Lamarckism (or Lamarckian inheritance) is the idea that an organism can pass on characteristics that it acquired during its lifetime to its offspring (also known as heritability of acquired characteristics or soft inheritance). It is named after the French biologist Jean-Baptiste Lamarck (1744–1829), who incorporated the action of soft inheritance into his evolutionary theories as a supplement to his concept of an inherent progressive tendency driving organisms continuously towards greater complexity, in parallel but separate lineages with no extinction. Lamarck did not originate the idea of soft inheritance, which proposes that individual efforts during the lifetime of the organisms were the main mechanism driving species to adaptation, as they supposedly would acquire adaptive changes and pass them on to offspring.
|Theories of Evolution|
When Charles Darwin published his theory of evolution by natural selection in On the Origin of Species, he continued to give credence to what he called "use and disuse inheritance", but rejected other aspects of Lamarck's theories. Later, Mendelian genetics supplanted the notion of inheritance of acquired traits, eventually leading to the development of the modern evolutionary synthesis, and the general abandonment of the Lamarckian theory of evolution in biology. Despite this abandonment, interest in Lamarckism has continued (2009) as studies in the field of epigenetics have highlighted the possible inheritance of behavioral traits acquired by the previous generation  .
Between 1794 and 1796 Erasmus Darwin wrote Zoonomia suggesting "that all warm-blooded animals have arisen 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. Subsequently Jean-Baptiste Lamarck repeated in his Philosophie Zoologique of 1809 the folk wisdom that characteristics which were "needed" were acquired (or diminished) during the lifetime of an organism then passed on to the offspring. He incorporated this mechanism into his thoughts on evolution, seeing it as resulting in the adaptation of life to local environments.
Lamarck founded a school of French Transformationism which included Étienne Geoffroy Saint-Hilaire, and which corresponded with a radical British school of anatomy based in the extramural anatomy schools in Edinburgh which included the surgeon Robert Knox and the comparative anatomist Robert Edmund Grant. In addition, the Regius Professor of Natural History, Robert Jameson, was the probable author of an anonymous paper in 1826 praising "Mr. Lamarck" for explaining how the higher animals had "evolved" from the "simplest worms" – this was the first use of the word "evolved" in a modern sense. As a young student, Charles Darwin was tutored by Grant, and worked with him on marine creatures.
The Vestiges of the Natural History of Creation, authored by Robert Chambers in St Andrews and published anonymously in England in 1844, proposed a theory which combined radical phrenology with Lamarckism, causing political controversy for its radicalism and unorthodoxy, but exciting popular interest and preparing a huge and prosperous audience for Darwin.
Darwin's Origin of Species proposed natural selection as the main mechanism for development of species, but did not rule out a variant of Lamarckism as a supplementary mechanism. Darwin called his Lamarckian hypothesis Pangenesis, and explained it in the final chapter of his book Variation in Plants and Animals under Domestication, after describing numerous examples to demonstrate what he considered to be 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. 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 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.
1880 to 1930 
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 sometimes called the eclipse of Darwinism by some historians of science, because during that time many scientists and philosophers accepted the reality of evolution but doubted whether natural selection was the main evolutionary mechanism. Theories involving the inheritance of acquired characteristics were among the most popular alternatives to natural selection, and scientists who felt that such Lamarckian mechanisms were the key to evolution were called neo-Lamarckians. Proponents included the British botanist George Henslow 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 Packard who studied blind animals living in caves and wrote a book in 1901 about Lamarck and his work. Also included were a number of paleontologists like Edward Drinker Cope and Alpheus Hyatt who felt 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 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. The German zoologist Theodor Eimer combined Larmarckism with ideas about orthogenesis. With the development of the modern synthesis of the theory of evolution and a lack of evidence for either a mechanism or even the heritability of acquired characteristics, Lamarckism largely fell from favor.
In the 1920s, experiments by Paul Kammerer on amphibians, particularly the midwife toad, appeared to find evidence supporting Lamarckism, but his specimens with supposedly acquired black foot-pads were found to have been tampered with. In The Case of the Midwife Toad Arthur Koestler surmised that the tampering had been done by a Nazi sympathiser to discredit Kammerer for his political views, and that his research might actually have been valid. However most biologists believe that Kammerer was a fraud and even among those who believe he was not dishonest most believe that he misinterpreted the results of his experiments.
