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Dr. Vilayanur Ramachandran on an episode of PBS's NOVA Television program.

Vilayanur Subramanian Ramachandran (born 1951) is a neuroscientist known for his work in the fields of behavioral neurology and visual psychophysics. He is the Director of the Center for Brain and Cognition,[1][2][3] and is currently a Professor in the Department of Psychology[4] and the Neurosciences Graduate Program[5] at the University of California, San Diego.

Ramachandran is noted for his use of experimental methods that rely relatively little on complex technologies such as neuroimaging. Despite the apparent simplicity of his approach, Ramachandran has generated many new ideas about the brain.[6] He has been called "The Marco Polo of neuroscience" by Richard Dawkins and "the modern Paul Broca" by Eric Kandel.[7] In 1997 Newsweek named him a member of "The Century Club", one of the "hundred most prominent people to watch" in the 21st century.[8] In 2011 Time listed him as one of "the most influential people in the world" on the "Time 100" list.[9][10] Ramachandran is the author of several books that have garnered widespread public interest. These include Phantoms In the Brain (1999) and more recently The Tell-Tale Brain (2010).

Early life and education

Vilayanur Subramanian Ramachandran (in accordance with some Tamil family name traditions, his family name, Vilayanur, is placed first) was born in 1951 in Tamil Nadu, India. His father, V.M. Subramanian, was an engineer who worked for the U.N. Industrial Development Organization and served as a diplomat in Bangkok, Thailand.[11] Ramachandran spent much of his youth moving among several different posts in India and other parts of Asia.[12][13] As a young man Ramachandran attended schools in Madras, and British schools in Bangkok.[14] He pursued many scientific interests, including conchology.[12] Ramachandran obtained an M.B.B.S. from Stanley Medical College in Madras, India, and subsequently obtained a Ph.D. from Trinity College at the University of Cambridge. While a graduate student at Cambridge, Ramachandran also collaborated on research projects with faculty at Oxford, including David Whitteridge of the Physiology Department. He then spent two years at Caltech, as a research fellow working with Jack Pettigrew. He was appointed Assistant Professor of Psychology at the University of California, San Diego in 1983, and has been a full professor there since 1998.

Ramachandran is the grandson of Sir Alladi Krishnaswamy Iyer, Advocate General of Madras and co-architect of the Constitution of India.[13][15] He is married to Diane Rogers-Ramachandran and they have two boys, Mani and Jaya.[12]

Scientific career

In order to understand how the brain works Ramachandran has studied neurological syndromes such as phantom limbs, body integrity identity disorder, and Capgras delusion. He has also contributed to the understanding of synesthesia.[9][12] In addition, Ramachandran is known for the invention of the mirror box.

He has published over 180 papers in scientific journals. Twenty of these have appeared in Nature, and others have appeared in Science, Nature Neuroscience, Perception and Vision Research. Ramachandran is a member of the editorial board of Medical Hypotheses (Elsevier) and has published 15 articles there.[16]

Ramachandran's work in behavioral neurology has been widely reported by the media. He has appeared in numerous Channel 4 and PBS documentaries. He has also been featured by the BBC, the Science Channel, Newsweek, Radio Lab, and This American Life, TED Talks, and Charlie Rose. His book Phantoms in the Brain formed the basis for a two part series on BBC Channel 4 TV (UK) and a 1-hour PBS special in the USA. He is the editor of the Encyclopedia of the Human Brain (2002).

Ramachandran has recently lamented that science has become too professionalized. In a 2010 interview with the British Neuroscience Association he stated: "But where I'd really like to go is back in time. I'd go to the Victorian age, before science had professionalized and become just another 9–5 job, with power-brokering and grants nightmares. Back then scientists just had fun. People like Darwin and Huxley; the whole world was their playground."[17]

In 2012, neuroscientist Peter Brugger criticized Ramachandran's book The Tell-Tale Brain as a pop-neuroscience book that provides vague answers to big questions.[18] Ramachandran responded that "I have—for better or worse—roamed the whole landscape of visual perception, stereopsis, phantom limbs, denial of paralysis, Capgras syndrome, synaesthesia, and many others."[19]

Human vision

Ramachandran’s early research was on human visual perception using psychophysical methods to draw clear inferences about the brain mechanisms underlying visual processing.

