marks, created by methylation, enabling a neo-Lamarckian pattern of inheritance for some generations.

Epigenetic inheritance has been argued by scientists including Eva Jablonka and Marion J. Lamb to be Lamarckian.[114] Epigenetics is based on hereditary elements other than genes that pass into the germ cells. These include methylation patterns in DNA and chromatin marks on histone proteins, both involved in gene regulation. These marks are responsive to environmental stimuli, differentially affect gene expression, and are adaptive, with phenotypic effects that persist for some generations. The mechanism may also enable the inheritance of behavioral traits, for example in chickens,[115][116][117] rats[118][119] and human populations that have experienced starvation, DNA methylation resulting in altered gene function in both the starved population and their offspring.[120] Methylation similarly mediates epigenetic inheritance in plants such as rice.[121][122] Small RNA molecules, too, may mediate inherited resistance to infection.[123][124][125] Handel and Romagopalan commented that "epigenetics allows the peaceful co-existence of Darwinian and Lamarckian evolution."[126]

Joseph Springer and Dennis Holley commented in 2013 that:[127]

Lamarck and his ideas were ridiculed and discredited. In a strange twist of fate, Lamarck may have the last laugh. Epigenetics, an emerging field of genetics, has shown that Lamarck may have been at least partially correct all along. It seems that reversible and heritable changes can occur without a change in DNA sequence (genotype) and that such changes may be induced spontaneously or in response to environmental factors—Lamarck's "acquired traits." Determining which observed phenotypes are genetically inherited and which are environmentally induced remains an important and ongoing part of the study of genetics, developmental biology, and medicine.[127]

The prokaryotic CRISPR system and Piwi-interacting RNA could be classified as Lamarckian, within a Darwinian framework.[128][129] However, the significance of epigenetics in evolution is uncertain. Critics such as the evolutionary biologist Jerry Coyne point out that epigenetic inheritance lasts for only a few generations, so it is not a stable basis for evolutionary change.[130][131][132][133]

The evolutionary biologist T. Ryan Gregory contends that epigenetic inheritance should not be considered Lamarckian. According to Gregory, Lamarck did not claim that the environment directly affected living things. Instead, Lamarck "argued that the environment created needs to which organisms responded by using some features more and others less, that this resulted in those features being accentuated or attenuated, and that this difference was then inherited by offspring." Gregory has stated that Lamarckian evolution in epigenetics is more like Darwin's point of view than Lamarck's.[110]

In 2007, David Haig wrote that research into epigenetic processes does allow a Lamarckian element in evolution but the processes do not challenge the main tenets of the modern evolutionary synthesis as modern Lamarckians have claimed. Haig argued for the primacy of DNA and evolution of epigenetic switches by natural selection.[134] Haig has written that there is a "visceral attraction" to Lamarckian evolution from the public and some scientists, as it posits the world with a meaning, in which organisms can shape their own evolutionary destiny.[135]

Thomas Dickens and Qazi Rahman (2012) have argued that epigenetic mechanisms such as DNA methylation and histone modification are genetically inherited under the control of natural selection and do not challenge the modern synthesis. They dispute the claims of Jablonka and Lamb on Lamarckian epigenetic processes.[136]

Edward J. Steele's disputed[137] Neo-Lamarckian mechanism involves somatic hypermutation and reverse transcription by a retrovirus to breach the Weismann barrier to germline DNA.

In 2015, Khursheed Iqbal and colleagues discovered that although "endocrine disruptors exert direct epigenetic effects in the exposed fetal germ cells, these are corrected by reprogramming events in the next generation."[138] Also in 2015, Adam Weiss argued that bringing back Lamarck in the context of epigenetics is misleading, commenting, "We should remember [Lamarck] for the good he contributed to science, not for things that resemble his theory only superficially. Indeed, thinking of CRISPR and other phenomena as Lamarckian only obscures the simple and elegant way evolution really works."[139]

Somatic hypermutation and reverse transcription to germline

In the 1970s, the Australian immunologist Edward J. Steele developed a neo-Lamarckian theory of somatic hypermutation within the immune system and coupled it to the reverse transcription of RNA derived from body cells to the DNA of germline cells. This reverse transcription process supposedly enabled characteristics or bodily changes acquired during a lifetime to be written back into the DNA and passed on to subsequent generations.[140][141]

