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^{[dubious – discuss]}. 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 recently increased, as several studies in the field of epigenetics have highlighted the possible inheritance of behavioral traits acquired by the previous generation. 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.

History[link]

Jean-Baptiste Lamarck

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.^[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]

1880 to 1930[link]

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.^[3] 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.^[4][5] 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.^[4] The German zoologist Theodor Eimer combined Larmarckism with ideas about orthogenesis.^[6] 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.^[7]

After 1930[link]

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.

A late defence of Lamarckism came from the British zoologist Herbert Graham Cannon, he published Lamarck and Modern Genetics in 1959.^[8]

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[link]

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."^[9] 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".^[10] 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.^[11]

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.^[12][13] 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.

Current views[link]

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,^[14] 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 ^[15] or 'Lamarckian inheritance'.^[14]

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.^[16] 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.^[17] 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.^{[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."

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.^[18]

Lamarckian elements also appear in the hologenome theory of evolution.

Neo-Lamarckism[link]

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.^[19] Oscar Werner Tiegs and Wilfred Eade Agar later showed McDougall's results to be incorrect, caused by poor experimental controls.^{[20][21][22][23][24]}

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.^{[citation needed]}

Peter Bowler wrote that Edward Drinker Cope was a neo-Lamarckian as he believed it allowed living things to be in charge of their own destiny as opposed to the Darwinian view of living things being a puppet at the mercy of the environment. Cope's neo-Lamarckism had claimed that the organism could respond to any challenge by choosing to adapt to a new way of life.^[25] The neo-Lamarckian version of evolution was 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 natural directed plan, thus the neo-Lamarckian view of evolution was often advocated by proponents of orthogenesis.^[26]

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.^[27]

Later supporters of Neo-Lamarckism were George Bernard Shaw and Arthur Koestler who both claimed Lamarckism is more humane, and optimistic than Darwinism.^[28]

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.^[29]

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^{[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.^[30]

Lamarckism and single-celled organisms[link]

While Lamarckism has been discredited as an evolutionary influence for larger lifeforms, some scientists controversially argue that it can be observed among microorganisms.^[31] 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).^[32] 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.^[33] 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 crossed 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.^[34]

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.

Lamarckism and societal change[link]

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.

See also[link]

Evolutionary biology portal

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

• Jean-Baptiste Lamarck
• Baldwinian evolution
• Darwinism
• 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

References[link]

Further reading[link]

• 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
• http://www.scientificamerican.com/article.cfm?id=longevity-inheritance-epigenetics

External links[link]

• 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.
• Essay on Meta-Lamarckism

Susan Lindquist
Susan Lindquist
Born	(1949-06-05) June 5, 1949 (age 63) Chicago
Nationality	United States
Fields	Molecular biology
Institutions	Whitehead Institute MIT University of Chicago
Alma mater	Harvard University University of Illinois at Urbana-Champaign
Doctoral advisor	Matthew Meselson[1]
Known for	protein folding heat-shock proteins prions
Notable awards	Genetics Society of America Medal, 2008[2] National Medal of Science, 2010

Susan Lindquist (born June 5, 1949) is a professor of biology at MIT^[3][4] specializing in molecular biology, particularly the protein folding problem^[1][5] within a family of molecules known as heat-shock proteins,^[6][7] and prions. Lindquist is a member and former Director of the Whitehead Institute and was awarded the National Medal of Science in 2010.

Research[link]

Lindquist is best known for her research that provided strong evidence for a new paradigm in genetics based upon the inheritance of proteins with new, self-perpetuating shapes rather than new DNA sequences. This research provided a biochemical framework for understanding other mysteries in biology, such as Alzheimer's disease and Creutzfeldt-Jakob disease. She is considered an expert in protein folding which, as explained by Lindquist in the following excerpt, is an ancient, fundamental problem in biology:

"What do "mad cows", people with neurodegenerative diseases, and an unusual type of inheritance in yeast have in common? They are all experiencing the consequences of misfolded proteins. ... In humans the consequences can be deadly, leading to such devastating illnesses as Alzheimer's Disease. In one case, the misfolded protein is not only deadly to the unfortunate individual in which it has appeared, but it can apparently be passed from one individual to another under special circumstances - producing infectious neurodegenerative diseases such as mad-cow disease in cattle and Creutzfeld-Jacob Disease in humans."