Ramachandran is credited with discovering several new visual effects and illusions; most notably perceived slowing of motion at equiluminance (when red and green are seen as equally bright), stereoscopic "capture" using illusory contours, stereoscopic learning, shape-from-shading, and motion capture. He invented (together with Richard Gregory) filling in of "artificial scotomas" and discovered a new "dynamic noise after effect." He also invented a class of stimuli (phantom contours) that selectively activate the magnocellular pathway in human vision and that have been used by Anne Sperling, and her colleagues, to evaluate aspects of dyslexia.[20]

Phantom limbs

Main article: Phantom limb

When an arm or leg is amputated, patients often continue to feel vividly the presence of the missing limb as a "phantom limb". Building on earlier work by Ronald Melzack (McGill University) and Timothy Pons (NIMH), Ramachandran theorized that there was a link between the phenomenon of phantom limbs and neural plasticity in the adult human brain. In particular, he theorized that the body image maps in the somatosensory cortex are re-mapped after the amputation of a limb. In 1993, working with T.T. Yang who was conducting MEG research at the Scripps Research Institute,[21] Ramachandran demonstrated that there had been measurable changes in the somatosensory cortex of several patients who had undergone arm amputations.[22][23] Ramachandran theorized that there was a relationship between the cortical reorganization evident in the MEG images and the referred sensations he observed in his subjects. He presented this theory in a paper titled "Perceptual correlates of massive cortical reorganization."[24] Although Ramachandran was one of the first scientists to emphasize the role of cortical reorganization as the basis for phantom limb sensations, subsequent research has demonstrated that referred sensations are not the perceptual correlate of cortical reorganization after amputation.[25] The question of which neural processes are related to non-painful referred sensations has not been resolved.

Mirror visual feedback

Main article: Mirror box

Ramachandran is credited with the invention of the mirror box and the introduction of mirror visual feedback as a treatment for a variety of conditions associated with phantom limb pain, stroke, and regional pain syndrome. Several research studies using mirror therapy have produced promising results.[26][27] However, mirror therapy has produced conflicting results in randomized controlled trials,[28] and the applications of mirror therapy are still under experimental evaluation.[29]


Synesthetes who experience color when viewing different symbols may quickly identify the presence of the "triangle" in the left-hand image.

Main article: Synesthesia

Ramachandran has studied the neural mechanisms of synesthesia, a condition in which stimulation in one sensory modality leads to experiences in a second, unstimulated modality. His initial studies focused on grapheme → color synesthesia, in which viewing black and white letters or numbers (collectively referred to as graphemes) on a page evokes the experience of seeing colors. Ramachandran (with then PhD student, Edward Hubbard) showed that some synesthetes were better able to detect "embedded figures" composed of one letter or number (for example a triangle composed of 2s) on a background of another number (for example 5s).[30][31]

Based on his previous work on phantom limbs, Ramachandran suggested that synesthesia may arise from a cross-activation between brain regions.[30]

Different subtypes of number–colour synaesthesia...are caused by hyperconnectivity between colour and number areas at different stages in processing; lower synaesthetes may have cross-wiring (or cross-activation) within the fusiform gyrus, whereas higher synaesthetes may have cross-activation in the angular gyrus.[30]

Consistent with this model, Ramachandran's group found increased activity in color selective areas in synesthetes compared to non-synesthetes using fMRI.[31][32] Using MEG, they also showed that differences between synesthetes and non-synesthetes begin very quickly after the grapheme is presented.[33]

However, recent research in the Netherlands (involving 19 grapheme-color synesthetes) demonstrated that synesthesia can be the result of either direct bottom-up cross-activation from grapheme processing areas within the fusiform gyrus, or indirectly via higher-order parietal areas. Tessa van Leeuwen and her colleagues concluded:

To summarize, in this first study of effective connectivity in synesthesia, we established that while the same network of regions is active in different types of synesthesia, individual differences in subjective color experience are determined by altered coupling.[34]

Recently, Ramachandran has also begun investigations of other forms of synesthesia, including number forms[35][36] and tactile → emotion synesthesia.[37] Ramachandran has helped to advance public awareness of synesthesia by hosting two meetings of the American Synesthesia Association at UCSD in 2002[38] and 2011.[39]

Conceptual metaphors

Ramachandran has speculated that synesthesia and conceptual metaphors may share a common basis in cortical cross-activation. In 2003 Ramachandran and Edward Hubbard published a paper in which they speculated that the angular gyrus is at least partially responsible for understanding metaphors.