The mechanism was meant to explain why homologous DNA sequences from the VDJ gene regions of parent mice were found in their germ cells and seemed to persist in the offspring for a few generations. The mechanism involved the somatic selection and clonal amplification of newly acquired antibody gene sequences generated via somatic hypermutation in B-cells. The messenger RNA products of these somatically novel genes were captured by retroviruses endogenous to the B-cells and were then transported through the bloodstream where they could breach the Weismann or soma-germ barrier and reverse transcribe the newly acquired genes into the cells of the germ line, in the manner of Darwin's pangenes.[104][103][142]

Neo-Lamarckian inheritance of hologenome[143]

The historian of biology Peter J. Bowler noted in 1989 that other scientists had been unable to reproduce his results, and described the scientific consensus at the time:[137]

There is no feedback of information from the proteins to the DNA, and hence no route by which characteristics acquired in the body can be passed on through the genes. The work of Ted Steele (1979) provoked a flurry of interest in the possibility that there might, after all, be ways in which this reverse flow of information could take place. ... [His] mechanism did not, in fact, violate the principles of molecular biology, but most biologists were suspicious of Steele's claims, and attempts to reproduce his results have failed.[137]

Bowler commented that "[Steele's] work was bitterly criticized at the time by biologists who doubted his experimental results and rejected his hypothetical mechanism as implausible."[137]

Hologenome theory of evolution

The hologenome theory of evolution, while Darwinian, has Lamarckian aspects. An individual animal or plant lives in symbiosis with many microorganisms, and together they have a "hologenome" consisting of all their genomes. The hologenome can vary like any other genome by mutation, sexual recombination, and chromosome rearrangement, but in addition it can vary when populations of microorganisms increase or decrease (resembling Lamarckian use and disuse), and when it gains new kinds of microorganism (resembling Lamarckian inheritance of acquired characteristics). These changes are then passed on to offspring.[144] The mechanism is largely uncontroversial, and natural selection does sometimes occur at whole system (hologenome) level, but it is not clear that this is always the case.[143]

Lamarckian use and disuse compared to Darwinian evolution, the Baldwin effect, and Waddington's genetic assimilation. All the theories offer explanations of how organisms respond to a changed environment with adaptive inherited change.

Baldwin effect

The Baldwin effect, named after the psychologist James Mark Baldwin by George Gaylord Simpson in 1953, proposes that the ability to learn new behaviours can improve an animal's reproductive success, and hence the course of natural selection on its genetic makeup. Simpson stated that the mechanism was "not inconsistent with the modern synthesis" of evolutionary theory,[145] though he doubted that it occurred very often, or could be proven to occur. He noted that the Baldwin effect provided a reconciliation between the neo-Darwinian and neo-Lamarckian approaches, something that the modern synthesis had seemed to render unnecessary. In particular, the effect allows animals to adapt to a new stress in the environment through behavioural changes, followed by genetic change. This somewhat resembles Lamarckism but without requiring animals to inherit characteristics acquired by their parents.[146] The Baldwin effect is broadly accepted by Darwinists.[147]

In sociocultural evolution

Within the field of cultural evolution, Lamarckism has been applied as a mechanism for dual inheritance theory.[148] Gould viewed culture as a Lamarckian process whereby older generations transmitted adaptive information to offspring via the concept of learning. In the history of technology, components of Lamarckism have been used to link cultural development to human evolution by considering technology as extensions of human anatomy.[149]