After 1930 
A form of Lamarckism was revived in the Soviet Union of the 1930s when Trofim Lysenko promoted Lysenkoism which suited the ideological opposition of Joseph Stalin to genetics. This ideologically driven research influenced Soviet agricultural policy which in turn was later blamed for crop failures.
Since 1988 certain scientists have produced work proposing that Lamarckism could apply to single celled organisms. A version of Lamarckian acquisition in higher order animals is still posited in certain branches of psychology, as, for example, in the Jungian racial memory.
Neo-Lamarckism is a theory of inheritance based on a modification and extension of Lamarckism, essentially maintaining the principle that genetic changes can be influenced and directed by environmental factors.
Lamarck's theory 
The identification of Lamarckism with the inheritance of acquired characteristics is regarded by some as an artifact of the subsequent history of evolutionary thought, repeated in textbooks without analysis. Stephen Jay Gould wrote that 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." 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". Gould advocated defining "Lamarckism" more broadly, in line with Lamarck's overall evolutionary theory.
Lamarck incorporated two ideas into his theory of evolution, in his day considered to be generally true:
- Use and disuse – Individuals lose characteristics they do not require (or use) and develop characteristics that are useful.
- Inheritance of acquired traits – Individuals inherit the traits of their ancestors.
Examples of what is traditionally called "Lamarckism" would include:
- Giraffes stretching their necks to reach leaves high in trees (especially Acacias), strengthen and gradually lengthen their necks. These giraffes have offspring with slightly longer necks (also known as "soft inheritance").
- A blacksmith, through his work, strengthens the muscles in his arms. His sons will have similar muscular development when they mature.
Lamarck stated the following two laws:
- 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.
- 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.
- 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.
- 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.
In essence, a change in the environment brings about change in "needs" (besoins), resulting in change in behavior, bringing change in organ usage and development, bringing change in form over time — and thus the gradual transmutation of the species.
However, as historians of science such as Michael Ghiselin and Stephen Jay Gould have pointed out, none of these views were original to Lamarck. On the contrary, Lamarck's contribution was a systematic theoretical framework for understanding evolution. He saw evolution as comprising two processes;
- Le pouvoir de la vie (a complexifying force) - in which the natural, alchemical movements of fluids would etch out organs from tissues, leading to ever more complex construction regardless of the organ's use or disuse. This would drive organisms from simple to complex forms.
- L'influence des circonstances (an adaptive force) - in which the use and disuse of characters led organisms to become more adapted to their environment. This would take organisms sideways off the path from simple to complex, specialising them for their environment.
Current views 
Interest in Lamarckism has recently increased, as several studies in the field of epigenetics have highlighted the possible inheritance of behavioral traits acquired by the previous generation. A recent such study examined foraging behavior in chickens as a function of stress, concluding:
Our findings suggest that unpredictable food access caused seemingly adaptive responses in feeding behavior, which may have been transmitted to the offspring by means of epigenetic mechanisms, including regulation of immune genes. This may have prepared the offspring for coping with an unpredictable environment.... Transmissions of information across generations which does not involve traditional inheritance of DNA-sequence alleles is often referred to as soft inheritance  or 'Lamarckian inheritance'.
The evolution of acquired characteristics has also been shown in human populations who have experienced starvation, resulting in altered gene function in both the starved population and their offspring. The process of DNA methylation is thought to be behind such changes.
In October 2010, further evidence linking food intake to traits inherited by the offspring were shown in a study of rats conducted by several Australian universities. The study strongly suggested that fathers can transfer a propensity for obesity to their daughters as a result of the fathers' food intake, and not their genetics (or specific genes), prior to the conception of the daughter. A "paternal high-fat diet" was shown to cause cell dysfunction in the daughter, which in turn led to obesity for the daughter.