Recent research by Krish Sathian (Emory University) using functional magnetic resonance imaging (fMRI) suggests that conceptual metaphors activate the texture-selective somatosensory cortex in the parietal operculum.[40] Sathian stated that "I don't think that there's only one area for metaphor processing...several recent lines of research indicate that engagement with abstract concepts is distributed around the brain."[41] Vilayanur Ramachandran commented that "the authors have paved the way" to study how different brain regions communicate. "This is a very ingenious and elegant approach to the problem."[41]

Evolution of language

Following Lakoff and Johnson,[42] Ramachandran argues that metaphors are non-arbitrary. Ramachandran and Hubbard suggest that "these rules [of metaphor production] are a result of strong anatomical constraints that permit certain types of cross-activation, but not others."[30]p. 18 Ramachandran has suggested that the evolution of language is the result of three types of non-arbitrary mappings: between sounds and visual shapes (the bouba-kiki effect), sensory-to-motor synesthesia, and motor-to-motor synesthesia (or "synkinesia").[30]p. 18–23

Mirror neurons

Main article: Mirror neuron

Ramachandran is known for advocating the importance of mirror neurons. Ramachandran has stated that the discovery of mirror neurons is the most important unreported story of the last decade.[43] (Mirror neurons were first reported in a paper published in 1992 by a team of researchers led by Giacomo Rizzolatti at the University of Parma.[44]) In 2000, Ramachandran made a prediction that “mirror neurons will do for psychology what DNA did for biology: they will provide a unifying framework and help explain a host of mental abilities that have hitherto remained mysterious and inaccessible to experiments.”[45][46]

Ramachandran has speculated that research into the role of mirror neurons will help explain a variety of human mental capacities such as empathy, imitation learning, and the evolution of language. Ramachandran has also theorized that mirror neurons may be the key to understanding the neurological basis of human self-awareness.[47][48]

Theories of autism

Main article: Autism#Pathophysiology

In 1999, Ramachandran, in collaboration with then post-doctoral fellow Eric Altschuler and colleague Jaime Pineda, was one of the first to suggest that a loss of mirror neurons might be the key deficit that explains many of the symptoms and signs of autism spectrum disorders.[49] Between 2000 and 2006 Ramachandran and his colleagues at UC San Diego published a number of articles in support of this theory, which became known as the "Broken Mirrors" theory of autism.[50][51][52] Ramachandran and his colleagues did not measure mirror neuron activity directly; rather they demonstrated that children with ASD showed abnormal EEG responses (known as Mu wave suppression) when they observed the activities of other people.

In 2006 Ramachandran conducted an interview with Frontline, India's National Magazine, in which he stated that "One of the things we have discovered in our lab is the cause of the cruel disorder called autism.....the impoverishment of the mirror neuron system explains the symptoms that are unique to autism alone and are not seen in any other short we found the cause for autism in 2000."[53]

Ramachandran's claim that dysfunctional mirror neuron systems (MNS) are the cause of autism remains controversial. In his 2011 review of The Tell-Tale Brain, Simon Baron-Cohen, Director of the Autism Research Center at Cambridge University, states that "As an explanation of autism, the [Broken Mirrors] theory offers some tantalizing clues; however, some problematic counter-evidence challenges the theory and particularly its scope."[54]

Recognizing that dysfunctional mirror neuron systems cannot account for the wide range of symptoms that are included in autism spectrum disorder (ASD), Ramachandran has theorized that childhood temporal lobe epilepsy and olfactory bulb dysgenesis may also play a role in creating the symptoms of ASD. In 2010 Ramachandran stated that "The olfactory bulb hypothesis has important clinical implications" and announced that his group would undertake a study "comparing olfactory bulb volumes in individuals with autism with those of normal controls."[55]