References
  1. Lamarck 1830, p. 235
  2. Ghiselin, Michael T. (1994). "The Imaginary Lamarck: A Look at Bogus "History" in Schoolbooks". The Textbook Letter (September–October 1994). Archived from the original on 12 October 2000. Retrieved 17 February 2006.
  3. Gould 2002, pp. 177–178
  4. Bowler 2003, pp. 245–246
  5. Medawar 1985, p. 168
  6. Zirkle, Conway (1935). "The Inheritance of Acquired Characters and the Provisional Hypothesis of Pangenesis". The American Naturalist. 69 (724): 417–445. doi:10.1086/280617. S2CID 84729069.
  7. Zirkle, Conway (January 1946). "The Early History of the Idea of the Inheritance of Acquired Characters and of Pangenesis". Transactions of the American Philosophical Society. 35 (2): 91–151. doi:10.2307/1005592. JSTOR 1005592.
  8. Diderot, Denis; Ballestero, Manuel (1992). El sueño de D'Alembert; y Suplemento al viaje de Bougainville (First ed.). Madrid: Debate. ISBN 84-7444-583-3. OCLC 433436276.
  9. Darwin 1794–1796, Vol I, section XXXIX
  10. Desmond & Moore 1991, p. 617: "But Darwin was loath to let go of the notion that a well-used and strengthened organ could be inherited."
  11. Darwin, Charles (April 27, 1871). "Pangenesis". Nature. 3 (78): 502–503. Bibcode:1871Natur...3..502D. doi:10.1038/003502a0.
  12. Holterhoff, Kate (2014). "The History and Reception of Charles Darwin's Hypothesis of Pangenesis". Journal of the History of Biology. 47 (4): 661–695. doi:10.1007/s10739-014-9377-0. PMID 24570302. S2CID 207150548.
  13. Liu, Yongsheng (2008). "A new perspective on Darwin's Pangenesis". Biological Reviews. 83 (2): 141–149. doi:10.1111/j.1469-185x.2008.00036.x. PMID 18429766. S2CID 39953275.
  14. Larson, Edward J. (2004). A Growing sense of progress. Evolution: The remarkable history of a Scientific Theory. Modern Library. pp. 38–41.
  15. Gould, Stephen (2001). The lying stones of Marrakech : penultimate reflections in natural history. Vintage. pp. 119–121. ISBN 978-0-09-928583-0.
  16. Lamarck 1914, p. 113
  17. Gould 2002, pp. 170–191
  18. Romanes, George John (1893). An examination of Weismannism. Open Court. OL 23380098M.
  19. Weismann 1889, "The Supposed Transmission of Mutilations" (1888), p. 432
  20. Gauthier, Peter (March–May 1990). "Does Weismann's Experiment Constitute a Refutation of the Lamarckian Hypothesis?". BIOS. 61 (1/2): 6–8. JSTOR 4608123.
  21. Winther, Rasmus (2001). "August Weismann on Germ-Plasm Variation". Journal of the History of Biology. 34 (3): 517–555. doi:10.1023/A:1012950826540. PMID 11859887. S2CID 23808208.
  22. Gould 1980, p. 66
  23. Gould, Stephen Jay (October 4, 1979). "Another Look at Lamarck". New Scientist. Vol. 84, no. 1175. pp. 38–40. Retrieved 2015-11-09.
  24. Quammen 2006, p. 216
  25. Bowler 2003, pp. 236–244
  26. Quammen 2006, pp. 218, 220
  27. Quammen 2006, p. 221
  28. Bowler, Peter J. (1989) [1983]. Evolution: The History of an Idea (Revised ed.). University of California Press. pp. 257, 264, 279–280. ISBN 978-0520063860.
  29. Bowler 1992
  30. Bowler 2003, p. 367
  31. Mumford 1921, p. 209
  32. Mason 1956, p. 343
  33. Burkhardt 1995, p. 166
  34. Raitiere 2012, p. 299
  35. Linville & Kelly 1906, p. 108
  36. Aminoff 2011, p. 192
  37. Kohler 2002, p. 167
  38. Cunningham, Joseph Thomas (1891). "An Experiment concerning the Absence of Color from the lower Sides of Flat-fishes". Zoologischer Anzeiger. 14: 27–32.
  39. Cunningham, Joseph Thomas (May 1893). "Researches on the Coloration of the Skins of Flat Fishes". Journal of the Marine Biological Association of the United Kingdom. 3 (1): 111–118. doi:10.1017/S0025315400049596. S2CID 84934811.