--from "From Mad Cows to 'Psi-chotic' Yeast: A New Paradigm in Genetics," NAS Distinguished Leaders in Science Lecture Series, November 10, 1999.

Lindquist worked on the PSI+ element in yeast (a prion) and how it can act as a switch that hides or reveals numerous mutations throughout the genome, thus acting as an evolutionary capacitor. She also proposed that a heat shock protein, hsp90, may act in the same way, normally preventing phenotypic consequences of genetic changes, but showing all changes at once when the HSP system is overloaded, either pharmacologically or under stressful environmental conditions. Most of these variations are likely to be harmful, but a few unusual combinations may produce valuable new traits, spurring the pace of evolution. Cancer cells too have an extraordinary ability to evolve. Lindquist's lab investigates closely related evolutionary mechanisms involved in the progression of cancerous tumors and in the evolution of antibiotic-resistant fungi.

Recently, Lindquist has made advances in nanotechnology, researching organic amyloid fibers capable of self-organizing into structures smaller than manufactured materials. Her group also developed a yeast “living test tube” model to study protein folding transitions in neurodegenerative diseases and to test therapeutic strategies through high-throughput screening. She is a co-founder of FoldRx, a company developing drug therapies for diseases of protein misfolding and amyloidosis.^[8]

Dr. Lindquist lectures nationally and internationally on a variety of scientific topics. In June 2006, she was the inaugural guest on the "Futures in Biotech" podcast on Leo Laporte's TWiT network.^[9] In 2007, she participated in the World Economic Forum in Davos, Switzerland with other MIT leaders.^[10]

Biography[link]

Although expected to become a housewife by her parents,^[11] Lindquist studied microbiology at the University of Illinois as an undergraduate and received her PhD in biology from Harvard in 1976. She was the Albert D. Lasker Professor of Medical Sciences in the Department of Molecular Genetics and Cell Biology at the University of Chicago, and the Director of the Whitehead Institute for Biomedical Research from 2001-2004. She is currently a member of the Whitehead Institute, a professor of biology at MIT, and an Investigator of the Howard Hughes Medical Institute.^[12]

Lindquist is married to Edward Buckbee and has two adult daughters, one of whom is an alumna of Northwestern University.

Significant Papers[link]