Rare neurological syndromes


Main article: Apotemnophilia

In 2008 Ramachandran, along with David Brang and Paul McGeoch, published the first paper to theorize that apotemnophilia is a neurological disorder caused by damage to the right parietal lobe of the brain.[56] This rare disorder, in which a person desires the amputation of a limb, was first identified by John Money in 1977. Building on Ramachandran's previous work identifying representations of body image in the brain, they argued that this disorder stems from a neural body image that is incomplete. Hence the person sees their limb as a foreign appendage that is outside their body.[56] Ramachandran has extended this theory to suggest that anorexia nervosa may be a body image disorder that has its basis in neurological representations of the body, rather than an appetite disorder of the hypothalamus.[57]

Capgras delusion

Main article: Capgras delusion

In collaboration with then post-doctoral fellow, William Hirstein, Ramachandran published a paper in 1997 in which he presented a theory describing the neural basis of Capgras delusion, a delusion in which family members and other loved ones are thought to be replaced by impostors. Previously, Capgras delusion was attributed to a disconnection between facial recognition and emotional arousal.[58] Ramachandran and Hirstein presented a more specific structural explanation that argued that Capgras delusion might be the result of a disconnection between the "fusiform face area", a region of the fusiform gyrus involved in face perception, and the amygdala, which is involved in the emotional responses to familiar faces. Additionally, based on their model and the specific responses of the patient they examined (a Brazilian man who had sustained a head injury in a traffic accident), Ramachandran and Hirstein proposed a general theory of memory formation. They speculated that a person suffering from Capgras delusion loses the ability to form a taxonomy of memories and hence they can no longer manage memories effectively. Instead of a continuum of memories that constitute a unified sense of self, each memory takes on its own categorical sense of self.[59]

Alternating gender incongruity (AGI)

In 2012, Case and Ramachandran reported the results of a survey of bigendered individuals who experience involuntary alternation between male and female states. Case and Ramachandran hypothesized that gender alternation may reflect an unusual degree (or depth) of hemispheric switching, and the corresponding suppression of sex appropriate body maps in the parietal cortex. They stated that "we hypothesize that tracking the nasal cycle, rate of binocular rivalry, and other markers of hemispheric switching will reveal a physiological basis for AGI individuals' subjective reports of gender switches... We base our hypotheses on ancient and modern associations between the left and right hemispheres and the male and female genders."[60][61]

Awards and honors

Ramachandran was elected to a visiting fellowship at All Souls College, Oxford (1998–1999). In addition he was a Hilgard visiting professor at Stanford University in 2005. He has received honorary doctorates from Connecticut College (2001) and the Indian Institute of Technology, Madras (2004).[62] Ramachandran received the annual Ramon y Cajal award (2004) from the International Neuropsychiatry Society, and the Ariens-Kappers medal from the Royal Netherlands Academy of Sciences for his contributions to Neuroscience (1999). He shared the 2005 Henry Dale Prize with Michael Brady of Oxford, and, as part of the award was elected an honorary life member of the Royal institution for "outstanding research of an interdisciplinary nature".[63] In 2007, the President of India conferred on him the third highest civilian award and honorific title in India, the Padma Bhushan.[64] In 2008, he was listed as number 50 in the Top 100 Public Intellectuals Poll.[65]

Public lectures

Ramachandran is known for his engaging style as a public lecturer. He has presented keynote addresses and public lectures in the U.S., Canada, Britain, Australia and India. He gave the Decade of the Brain lecture at the 25th annual meeting of the Society for Neuroscience in 1995,[66] the Beatty Memorial Talk in 1997 (McGill University),[67] and the Keynote Lecture at the 1999 Decade of the Brain meeting before the NIH and the Library of Congress,[68][69] as well as the Rabindranath Tagore lecture at the Centre for Philosophy and Foundations of Science in New Delhi.[70] In 2003 he gave the annual BBC Reith Lectures.[71] In 2007 he gave a public lecture that was part of a series sponsored by the Templeton Foundation at the Royal Society in London.[72] He gave the 2010 IAS Distinguished Lecture at the University of Bristol's Institute of Advanced Studies dedicated to the memory of his longtime friend and collaborator, Richard Gregory.[73] In October 2011, Ramachandran delivered a lecture titled "The Neurology of Human Nature" at the 47th Nobel Conference at Gustavus Adolphus College in Saint Peter, Minnesota.[74] In 2012, he gave the Gifford Lectures (May 28, 2012 - May 30, 2012) at the University of Glasgow.[75]