  40. Cunningham, Joseph Thomas (May 1895). "Additional Evidence on the Influence of Light in producing Pigments on the Lower Sides of Flat Fishes" (PDF). Journal of the Marine Biological Association of the United Kingdom. 4: 53–59. doi:10.1017/S0025315400050761. S2CID 86159587.
  41. Moore, Eldon (September 15, 1928). "The New View of Mendelism". The Spectator (Book review). Vol. 141, no. 5229. p. 337. Retrieved 2015-10-24. Review of Modern Biology (1928) by J. T. Cunningham.
  42. Cock & Forsdyke 2008, pp. 132–133
  43. Morgan 1903, pp. view=1up, seq=277 257–259
  44. Goldschmidt 1940, pp. 266–267
  45. Burkhardt 1998, "Lamarckism in Britain and the United States", p. 348
  46. Forel 1934, p. 36
  47. Packard, A. S. (July 10, 1896). "Handbuch der paläarktischen Gross-Schmetterlinge für Forscher und Sammler. Zweite gänzlich umgearbeitete und durch Studien zur Descendenztheorie erweitete Auflage, etc". Science (Book review). 4 (80): 52–54. doi:10.1126/science.4.80.52-c. Review of Handbuch der paläarktischen Gross-Schmetterlinge für Forscher und Sammler (1896) by Maximilian Rudolph Standfuss.
  48. Delage & Goldsmith 1912, p. 210
  49. Kohler 2002, pp. 202–204
  50. Mitman 1992, p. 219
  51. Rignano 1906
  52. Rignano & Harvey 1911
  53. Eastwood, M. Lightfoot (October 1912). "Reviewed Work: Eugenio Rignano Upon the Inheritance of Acquired Characters by C.H. Harvey". International Journal of Ethics. 23 (1): 117–118. doi:10.1086/206715. JSTOR 2377122.
  54. Newman 1921, p. 335
  55. Rignano 1926
  56. Carmichael, Leonard (December 23, 1926). "Reviewed Work: Biological Memory by Eugenio Rignano, E. W. MacBride". The Journal of Philosophy. 23 (26): 718–720. doi:10.2307/2014451. JSTOR 2014451.
  57. "(1) Upon the Inheritance of Acquired Characters (2) Biological Aspects of Human Problems". Nature (Book review). 89 (2232): 576–578. August 8, 1912. Bibcode:1912Natur..89..576.. doi:10.1038/089576a0. S2CID 3984855.
  58. Bateson, William (July 3, 1919). "Dr. Kammerer's Testimony to the Inheritance of Acquired Characters". Nature (Letter to editor). 103 (2592): 344–345. Bibcode:1919Natur.103..344B. doi:10.1038/103344b0. S2CID 4146761.
  59. Bateson 1913, pp. 219–227
  60. Weinstein 1998, "A Note on W. L. Tower's Lepinotarsa Work," pp. 352–353
  61. MacBride, Ernest (January 1924). "The work of tornier as affording a possible explanation of the causes of mutations". The Eugenics Review. 15 (4): 545–555. PMC 2942563. PMID 21259774.
  62. Cunningham 1928, pp. 84–97
  63. Sladden, Dorothy E. (May 1930). "Experimental Distortion of Development in Amphibian Tadpoles". Proceedings of the Royal Society B. 106 (744): 318–325. doi:10.1098/rspb.1930.0031.
  64. Sladden, Dorothy E. (November 1932). "Experimental Distortion of Development in Amphibian Tadpoles. Part II". Proceedings of the Royal Society B. 112 (774): 1–12. Bibcode:1932RSPSB.112....1S. doi:10.1098/rspb.1932.0072.
  65. Blumberg 2010, pp. 69–70
  66. Young 1922, p. 249
  67. Child 1945, pp. 146–173
  68. Guyer, Michael F.; Smith, E. A. (March 1920). "Transmission of Eye-Defects Induced in Rabbits by Means of Lens-Sensitized Fowl-Serum". PNAS. 6 (3): 134–136. Bibcode:1920PNAS....6..134G. doi:10.1073/pnas.6.3.134. PMC 1084447. PMID 16576477.
  69. Medawar 1985, p. 169
  70. Moore 2002, p. 330
  71. McDougall, William (April 1938). "Fourth Report on a Lamarckian Experiment". General Section. British Journal of Psychology. 28 (4): 365–395. doi:10.1111/j.2044-8295.1938.tb00882.x.
  72. Pantin, Carl F. A. (November 1957). "Oscar Werner Tiegs. 1897-1956". Biographical Memoirs of Fellows of the Royal Society. 3: 247–255. doi:10.1098/rsbm.1957.0017.