• Jarosz, D. F.; Lindquist, S. (2010). "Hsp90 and Environmental Stress Transform the Adaptive Value of Natural Genetic Variation". Science 330 (6012): 1820–1824. DOI:10.1126/science.1195487. PMID 21205668.  edit
• Alberti, S.; Halfmann, R.; King, O.; Kapila, A.; Lindquist, S. (2009). "A Systematic Survey Identifies Prions and Illuminates Sequence Features of Prionogenic Proteins". Cell 137 (1): 146–158. DOI:10.1016/j.cell.2009.02.044. PMC 2683788. PMID 19345193. //www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=2683788.  edit.
• Gitler, A. D.; Chesi, A.; Geddie, M. L.; Strathearn, K. E.; Hamamichi, S.; Hill, K. J.; Caldwell, K. A.; Caldwell, G. A. et al. (2009). "Α-Synuclein is part of a diverse and highly conserved interaction network that includes PARK9 and manganese toxicity". Nature Genetics 41 (3): 308–315. DOI:10.1038/ng.300. PMC 2683786. PMID 19182805. //www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=2683786.  edit
• Dai, C.; Whitesell, L.; Rogers, A. B.; Lindquist, S. (2007). "Heat Shock Factor 1 is a Powerful Multifaceted Modifier of Carcinogenesis". Cell 130 (6): 1005–1018. DOI:10.1016/j.cell.2007.07.020. PMC 2586609. PMID 17889646. //www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=2586609.  edit
• Cooper, A. A.; Gitler, A.; Cashikar, A.; Haynes, C.; Hill, K.; Bhullar, B.; Liu, K.; Xu, K. et al. (2006). " -Synuclein Blocks ER-Golgi Traffic and Rab1 Rescues Neuron Loss in Parkinson's Models". Science 313 (5785): 324–328. DOI:10.1126/science.1129462. PMC 1983366. PMID 16794039. //www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1983366.  edit
• Cowen, L. E.; Lindquist, S. (2005). "Hsp90 Potentiates the Rapid Evolution of New Traits: Drug Resistance in Diverse Fungi". Science 309 (5744): 2185–2189. DOI:10.1126/science.1118370. PMID 16195452.  edit
• Krishnan, R.; Lindquist, S. L. (2005). "Structural insights into a yeast prion illuminate nucleation and strain diversity". Nature 435 (7043): 765–772. DOI:10.1038/nature03679. PMC 1405905. PMID 15944694. //www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1405905.  edit
• Si, K.; Lindquist, S.; Kandel, E. (2003). "A neuronal isoform of the aplysia CPEB has prion-like properties". Cell 115 (7): 879–891. DOI:10.1016/S0092-8674(03)01020-1. PMID 14697205.  edit
• Queitsch, C.; Sangster, T. A.; Lindquist, S. (2002). "Hsp90 as a capacitor of phenotypic variation". Nature 417 (6889): 618–624. DOI:10.1038/nature749. PMID 12050657.  edit
• Serio, T.; Cashikar, A.; Kowal, A.; Sawicki, G.; Moslehi, J.; Serpell, L.; Arnsdorf, M.; Lindquist, S. (2000). "Nucleated conformational conversion and the replication of conformational information by a prion determinant". Science 289 (5483): 1317–1321. DOI:10.1126/science.289.5483.1317. PMID 10958771.  edit
• Patino, M. M.; Liu, J. -J.; Glover, J. R.; Lindquist, S. (1996). "Support for the Prion Hypothesis for Inheritance of a Phenotypic Trait in Yeast". Science 273 (5275): 622–626. DOI:10.1126/science.273.5275.622. PMID 8662547.  edit
• Lindquist, S. (1981). "Regulation of protein synthesis during heat shock". Nature 293 (5830): 311–314. DOI:10.1038/293311a0. PMID 6792546.  edit

Awards[link]

• Elected to the American Academy of Arts and Sciences in 1996.
• Elected to the National Academy of Sciences in 1997.
• Named a fellow of the American Academy of Microbiology in 1997.
• Named Albert D. Lasker Professor of Medical Sciences in 1999.
• Received the Novartis/Drew Award in Biomedical Research in 2000.
• Served as director of the Whitehead Institute for Biomedical Research from 2001-2004.
• Received the Dickson Prize in Medicine in 2002.
• Named one of the 50 most important women in science by Discover Magazine in 2002.
• Awarded the Sigma Xi William Procter Prize for Scientific Achievement in 2006.
• Elected to the Institute of Medicine of the National Academies in 2006.
• Awarded the Genetics Society of America Medal in 2008.^[2]
• Awarded the Otto Warburg Prize in 2008.
• Awarded the FASEB Excellence in Science Award in 2009.
• Awarded the Max-Delbrück-Medal, Berlin, Germany, in 2010.^[13]
• Awarded the Mendel Medal, Genetics Society UK, in 2010.^[14]
• Awarded the National Medal of Science (for 2009) in 2010.^[15]

References[link]

External links[link]

• Susan Lindquist iBioseminar Lectures, I. Protein Folding and Prions, II. Prions and Evolution
• http://www.wi.mit.edu/research/faculty/lindquist.html

M. Night Shyamalan
M. Night Shyamalan at a press conference announcing The Happening in 2008
Born	Manoj Shyamalan (1970-08-06) 6 August 1970 (age 41) Mahé, India[1]
Occupation	Director, producer, screenwriter, actor
Years active	1992–present
Religion	Hinduism
Spouse	Bhavna Vaswani (1993–present)
Website
www.mnightshyamalan.com

Manoj Shyamalan ( /ˈʃæməlɑːn/;^[2]Maṉōj Śyāmaḷaṉ; Malayalam: മനോജ് ശ്യാമളന്‍; Tamil: மனோஜ் ஷியாமளன்; born 6 August 1970), known professionally as M. Night Shyamalan, is an American screenwriter, film director, and producer known for making movies with contemporary supernatural plots. He is also known for filming his movies (and staging his plots) in and around Philadelphia, Pennsylvania, where he was raised.