Testimony as an expert witness

Ramachandran has served as an expert witness on the delusions associated with pseudocyesis (false pregnancy). At the 2007 trial of Lisa M. Montgomery he testified that Montgomery suffered from severe pseudocyesis disorder and that she was unable to appreciate the nature and quality of her acts.[76][77]

Minotaurasaurus ramachandrani

An interest in paleontology led him to purchase a fossil dinosaur skull from the Gobi Desert, which was named after him as Minotaurasaurus ramachandrani in 2009.[78] A minor controversy surfaced around the provenance of this skull. Some paleontologists claim that this fossil was removed from the Gobi desert without the permission of the Chinese government and sold without proper documentation. V.S. Ramachandran, who purchased the fossil in Tucson, Arizona, says that he would be happy to repatriate the fossil to the appropriate nation, if someone shows him "evidence it was exported without permit". For now, the specimen rests at the Victor Valley Museum, an hour's drive east of Los Angeles.[79]

Books authored

See also


  1. Center for Brain and Cognition website
  2. Ramachandran Bio on CBC website
  3. Psychology Department Webage with link to CBC
  4. UCSD Psychology Faculty Directory
  5. Ramachandran Neurosciences Graduate Program Webpage
  6. Anthony, VS Ramachandran: The Marco Polo of neuroscience, The Observer, January 29, 2011.
  7. A Brief Tour of Human Consciousness, 2004, Back Cover
  8. The Century Club. Newsweek. URL accessed on February 16, 2011.
  9. 9.0 9.1 V.S. Ramachandran - Time 100. URL accessed on April 21, 2011. The citation by Tom Insel, Director of NIH, reads: "Once described as the Marco Polo of neuroscience, V.S. Ramachandran has mapped some of the most mysterious regions of the mind. He has studied visual perception and a range of conditions, from synesthesia to autism. V.S. Ramachandran is changing how our brains think about our minds."
  10. In public polling of the people included in the 2011 list, Ramachandran ranked 97 out of 100.[1]
  11. The Science Studio Interview, June 10, 2006, transcript
  12. 12.0 12.1 12.2 12.3 includeonly>Colapinto, J. "Brain Games; The Marco Polo of Neuroscience", May 11, 2009. Retrieved on March 11, 2011. Full text via Lexis-Nexus here
  13. 13.0 13.1 Andrew Anthony. date = January 30, 2011 VS Ramachandran: The Marco Polo of neuroscience. URL accessed on August 8, 2011.
  14. Ramachandran V.S.,The Making of a Scientist,essay included in Curious Minds:How a Child Becomes a Scientist,page 211[2]
  15. includeonly>Ravi, Y.V.. "Legal luminary", The Hindu, 2003-09-23. Retrieved on 2011-04-21.
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  17. Bulletin, BNA, page 14, issue 62, Autumn 2010
  18. Brugger, Peter, Book Review, Cognitive Neuropsychiatry, Vol. 17, Issue 4, 2012
  19. Ramachandran,V.S. Author Response, Cognitive Neuropsychiatry, Vol. 17, Issue 4, 2012
  20. Sperling et al., Selective magnocellular deficits in dyslexia: a "phantom contour" study, Neuropsychologia, 41,(2003)1422-1429
  21. Yang, UCSD Faculty web page
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  23. For a competing view, see: Flor et al., Nature Reviews, Vol 7, November 2006 [3]
  24. Ramachandran, Rogers-Ramachandran, Stewart, Perceptual correlates of massive cortical reorganization, Science, 1992, Nov 13, 1159-1160
  25. Reprogramming the cerebral cortex: plasticity following central and peripheral lesions, Oxford, 2006, Edited by Stephen Lomber, pages 334
  26. Ramachandran VS, Rogers-Ramachandran D (April 1996). Synaesthesia in phantom limbs induced with mirrors. Proc. Biol. Sci. 263 (1369): 377–86.
  27. Chan, B; Witt, R; Charrow, A; Magee, A; Howard, R; Pasquina, P. Mirror Therapy for Phantom Limb Pain, N Engl J Med 2007; 357:2206–2207November 22, 2007.