  73. Agar, Wilfred E.; Drummond, Frank H.; Tiegs, Oscar W. (July 1935). "A First Report on a Test of McDougall'S Lamarckian Experiment on the Training of Rats". The Journal of Experimental Biology. 12 (3): 191–211. doi:10.1242/jeb.12.3.191.
  74. Agar, Wilfred E.; Drummond, Frank H.; Tiegs, Oscar W. (October 1942). "Second Report on a Test of McDougall's Lamarckian Experiment on the Training of Rats". The Journal of Experimental Biology. 19 (2): 158–167. doi:10.1242/jeb.19.2.158.
  75. Agar, Wilfred E.; Drummond, Frank H.; Tiegs, Oscar W. (June 1948). "Third Report on a Test of McDougall'S Lamarckian Experiment on the Training of Rats". The Journal of Experimental Biology. 25 (2): 103–122. doi:10.1242/jeb.25.2.103. Retrieved 2015-10-28.
  76. Agar, Wilfred E.; Drummond, Frank H.; Tiegs, Oscar W.; Gunson, Mary M. (September 1954). "Fourth (Final) Report on a Test of McDougall'S Lamarckian Experiment on the Training of Rats". The Journal of Experimental Biology. 31 (3): 308–321. doi:10.1242/jeb.31.3.307.
  77. Hagen 2002, p. 144: "During the 1920s, the entomologist J. W. Heslop-Harrison published experimental data supporting his claim that chemicals in soot caused widespread mutations from light winged to the dark winged form. Because these mutations were supposedly passed on to subsequent generations, Harrison claimed that he had documented a case of inheritance of acquired traits. Other biologists failed to replicate Harrison's results, and R. A. Fisher pointed out that Harrison's hypothesis required a mutation rate far higher than any previously reported."
  78. Moore & Decker 2008, p. 203
  79. McDougall 1934, p. 180
  80. Macdowell, E. Carleton; Vicari, Emilia M. (May 1921). "Alcoholism and the behavior of white rats. I. The influence of alcoholic grandparents upon maze-behavior". Journal of Experimental Zoology. 33 (1): 208–291. doi:10.1002/jez.1400330107.
  81. Griffith, Coleman R. (November–December 1920). "The Effect upon the White Rat of Continued Bodily Rotation". The American Naturalist. 54 (635): 524–534. doi:10.1086/279783. JSTOR 2456346. S2CID 84453628.
  82. Griffith, Coleman R. (December 15, 1922). "Are Permanent Disturbances of Equilibration Inherited?". Science. 56 (1459): 676–678. Bibcode:1922Sci....56..676G. doi:10.1126/science.56.1459.676. PMID 17778266.
  83. Detlefsen, John A. (1923). "Are the Effects of Long-Continued Rotation in Rats Inherited?". Proceedings of the American Philosophical Society. 62 (5): 292–300. JSTOR 984462.
  84. Detlefsen, John A. (April 1925). "The inheritance of acquired characters". Physiological Reviews. 5 (2): 224–278. doi:10.1152/physrev.1925.5.2.244.
  85. Dorcus, Roy M. (June 1933). "The effect of intermittent rotation on orientation and the habituation of nystagmus in the rat, and some observations on the effects of pre-natal rotation on post-natal development". Journal of Comparative Psychology. 15 (3): 469–475. doi:10.1037/h0074715.
  86. Otho S. A. Sprague Memorial Institute 1940, p. 162
  87. Jollos, Victor (September 1934). "Inherited changes produced by heat-treatment in Drosophila melanogaster". Genetica. 16 (5–6): 476–494. doi:10.1007/BF01984742. S2CID 34126149.
  88. Harwood 1993, pp. 121–131
  89. Wood 2013
  90. Cannon 1975
  91. Boesiger 1974, p. 29
  92. Loison, Laurent (November 2011). "French Roots of French Neo-Lamarckisms, 1879–1985". Journal of the History of Biology. 44 (4): 713–744. doi:10.1007/s10739-010-9240-x. PMID 20665089. S2CID 3398698.
  93. Pearson, Roy Douglas (March 1988). "Reviews". Acta Biotheoretica (Book review). 37 (1): 31–36. doi:10.1007/BF00050806. Book reviews of Animal Evolution in Changing Environments: With Special Reference to Abnormal Metamorphosis (1987) by Ryuichi Matsuda and The Evolution of Individuality (1987) by Leo W. Buss.
