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. He is often incorrectly cited^{[citation needed]} as the founder 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.

After publication of Charles Darwin's theory of natural selection, the importance of individual efforts in the generation of adaptation was considerably diminished. 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. In a wider context, soft inheritance is of use when examining the evolution of cultures and ideas, and is related to the theory of Memetics.

[edit] History

Jean-Baptiste Lamarck

Between 1794 and 1796 Erasmus Darwin wrote Zoönomia 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.^[1] 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.^[2] 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 specimens had been faked by a Nazi sympathiser to discredit Kammerer for his political views.

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.

[edit] Lamarck's theory

The evolution of necks is often used as the example in explanations of Lamarckism.

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."^[3] 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"^[4]. 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:

1. Use and disuse ' Individuals lose characteristics they do not require (or use) and develop characteristics that are useful.
2. 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.

With this in mind, Lamarck has been credited in some textbooks^[which?] and popular culture^[which?] with developing two laws:

1. 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. 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.^[5]

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.^[6][7] On the contrary, Lamarck's contribution was a systematic theoretical framework for understanding evolution. He saw evolution as comprising two processes;

1. 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.
2. 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.

[edit] Current views on "Lamarckism"

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. Some recent notable studies include those made by the University of Linköping, Sweden, which have looked at foraging behavior in chickens as well as stress factors^[8]. The conclusion of the referenced study is as follows:

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 ^[9] or 'Lamarckian inheritance'.^[8]

On a much smaller scale, genetic exchange without reproduction between single-celled organisms has been well-researched and is termed horizontal gene transfer.

The group of researchers at The University of Linköping again highlighted the apparent link between food intake and cross-generational inheritance of acquired traits. This link has been shown before in studies of human populations who have experienced starvation, where epigenetic factors have altered the functioning of genes^[10]. These changed epigenetic factors appear to show traits in the next generation such as an increased occurrence of diabetes. The process of methylation is thought to be behind such changes.

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.^{[citation needed]}

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."

[edit] Neo-Lamarckism

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 children 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.^{[citation needed]} Oscar Werner Tiegs and Wilfred Eade Agar later showed McDougall's results to be incorrect, caused by poor experimental controls.^{[11][12][13][14][15]}

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.

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.^[16]

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 countless experiments have validated it), its significance to the evolutionary process is uncertain. Most neo-Darwinians consider^{[citation needed]} 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.^[17]

[edit] Lamarckism and single-celled organisms

While Lamarckism has been discredited as an evolutionary influence for larger lifeforms, some scientists controversially argue that it can be observed among microorganisms.^[18] Whether such mutations are directed or not also remains in contention. (see Horizontal Gene Transfer)

In 1988, John Cairns at the Radcliffe Infirmary in Oxford, England, and a group of other scientists renewed the Lamarckian controversy (which at that point had been a dead debate for many years).^[19] The group took a mutated strain of E. coli that was unable to consume the sugar lactose and placed it in an environment where lactose was the only food source. They observed over time that mutations occurred within the colony at a rate that suggested the bacteria were overcoming their handicap by altering their own genes. Cairns, among others, dubbed the process adaptive mutation.

If bacteria that had overcome their own inability to consume lactose passed on this "learned" trait to future generations, it could be argued as a form of Lamarckism; though Cairns later chose to distance himself from such a position.^[20] More typically, it might be viewed as a form of ontogenic evolution.

There has been some research into Lamarckism and prions. A group of researchers, for example, discovered that in yeast cells containing a specific prion protein Sup35, the yeast were able to gain new genetic material, some of which gave them new abilities such as resistance to a particular herbicide. When the researchers mated the yeast cells with cells not containing the prion, the trait reappeared in some of the resulting offspring, indicating that some information indeed was passed down, though whether or not the information is genetic is debatable: trace prion amounts in the cells may be passed to their offspring, giving the appearance of a new genetic trait where there is none.^[21]

Finally, there is growing evidence that cells can activate low-fidelity DNA polymerases in times of stress to induce mutations. While this does not directly confer advantage to the organism on the organismal level, it makes sense at the gene-evolution level. While the acquisition of new genetic traits is random, and selection remains Darwinian, the active process of identifying the necessity to mutate is considered to be Lamarckian.

[edit] Lamarckism and societal change

Jean Molino (2000) has proposed that Lamarckian evolution may be accurately applied to cultural evolution. This was also previously suggested by Peter Medawar (1959) and Conrad Waddington (1961). K. N. Laland and colleagues have recently suggested that human culture can be looked upon as an ecological niche like phenomena, where the effects of cultural niche construction are transmissible from one generation to the next. One interpretation of the Meme theory is that memes are both Darwinian and Lamarckian in nature, as in addition to being subject to selection pressures based on their ability to differentially influence Human minds, memes can be modified and the effects of that modification passed on. Richard Dawkins notes (in Blackmore 2000: The Meme machine, page 13), that Memes can be copied in a Lamarckian way (copying of the product) or in a Weismann-type evolutionary way (copying of the instruction) which is much more resistant against changes.

[edit] See also

Evolutionary biology portal

	Book:Evolution
Books are collections of articles that can be downloaded or ordered in print.

• Jean-Baptiste Lamarck
• Baldwinian evolution
• Darwinism
• Epigenetic inheritance
• Epigenetics
• Evolution
• Inheritance of acquired characters
• Lysenkoism
• Memetics
• Obsolete scientific theories
• Orthogenesis
• Marcus Pembrey
• Racial memory
• Ted Steele
• History of evolutionary thought
• Eclipse of Darwinism

[edit] Notes

[edit] Further references

• 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.
• 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

[edit] External links

• 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. http://www.ucmp.berkeley.edu/history/lamarck.html. 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.