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  30. 30.0 30.1 30.2 30.3 30.4 Ramachandran VS and Hubbard EM (2001). Synaesthesia: A window into perception, thought and language. Journal of Consciousness Studies 8 (12): 3–34.
  31. 31.0 31.1 Hubbard EM, Arman AC, Ramachandran VS, Boynton GM (March 2005). Individual differences among grapheme-color synesthetes: brain-behavior correlations. Neuron 45 (6): 975–85.
  32. Hubbard EM, Ramachandran VS (2005). Neurocognitive mechanisms of synesthesia. Neuron 48 (3): 509–520.
  33. Brang D, Hubbard EM, Coulson S, Huang M, Ramachandran VS (2010). Magnetoencephalography reveals early activation of V4 in grapheme-color synesthesia. Neuroimage 53 (1): 268–274.
  34. Leeuwen,T, den Ouden,H, Hagoort,P, Effective Connectivity Determines the Nature of Subjective Experience in Graphene-Color Synesthesia, The Journal of Neuroscience, May 17, 2011 [4]
  35. Brang D, Teuscher U, Ramachandran VS, Coulson S. (2010). Temporal sequences, synesthetic mappings, and cultural biases: the geography of time. Consciousness and Cognition 19 (1): 311–320.
  36. Teuscher U, Brang D, Ramachandran VS, Coulson S. (2010). Spatial cueing in time-space synesthetes: An event-related brain potential study. Brain and Cognition 74 (1): 35–46.
  37. Ramachandran VS, Brang D. (2008). Tactile-emotion synesthesia. Neurocase 14 (5): 390–399.
  38. UCSD ASA Program. American Synesthesia Association. URL accessed on June 6, 2011.
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  41. 41.0 41.1 Metaphorically Feeling:Warren, Tracy,
  42. Lakoff, G & Johnson, M (1980). Metaphors We Live By, Chicago, IL: University of Chicago Press.
  43. Ramachandran, V.S.. Mirror neurons and imitation learning as the driving force behind "the great leap forward" in human evolution. Edge Foundation web site. URL accessed on October 19, 2011.
  44. Rizzolatti,G.,Fabbri-Destro,M,"Mirror Neurons:From Discovery to Autism" Experimental Brain Research, (2010)200:223–237 [6]
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  46. Baron-Cohen, Making Sense of the Brain's Mysteries, American Scientist, On-line Book Review, July–August, 2011 [8]
  47. Oberman, L.M. & Ramachandran, V.S. (2008). "Reflections on the Mirror Neuron System: Their Evolutionary Functions Beyond Motor Representation" Pineda, J. A. Mirror Neuron Systems: The Role of Mirroring Processes in Social Cognition, 39–62, Humana Press.
  48. Ramachandran, V.S.. Self Awareness: The Last Frontier. Edge Foundation web site. URL accessed on October 19, 2011.
  49. E.L. Altschuler, A. Vankov, E.M. Hubbard, E. Roberts, V.S. Ramachandran and J.A. Pineda (2000). "Mu wave blocking by observer of movement and its possible use as a tool to study theory of other minds". 30th Annual Meeting of the Society for Neuroscience, Society for Neuroscience. 
  50. Oberman LM, Hubbard EM, McCleery JP, Altschuler EL, Ramachandran VS & Pineda JA. (2005). EEG evidence for mirror neuron dysfunction in autism spectrum disorders.. Cognitive Brain Research 24 (2): 190–198.
  51. Ramachandran, V.S. & Oberman, L.M. (October 16, 2006). Broken Mirrors: A Theory of Autism. Scientific American 295 (5): 62–69.
  52. Oberman LM & Ramachandran VS. (2007). The simulating social mind: the role of the mirror neuron system and simulation in the social and communicative deficits of autism spectrum disorders. Psychological Bulletin 133 (2): 310–327.
  53. Interview with Sashi Kumar, Frontline,Vol 23,Issue 06,Mar 25, 2006
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  57. Ramachandran VS, Brang D, McGeoch PD, Rosar W. (2009). Sexual and food preference in apotemnophilia and anorexia: interactions between 'beliefs' and 'needs' regulated by two-way connections between body image and limbic structures.. Perception 38 (5): 775–777.
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