  94. Shapiro, Arthur M. (1988). "Book Review: Animal Evolution in Changing Environments with Special Reference to Abnormal Metamorphosis" (PDF). Journal of the Lepidopterists' Society (Book review). 42 (2): 146–147. Retrieved 2015-12-11.
  95. Baird, Scerri & McIntyre 2006, p. 166
  96. Simpson 1944, p. 75
  97. Simpson 1964, pp. 14–60
  98. Simpson 1965, p. 451
  99. Medawar 1985, pp. 166–169
  100. Gardner 1957, pp. 142–143
  101. Mayr 1997, p. 222: "...the recognition that DNA does not directly participate in the making of the phenotype and that the phenotype, in turn, does not control the composition of the DNA represents the ultimate invalidation of all theories involving the inheritance of acquired characters. This definitive refutation of Lamarck's theory of evolutionary causation clears the air."
  102. Bowler 2013, p. 21
  103. Steele, E. J. (2016). "Somatic hypermutation in immunity and cancer: Critical analysis of strand-biased and codon-context mutation signatures". DNA Repair. 45 (2016): 1–2 4. doi:10.1016/j.dnarep.2016.07.001. PMID 27449479.
  104. Steele, E. J. (1981). Somatic selection and adaptive evolution: on the inheritance of acquired characters (2nd ed.). University of Chicago Press.
  105. Roth, Tania L.; Lubin, Farah D.; Funk, Adam J.; et al. (May 2009). "Lasting Epigenetic Influence of Early-Life Adversity on the BDNF Gene". Biological Psychiatry. 65 (9): 760–769. doi:10.1016/j.biopsych.2008.11.028. PMC 3056389. PMID 19150054.
  106. Arai, Junko A.; Shaomin Li; Hartley, Dean M.; et al. (February 4, 2009). "Transgenerational Rescue of a Genetic Defect in Long-Term Potentiation and Memory Formation by Juvenile Enrichment". The Journal of Neuroscience. 29 (5): 1496–1502. doi:10.1523/JNEUROSCI.5057-08.2009. PMC 3408235. PMID 19193896.
  107. Hackett, Jamie A.; Sengupta, Roopsha; Zylicz, Jan J.; et al. (January 25, 2013). "Germline DNA Demethylation Dynamics and Imprint Erasure Through 5-Hydroxymethylcytosine". Science. 339 (6118): 448–452. Bibcode:2013Sci...339..448H. doi:10.1126/science.1229277. PMC 3847602. PMID 23223451.
  108. Bonduriansky, Russell (June 2012). "Rethinking heredity, again". Trends in Ecology & Evolution. 27 (6): 330–336. doi:10.1016/j.tree.2012.02.003. PMID 22445060.
  109. Skinner, Michael K. (May 2015). "Environmental Epigenetics and a Unified Theory of the Molecular Aspects of Evolution: A Neo-Lamarckian Concept that Facilitates Neo-Darwinian Evolution". Genome Biology and Evolution. 7 (5): 1296–1302. doi:10.1093/gbe/evv073. PMC 4453068. PMID 25917417.
  110. Gregory, T. Ryan (March 8, 2009). "Lamarck didn't say it, Darwin did". Genomicron (Blog). Archived from the original on 1 February 2015. Retrieved 2015-11-04.
  111. Wilkins 2009, pp. 295–315
  112. Burkhardt, Richard W. Jr. (August 2013). "Lamarck, Evolution, and the Inheritance of Acquired Characters". Genetics. 194 (4): 793–805. doi:10.1534/genetics.113.151852. PMC 3730912. PMID 23908372.
  113. Penny, David (June 2015). "Epigenetics, Darwin, and Lamarck". Genome Biology and Evolution. 7 (6): 1758–1760. doi:10.1093/gbe/evv107. PMC 4494054. PMID 26026157.
  114. Jablonka & Lamb 1995
  115. Moore 2015
  116. Richards, Eric J. (May 2006). "Inherited epigenetic variation — revisiting soft inheritance". Nature Reviews Genetics. 7 (5): 395–401. doi:10.1038/nrg1834. PMID 16534512. S2CID 21961242.
  117. Nätt, Daniel; Lindqvist, Niclas; Stranneheim, Henrik; et al. (July 28, 2009). Pizzari, Tom (ed.). "Inheritance of Acquired Behaviour Adaptations and Brain Gene Expression in Chickens". PLOS ONE. 4 (7): e6405. Bibcode:2009PLoSO...4.6405N. doi:10.1371/journal.pone.0006405. PMC 2713434. PMID 19636381.