Several historians have argued that Lamarck's name is linked somewhat unfairly to the theory that has come to bear his name, and that Lamarck deserves credit for being an influential early proponent of the concept of biological evolution, far more than for the mechanism of evolution, in which he simply followed the accepted wisdom of his time. Lamarck died 30 years before the first publication of Charles Darwin's Origin of Species. According to Stephen Jay Gould, if Lamarck had been aware of Darwin's proposed mechanism of natural selection, there is no reason to assume he would not have accepted it as a more likely alternative to his own mechanism. Note also that Darwin, like Lamarck, lacked a plausible alternative mechanism of inheritance - the particulate nature of inheritance was only observed by Gregor Mendel somewhat later, and published in 1866. Its full significance was not appreciated until the Modern evolutionary synthesis in the early 1920s. An important point in its favour at the time was that Lamarck's theory contained a mechanism describing how variation is maintained, which Darwin’s own theory lacked.
Several recent studies, one conducted by researchers at MIT and another by researchers at the Tufts University School of Medicine, have rekindled the debate once again. As reported in MIT's Technology Review in February 2009, "The effects of an animal's environment during adolescence can be passed down to future offspring ... The findings provide support for a 200-year-old theory of evolution that has been largely dismissed: Lamarckian evolution, which states that acquired characteristics can be passed on to offspring."
A report investigating the inheritance of resistance to viral infection in the nematode Caenorhabditis elegans suggests that small RNA molecules may be inherited in a non Mendelian fashion and provide resistance to infection.
Lamarckian elements also appear in the hologenome theory of evolution.
Unlike neo-Darwinism, the term neo-Lamarckism refers more to a loose grouping of largely heterodox theories and mechanisms that emerged after Lamarck's time, than to any coherent body of theoretical work.
In the 1920s, Harvard University researcher William McDougall studied the abilities of rats to correctly solve mazes. He found that offspring of rats that had learned the maze were able to run it faster. The first rats would get it wrong 165 times before being able to run it perfectly each time, but after a few generations it was down to 20. McDougall attributed this to some sort of Lamarckian evolutionary process. Oscar Werner Tiegs and Wilfred Eade Agar later showed McDougall's results to be incorrect, caused by poor experimental controls.
At around the same time, Ivan Pavlov, who was also a Lamarckist, claimed to have observed a similar phenomenon in animals being subject to conditioned reflex experiments. He claimed that with each generation, the animals became easier to condition. However, Pavlov never suggested a mechanism to explain these observations.
Ideological neo-Lamarckism 
Neo-Lamarckian versions of evolution were widespread in the late 19th century. According to Peter Bowler the idea that living things could to some degree choose the characteristics that would be inherited allowed them things to be in charge of their own destiny as opposed to the Darwinian view, which made them puppets at the mercy of the environment. Such ideas were more popular than Darwin's 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.
According to Bowler, Edward Drinker Cope was a neo-Lamarckian as he believed that an organism could respond to any challenge by choosing to adapt to a new way of life. In the early 20th century Frederic Wood Jones and Robert Broom had supported a neo-Lamarckian view of human evolution as opposed to the Darwinian view. Neo-Lamarckism often argued for parallel evolution and independent evolution of species. Hermann Klaatsch had relied on a neo-Lamarckian model of evolution to try and explain the origin of bipedalism. Klaatsch's model had claimed that human races had evolved independent of each other and because of this his model had influenced the belief of polygenism. 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.
Proposed mechanisms of Lamarkian inheritance 
Soma to germ-line feedback 
In the 1970s the immunologist Ted Steele, formerly of the University of Wollongong, and colleagues, proposed a neo-Lamarckian mechanism to try 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 that were generated via somatic hyper-mutation in B-cells. The mRNA products of these somatically novel genes were captured by retroviruses endogenous to the B-cells and were then transported through the blood stream where they could breach the soma-germ barrier and retrofect (reverse transcribe) the newly acquired genes into the cells of the germ line. Although Steele was advocating this theory for the better part of two decades, little more than indirect evidence was ever acquired to support it. An interesting attribute of this idea is that it strongly resembles Darwin's own theory of pangenesis, except in the soma to germ line feedback theory, pangenes are replaced with realistic retroviruses.