Most of Shyamalan's commercially successful films were co-produced by Disney's Touchstone Pictures film division.

In 2008, Shyamalan was awarded the Padma Shri by the government of India.^[3]

Early life[link]

M. Night Shyamalan and Bryce Dallas Howard at the Spanish premiere of The Village (in the San Sebastián International Film Festival, 2006).

Shyamalan was born in a Hindu household in Mahé, Pondicherry, India.^[1][4] His father, Nelliate C. Shyamalan, is a Malayali physician from Mahé. His mother, Jayalakshmi, speaks Tamil and is an obstetrician and gynecologist by profession.^[5] In the 1960s, after medical school (at the Jawaharlal Institute of Postgraduate Medical Education & Research in Pondicherry) and the birth of their first child, Veena, his parents moved to the United States. His mother returned to India to spend the last five months of her pregnancy at her parents’ home in Chennai.

Shyamalan spent his first six weeks in Pondicherry, and then was raised in Penn Valley, Pennsylvania, an affluent suburb of Philadelphia. He attended the private Roman Catholic grammar school Waldron Mercy Academy, though he was a Hindu (they chose it for disciplinary reasons), followed by the Episcopal Academy, a private Episcopal high school located at the time in Merion, Pennsylvania. Shyamalan earned the New York University Merit Scholarship in 1988.^[6] Shyamalan is an alumnus of New York University’s Tisch School of the Arts, in Manhattan,^[7] graduating in 1992. It was while studying there that he adopted Night as his second name.^[8]

Shyamalan had an early desire to be a filmmaker when he was given a Super-8 camera at a young age. Though his father wanted him to follow in the family practice of medicine, his mother encouraged him to follow his passion.^[9] By the time he was 17 the Steven Spielberg fan had made 45 home movies. On each DVD release of his films (beginning with The Sixth Sense and with the exception of Lady in the Water), he has included a scene from one of these childhood movies which he feels represents his first attempt at the same kind of film.

Career[link]

M. Night Shyamalan and Mark Wahlberg (right) at the presentation of the film The Happening in Madrid.

Shyamalan made his first film, the semi-autobiographical drama Praying with Anger, while still a student at NYU, using money borrowed from family and friends.^[10] It was screened at the Toronto International Film Festival on September 12, 1992,^[11] and played commercially at one theater for a week in rural Woodstock, Illinois.^[11] When the film debuted at the Toronto Film Festival, Shyamalan was introduced by David Overbey who predicted that the world would see more of Shyamalan in the years to come. Praying with Anger has also been shown on Canadian television. Filmed in Chennai, it is his only film to be shot outside of Pennsylvania.

Shyamalan wrote and directed his second movie, Wide Awake, in 1995, though it was not released until 1998.^[12] His parents were the film's associate producers. The drama dealt with a ten-year-old Catholic schoolboy (Joseph Cross) who, after the death of his grandfather (Robert Loggia), searches for God. The film's supporting cast included Dana Delany and Denis Leary as the boy's parents, as well as Julia Stiles, and Camryn Manheim. Wide Awake was filmed in a school Shyamalan attended as a child^[13] and earned 1999 Young Artist Award nominations for Best Drama, and, for Cross, Best Performance.^[14] Only in limited release, the film grossed $305,704 in theaters, against a $6 million budget.^[15]

That same year Shyamalan co-wrote the screenplay for Stuart Little with Greg Brooker.