  118. Sheau-Fang Ng; Lin, Ruby C. Y.; Laybutt, D. Ross; et al. (October 21, 2010). "Chronic high-fat diet in fathers programs β-cell dysfunction in female rat offspring". Nature. 467 (7318): 963–966. Bibcode:2010Natur.467..963N. doi:10.1038/nature09491. PMID 20962845. S2CID 4308799.
  119. Gibson, Andrea (June 16, 2013). "Obese male mice father offspring with higher levels of body fat" (Press release). Ohio University. Retrieved 2015-11-02.
  120. Lumey, Lambert H.; Stein, Aryeh D.; Ravelli, Anita C. J. (July 1995). "Timing of prenatal starvation in women and birth weight in their first and second born offspring: The Dutch famine birth cohort study". European Journal of Obstetrics & Gynecology and Reproductive Biology. 61 (1): 23–30. doi:10.1016/0028-2243(95)02149-M. PMID 8549843. INIST:3596539.
  121. Akimoto, Keiko; Katakami, Hatsue; Hyun-Jung Kim; et al. (August 2007). "Epigenetic Inheritance in Rice Plants". Annals of Botany. 100 (2): 205–217. doi:10.1093/aob/mcm110. PMC 2735323. PMID 17576658.
  122. Sano, Hiroshi (April 2010). "Inheritance of acquired traits in plants: Reinstatement of Lamarck". Plant Signaling & Behavior. 5 (4): 346–348. doi:10.4161/psb.5.4.10803. PMC 2958583. PMID 20118668.
  123. Singer, Emily (February 4, 2009). "A Comeback for Lamarckian Evolution?". MIT Technology Review (Biomedicine news).
  124. Rechavi, Oded; Minevich, Gregory; Hobert, Oliver (December 9, 2011). "Transgenerational Inheritance of an Acquired Small RNA-Based Antiviral Response in C. Elegans". Cell. 147 (6): 1248–1256. doi:10.1016/j.cell.2011.10.042. PMC 3250924. PMID 22119442.
  125. Rechavi, O.; Houri-Ze'evi, L.; Anava, S.; Goh, W.S.; Kerk, S.Y.; Hannon, G.J.; Hobert, O. (17 July 2014). "Starvation-induced transgenerational inheritance of small RNAs in C. elegans". Cell. 158 (2): 277–287. doi:10.1016/j.cell.2014.06.020. PMC 4377509. PMID 25018105.
  126. Handel, Adam E.; Ramagopalan, Sreeram V. (May 13, 2010). "Is Lamarckian evolution relevant to medicine?". BMC Medical Genetics. 11: 73. doi:10.1186/1471-2350-11-73. PMC 2876149. PMID 20465829.
  127. Springer & Holley 2013, p. 94
  128. Koonin, Eugene V.; Wolf, Yuri I. (November 11, 2009). "Is evolution Darwinian or/and Lamarckian?". Biology Direct. 4: 42. doi:10.1186/1745-6150-4-42. PMC 2781790. PMID 19906303.
  129. Koonin, Eugene V. (February 2019). "CRISPR: a new principle of genome engineering linked to conceptual shifts in evolutionary biology". Biology & Philosophy. 34 (9): 9. doi:10.1007/s10539-018-9658-7. PMC 6404382. PMID 30930513.
  130. Coyne, Jerry (October 24, 2010). "Epigenetics: the light and the way?". Why Evolution Is True (Blog). Retrieved 2015-11-04.
  131. Coyne, Jerry (September 23, 2013). "Epigenetics smackdown at the Guardian". Why Evolution is True (Blog). Retrieved 2015-11-04.
  132. González-Recio, O.; Toro, M. A.; Bach, A. (2015). "Past, present, and future of epigenetics applied to livestock breeding". Frontiers in Genetics. 6: 305. doi:10.3389/fgene.2015.00305. PMC 4585102. PMID 26442117.
  133. Varona, Luis; Munilla, Sebastián; Mouresan, Elena Flavia; González-Rodríguez, Aldemar; Moreno, Carlos; Altarriba, Juan (2015). "A Bayesian Model for the Analysis of Transgenerational Epigenetic Variation". G3: Genes, Genomes, Genetics. 5 (4): 477–485. doi:10.1534/g3.115.016725. PMC 4390564. PMID 25617408.