Epigenetic inheritance 
Forms of 'soft' or epigenetic inheritance within organisms have been suggested as neo-Lamarckian in nature by such scientists as Eva Jablonka and Marion J. Lamb. In addition to 'hard' or genetic inheritance, involving the duplication of genetic material and its segregation during meiosis, there are other hereditary elements that pass into the germ cells also. These include things like methylation patterns in DNA and chromatin marks, both of which regulate the activity of genes. These are considered "Lamarckian" in the sense that they are responsive to environmental stimuli and can differentially affect gene expression adaptively, with phenotypic results that can persist for many generations in certain organisms. Although the reality of epigenetic inheritance is not doubted (as many experiments have validated it), its significance to the evolutionary process is uncertain. Most neo-Darwinians consider epigenetic inheritance mechanisms to be little more than a specialized form of phenotypic plasticity, with no potential to introduce evolutionary novelty into a species lineage.
See also 
- Jean-Baptiste Lamarck
- Baldwinian evolution
- Inheritance of acquired characters
- Obsolete scientific theories
- Marcus Pembrey
- Racial memory
- Ted Steele
- History of evolutionary thought
- Eclipse of Darwinism
- Roth, Tania; Lubin, Funk, Sweatt (February 2009). "Lasting Epigenetic Influence of early-life adversity on the BDNF Gene". Biological Psychiatry 65: 760–769. Retrieved 13 May 2013.
- Arai, J; Li, Hartley, Feig (February 2009). "Transgenerational rescue of a genetic deficit in Long-Term Potentiation and memory formation by juvenile enrichment". The Journal of Neuroscience 29 (5): 1496–1502. Retrieved 13 May 2013.
- Desmond A. & Moore, J. (1991) Darwin Penguin Books p.617 "Darwin was loathe [sic?] to let go of the notion that a well-used and strengthened organ could be inherited"
- Charles Darwin (27 April 1871). "Pangenesis". Nature. A Weekly Illustrated Journal of Science. pp. 502–503. Retrieved 2007-11-08.
- Quammen, David (2006). The Reluctant Mr. Darwin. Atlas Books. p. 216. ISBN 0-393-05981-2.
- Bowler, Peter J. (2003). Evolution:The History of an Idea. University of California Press. pp. 236–244. ISBN 0-520-23693-9.
- Quammen, David (2006). The Reluctant Mr. Darwin. Atlas Books. pp. 218, 220. ISBN 0-393-05981-2.
- Quammen, David (2006). The Reluctant Mr. Darwin. Atlas Books. p. 221. ISBN 0-393-05981-2.
- Bowler, Peter J. (2003). Evolution:The History of an Idea. University of California Press. pp. 245–246. ISBN 0-520-23693-9.
- Herbert Graham Cannon Lamarck and Modern Genetics Greenwood Press Reprint, 1975 ISBN 0-8371-8173-9
- Gould, Stephen J. "Shades of Lamarck", reprinted in The Panda's Thumb (1980) pp.65-71. Quote from page 66.
- Gould, Stephen J. (2002). The Structure of Evolutionary Theory. Harvard: Belknap Harvard. pp. 177–178. ISBN 0-674-00613-5.
- Jean-Baptiste Lamarck Philosophie zoologique ch.7, p.235
- Jean-Baptiste Lamarck Zoological Philosophy trans. Hugh Elliot, 1914, p.113
- The Imaginary Lamarck: a look at bogus "history" in schoolbooks by Michael Ghiselin
- Gould, S.J. (2002) The Structure of Evolutionary Theory
- "Inheritance of Acquired Behaviour Adaptations and Brain Gene Expression in Chickens". Plos One. Retrieved 2011-04-12.
- Richards EJ (2006) Inherited epigenetic variation–revisiting soft inheritance. Nat Rev Genet 7(5): 395–401.
- "Timing of prenatal starvation in women and birth weight in their first and second born offspring : the Dutch famine birth cohort study". Cat.inist.fr. Retrieved 2011-04-12.
- Ng SF, Lin R, Laybutt DR, Barres R, Owens JA, Morris MJ. "Chronic high-fat diet in fathers programs [bgr]-cell dysfunction in female rat offspring : Nature : Nature Publishing Group". Nature. Retrieved 2011-04-12.