Shyamalan gained international recognition when he wrote and directed 1999's The Sixth Sense, which was a commercial success and later nominated for six Academy Awards, including Best Picture, Best Director and Best Original Screenplay. He followed The Sixth Sense by writing and directing Unbreakable, released in 2000, which received positive reviews. His 2002 film Signs, where he also played Ray Reddy, gained both critical and financial success. His next movie The Village (2004) received mixed reviews from the critics, but turned out to be a financial success. His later fantasy film Lady in the Water (2006) performed worse critically and financially. The film The Happening (2008) was a financial success but also received negative reviews. In 2010, he directed The Last Airbender, based on the Nickelodeon TV show Avatar: The Last Airbender. The Last Airbender (2010) received extremely negative reviews in the United States and won 5 Razzie Awards, but has gone on to make nearly $320 million internationally at the box office. He wrote the story for and produced his latest film, Devil (2010), but did not direct it. The film was not previewed by critics before its release, eventually receiving mixed reviews. Devil was not a blockbuster hit, but has become a commercial success relative to its budget.

Although Shyamalan has stated he is in the middle of directing his next thriller, it has been confirmed that his next film will be a science fiction epic titled After Earth, to star Jaden Smith. When asked about it in an interview, Shyamalan denied the idea of shooting it in 3D but stated he was planning something technically special.

Personal life[link]

In 1993, Shyamalan married psychologist Bhavna Vaswani, a fellow student whom he met at NYU^[16] and with whom he has two daughters. The family resides on a sprawling estate in Willistown, Pennsylvania, near Shyamalan's usual shooting site of Philadelphia. His production company, Blinding Edge Pictures is located in Berwyn, PA.^[17]

Filmography[link]

Other films[link]

In July 2000, on The Howard Stern Show, Shyamalan said he had met with Spielberg and was in early talks to write the script for the fourth Indiana Jones film. This would have given Shyamalan a chance to work with his longtime idol, Steven Spielberg.^[18] After the film fell through, Shyamalan later said it was too "tricky" to arrange and "not the right thing" for him to do.^[19]

Shyamalan's name was linked with the 2001 film Harry Potter and the Sorcerer's Stone, but it conflicted with the production of Unbreakable. In July 2006, while doing press tours for Lady in the Water, Shyamalan had said he was still interested in directing one of the last two Harry Potter films. "The themes that run through it...the empowering of children, a positive outlook...you name it, it falls in line with my beliefs", Shyamalan said. "I enjoy the humor in it. When I read the first Harry Potter and was thinking about making it, I had a whole different vibe in my head of it".^[20][21]

After the release of The Village in 2004, Shyamalan had been planning a film adaptation of Yann Martel's novel Life of Pi with 20th Century Fox, but later backed out so that he could make Lady in the Water. "I love that book. I mean, it's basically [the story of] a kid born in the same city as me [Pondicherry, India] — it almost felt predestined", Shyamalan said. "But I was hesitant because the book has kind of a twist ending. And I was concerned that as soon as you put my name on it, everybody would have a different experience. Whereas if someone else did it, it would be much more satisfying, I think. Expectations, you've got to be aware of them. I'm wishing them all great luck. I hope they make a beautiful movie".^[22]

In July 2008, it was announced that Shyamalan had partnered with Media Rights Capital to form a production company called Night Chronicles. Shyamalan would produce, but not direct, one film a year for three years.^[23] The first of the three films was Devil, a supernatural thriller directed by siblings John and Drew Dowdle. The script was written by Brian Nelson, based on an original idea from Shyamalan.^[24] The movie was about a group of people stuck in an elevator with the devil, and starred Chris Messina.^[25] The next film in the Night Chronicles series will be called Reincarnate. It will be scripted by Chris Sparling and directed by Daniel Stamm.

Television[link]

Sci-Fi Channel[link]

In 2004, Shyamalan was involved in a media hoax with Sci-Fi Channel, which was eventually uncovered by the press. Sci-Fi claimed in its "documentary" special — The Buried Secret of M. Night Shyamalan, shot on the set of The Village — that Shyamalan was legally dead for nearly a half-hour while drowned in a frozen pond in a childhood accident, and that upon being rescued he had experiences of communicating with spirits, fueling an obsession with the supernatural. The Sci-Fi Channel also claimed that Shyamalan had grown "sour" when the "documentary" filmmakers' questions got too personal, and had therefore withdrawn from participating and threatened to sue the filmmakers.