  134. Haig, David (June 2007). "Weismann Rules! OK? Epigenetics and the Lamarckian temptation". Biology and Philosophy. 22 (3): 415–428. doi:10.1007/s10539-006-9033-y. S2CID 16322990. Modern neo-Darwinists do not deny that epigenetic mechanisms play an important role during development nor do they deny that these mechanisms enable a variety of adaptive responses to the environment. Recurrent, predictable changes of epigenetic state provide a useful set of switches that allow genetically-identical cells to acquire differentiated functions and allow facultative responses of a genotype to environmental changes (provided that 'similar' changes have occurred repeatedly in the past). However, most neo-Darwinists would claim that the ability to adaptively switch epigenetic state is a property of the DNA sequence (in the sense that alternative sequences would show different switching behavior) and that any increase of adaptedness in the system has come about by a process of natural selection.
  135. Haig, David (November 2011). "Lamarck Ascending!". Philosophy and Theory in Biology (Book essay). 3 (e204). doi:10.3998/ptb.6959004.0003.004. "A Review of Transformations of Lamarckism: From Subtle Fluids to Molecular Biology, edited by Snait B. Gissis and Eva Jablonka, MIT Press, 2011"
  136. Dickins, Thomas E.; Rahman, Qazi (August 7, 2012). "The extended evolutionary synthesis and the role of soft inheritance in evolution". Proceedings of the Royal Society B. 279 (1740): 2913–2921. doi:10.1098/rspb.2012.0273. PMC 3385474. PMID 22593110.
  137. Bowler, Peter J. (1989) [1983]. Evolution: The History of an Idea (Revised ed.). University of California Press. pp. 179, 341. ISBN 978-0520063860.
  138. Whitelaw, Emma (March 27, 2015). "Disputing Lamarckian Epigenetic Inheritance in Mammals". Genome Biology. 16 (60): 60. doi:10.1186/s13059-015-0626-0. PMC 4375926. PMID 25853737.
  139. Weiss, Adam (October 2015). "Lamarckian Illusions". Trends in Ecology & Evolution. 30 (10): 566–568. doi:10.1016/j.tree.2015.08.003. PMID 26411613.
  140. Steele, E. J. (2016). "Somatic hypermutation in immunity and cancer: Critical analysis of strand-biased and codon-context mutation signatures". DNA Repair. 45: 1–24. doi:10.1016/j.dnarep.2016.07.001. PMID 27449479.
  141. Steele, E. J.; Pollard, J. W. (1987). "Hypothesis : Somatic Hypermutation by gene conversion via the error prone DNA-to-RNA-to-DNA information loop". Molecular Immunology. 24 (6): 667–673. doi:10.1016/j.dnarep.2016.07.001. PMID 2443841.
  142. Steele, Lindley & Blanden 1998
  143. Moran, Nancy A.; Sloan, Daniel B. (2015-12-04). "The Hologenome Concept: Helpful or Hollow?". PLOS Biology. 13 (12): e1002311. doi:10.1371/journal.pbio.1002311. PMC 4670207. PMID 26636661.
  144. Rosenberg, Eugene; Sharon, Gill; Zilber-Rosenberg, Ilana (December 2009). "The hologenome theory of evolution contains Lamarckian aspects within a Darwinian framework". Environmental Microbiology. 11 (12): 2959–2962. doi:10.1111/j.1462-2920.2009.01995.x. PMID 19573132.
  145. Depew, David J. (2003), "Baldwin Boosters, Baldwin Skeptics" in: Weber, Bruce H.; Depew, David J. (2003). Evolution and learning: The Baldwin effect reconsidered. MIT Press. pp. 3–31. ISBN 978-0-262-23229-6.
  146. Simpson, George Gaylord (1953). "The Baldwin effect". Evolution. 7 (2): 110–117. doi:10.2307/2405746. JSTOR 2405746.
  147. Dennett, Daniel (2003), "The Baldwin Effect, a Crane, not a Skyhook" in: Weber, Bruce H.; Depew, David J. (2003). Evolution and learning: The Baldwin effect reconsidered. MIT Press. pp. 69–106. ISBN 978-0-262-23229-6.
  148. Kronfeldner, Maria (December 13, 2005). "Is cultural evolution Lamarckian?". Biology & Philosophy. 22 (4): 493–512. doi:10.1007/s10539-006-9037-7. S2CID 85411375.
  149. Cullen 2000, pp. 31–60

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