- Rechavi O, Minevich G, Hobert O (2011) Transgenerational inheritance of an acquired small RNA-Based antiviral response in C. elegans. Cell 147(6):1248-1256
- McDougall, 1938. British Journal of Psychology 28:321-345
- Pantin, C F A (November 1957). "Oscar Werner Tiegs". Biographical Memiors of Fellows of the Royal Society (The Royal Society) 3: 247. doi:10.1098/rsbm.1957.0017.
- W E Agar, F H Drummond (1935). "First report on a test of McDougall's Lamarckian experiment on the training of rats". Journal of Experimental Biology 12: 191.
- W E Agar, F H Drummond (1942). "Second report on a test of McDougall's Lamarckian experiment on the training of rats". Journal of Experimental Biology 19: 158.
- W E Agar, F H Drummond (1948). "Third report on a test of McDougall's Lamarckian experiment on the training of rats". Journal of Experimental Biology 25: 103.
- W E Agar, F H Drummond, M M Gunson (1954). "Fourth (final) report on a test of McDougall's Lamarckian experiment on the training of rats". Journal of Experimental Biology 31: 308.
- Peter Bowler The eclipse of Darwinism: anti-Darwinian evolution theories in the decades around 1990 1992
- Peter J. Bowler Evolution: the history of an idea 1989 p. 227
- Bernard A. Wood Wiley-Blackwell encyclopedia of human evolution 2011, Blackwell Publishing Ltd
- James Richard Moore, History, Humanity and Evolution: Essays for John C. Greene, 1992, p. 330
- Lamarck's Signature: how retrogenes are changing Darwin's natural selection paradigm. Edward J. Steele, Robyn A. Lindley, Robert V. Blanden. Perseus Books, 1998
- Epigenetic Inheritance and Evolution: The Lamarckian Dimension. Eva Jablonka, Marion J. Lamb. Oxford University Press, 1995
Further reading 
- Burkeman, Oliver. Why everything you've been told about evolution is wrong, The Guardian, March 19, 2010.
- Desmond, Adrian (1989). The Politics of Evolution: Morphology, Medicine, and Reform in Radical London. Chicago: University of Chicago Press. ISBN 0-226-14374-0.
- Gould, Stephen J. (2002). The Structure of Evolutionary Theory. Harvard: Belknap Harvard. pp. 170–197 on Lamarck. ISBN 0-674-00613-5.
- Medawar, Peter (1959). "The threat and the glory". BBC Reith Lectures No. 6.
- Molino, Jean (2000). "Toward an Evolutionary Theory of Music and Language". In Brown, Merker & Wallin (Eds.), The Origins of Music, ISBN 0-262-23206-5.
- Waddington, Conrad (1961). "The human evolutionary system". In: Michael Banton (Ed.), Darwinism and the Study of Society. London: Tavistock.
- Honeywill, Ross (2008). "Lamarck's Evolution: two centuries of genius and jealousy". Murdoch Books, Sydney See website
- Cairns, J., J. Overbaugh, and S. Miller. 1988. Nature 335: 142-145
- Culotta, Elizabeth; "A Boost for 'Adaptive' Mutation", Science, 265:318, 1994.
- Vetsigian K, Woese C, Goldenfeld N. 2006. "Collective Evolution and the Genetic Code." PNAS 103: 10696-10701.
- Hall Barry G., Adaptive Evolution That Requires Multiple Spontaneous Mutations. I. Mutations Involving an Insertion Sequence
- Nonsense in Schoolbooks - The Imaginary Lamarck:Michael T. Ghiselin recounts Lamarck's times and writings.
- Jean-Baptiste Lamarck : works and heritage: an English/French web site edited by Pietro Corsi (Oxford Univ.) and realised by CNRS (France - IT team of CRHST). This web site contents all books, texts, manuscripts and the lamarck's herbarium.
- Guralnick, Rob, et. al (2006). "Jean-Baptiste Lamarck (1744-1829)". A History of Evolutionary Thought. University of California Museum of Paleontology. Retrieved 3 July 2010.
- The Sins of the Fathers, Take 2: "At tributes to Darwin, Lamarckism — inheritance of acquired traits — will be the skunk at the party." By Sharon Begley, Newsweek. From the magazine issue dated January 26, 2009.
- Essay on Meta-Lamarckism