In truth, Shyamalan developed the hoax with Sci-Fi, going so far as having Sci-Fi staffers sign non disclosure agreements with a $5-million fine attached and requiring Shyamalan's office to formally approve each step. Neither the childhood accident nor the supposed rift with the filmmakers ever occurred. The hoax included a non-existent Sci-Fi publicist, "David Westover", whose name appeared on press releases regarding the special. Sci-Fi also fed false news stories to the Associated Press^[26] and Zap2It.com,^[27] among others. A New York Post news item, based on a Sci Fi press release, referred to Shyamalan's attorneys threatening to sue the filmmakers; the attorneys named were non-existent.

After an AP reporter confronted Sci-Fi Channel president Bonnie Hammer at a press conference, Hammer admitted the hoax, saying it was part of a guerrilla marketing campaign to generate pre-release publicity for The Village. This prompted Sci-Fi's parent company, NBC Universal, to state that the undertaking was "not consistent with our policy at NBC. We would never intend to offend the public or the press and we value our relationship with both."^[28]

Critical reception[link]

Collaborations[link]

Actors	The Sixth Sense	Unbreakable	Signs	The Village	Lady in the Water	The Happening
Frank Collison				Y		Y
Bryce Dallas Howard				Y	Y
Cherry Jones			Y	Y
Joaquin Phoenix			Y	Y
Bruce Willis	Y	Y

Other[link]

• James Newton Howard has been Shyamalan's music composer since The Sixth Sense, with Edmund Choi providing music for Praying with Anger and Wide Awake.
• Tak Fujimoto was cinematographer for The Sixth Sense, Signs and The Happening.
• Andrew Mondshein edited Wide Awake and The Sixth Sense, having Barbara Tulliver editing Signs and Lady in the Water, and finally, Conrad Buff edited The Happening and The Last Airbender.
• Sam Mercer produced all of Shyamalan's works since Unbreakable.
• Barry Mendel & Roger Birnbaum produced Unbreakable and The Happening, while Mendel produced The Sixth Sense without Birnbaum.

Criticism and controversy[link]

Shyamalan twists[link]

With the exception of The Sixth Sense, a common criticism of Shyamalan's works is that they feature better direction than screenwriting.^[38][39] He has also been labeled a "one-trick pony" for his continuous use of the "twist" element in his screenplays.^[38] After the release of The Village, Slate's Michael Agger noted that Shyamalan was following "an uncomfortable pattern" of "making fragile, sealed-off movies that fell apart when exposed to outside logic."^[40]

In a May 31, 2008, interview with the London Independent, Shyamalan offered this answer to the question about his "one-trick" movies: "[A common misperception of me is] that all my movies have twist endings, or that they're all scary. All my movies are spiritual and all have an emotional perspective."^[41]

Plagiarism accusations[link]

Robert McIlhinney, a Pennsylvania screenwriter, sued Shyamalan in 2003, alleging similarity of Signs to his unpublished script Lord of the Barrens: The Jersey Devil.^[42][43][44]

In 2004, Margaret Peterson Haddix noted that The Village has numerous similarities to her children's novel Running Out of Time, prompting discussions with publisher Simon & Schuster about filing a lawsuit^[43][44][45]

In response to both allegations, Disney and Shyamalan's production company, Blinding Edge, issued statements calling the claims "meritless".^[45]

See also[link]

• Satelli D'Or Film Festival

References[link]

External links[link]

Wikimedia Commons has media related to: Night Shyamalan

• M. Night Shyamalan at the Internet Movie Database
• M. Night Shyamalan: The Official Site
• M. Night Shyamalan Online
• 2006 M. Night Shyamalan interview (Interview with Jon Niccum)
• Interview with Rajeev Masand on CNN-IBN/ibnlive.com
• M. Night Shyamalan Interview at www.sci-fi-online.com
• 2010 M. Night Shyamalan Time Magazine Interview