Jump to content

Michael Graziano

From Wikipedia, the free encyclopedia

Michael Steven Anthony Graziano (born May 22, 1967[1]) is an American scientist and novelist who is currently a professor of Psychology and Neuroscience at Princeton University.[2] His scientific research focuses on the brain basis of awareness. He has proposed the "attention schema" theory, an explanation of how, and for what adaptive advantage, brains attribute the property of awareness to themselves.[3][4] His previous work focused on how the cerebral cortex monitors the space around the body[5][6][7][8][9][10][11] and controls movement within that space.[12][13][14][15][16][17][18][19] Notably he has suggested that the classical map of the body in motor cortex, the homunculus, is not correct and is better described as a map of complex actions that make up the behavioral repertoire.[20] His publications on this topic have had a widespread impact among neuroscientists but have also generated controversy.[21][22][23] His novels[24][25] rely partly on his background in psychology and are known for surrealism or magic realism.[26][27][28] Graziano also composes music including symphonies[29] and string quartets.[30]


Graziano was born in Bridgeport Connecticut in 1967 and spent his childhood in Buffalo, New York. He received his Bachelor of Arts degree from Princeton University in 1989 in Psychology. He attended graduate school in neuroscience at MIT from 1989 to 1991 and then returned to Princeton University to complete his doctoral degree in 1996, in Neuroscience and Psychology. He remained at Princeton University as a postdoctoral researcher and then as a professor of neuroscience and psychology.

Contributions in neuroscience[edit]

Graziano has made contributions in three areas of neuroscience: how neurons in the primate brain encode peripersonal space, how the motor cortex controls complex movement, and the possible neuronal basis of consciousness. These contributions are detailed in the following sections.

Peripersonal space[edit]

In the 1990s, Graziano with Charles Gross described the properties of a set of multisensory neurons in the monkey brain. Building on the work of Hyvarinen and colleagues[31][32] and Rizzolatti and colleagues[33][34] Graziano and Gross described a network of brain areas that appeared to encode the space immediately surrounding the body.[7][8][9][10][11]

Multimodal neurons in the monkey brain that encode the space near the body. Each neuron responds to touching a specific part of the body called the neuron's tactile receptive field. The same neuron responds to visual stimuli in the space near the tactile receptive field. Two examples are depicted. From Graziano MSA and Gross CG (1998) Spatial maps for the control of movement. Current Opinion in Neurobiology, 8: 195 -201.

Each multisensory neuron responded to a touch within a specific "tactile receptive field" on the body surface. Each neuron also responded to a visual stimulus near or approaching the tactile receptive field. The "visual receptive field" was therefore a region of nearby space affixed to the relevant body part. Some neurons responded to sound sources near the tactile receptive field.[7] Some neurons also responded mnemonically, becoming active when a part of the body moved through space and approached the remembered location of an object in the dark.[8] The activity of these multisensory neurons therefore signaled the presence of an object near or touching a part of the body, regardless of whether the object was felt, seen, heard, or remembered.

Electrical stimulation of these multisensory neurons almost always evoked a complex, coordinated movement that resembled a flinching, blocking, or protecting action.[16][17][18][19] Chemical inhibition of these neurons produced a "nerves of steel" state in which defensive reactions were inhibited.[16] Chemical enhancement of these neurons produced a "super flincher" state in which any mild stimulus, such as an object gently moved toward the face, evoked a full-blown flinching reaction.[16]

In Graziano's interpretation,[35] these multisensory neurons form a specialized brain-wide network that encodes the space near the body, computes a margin of safety, and helps to coordinate movements in relation to nearby objects with an emphasis on withdrawal or blocking movements. A subtle level of activation might bias ongoing behavior to avoid collision, whereas a strong level of activation evidently causes an overt defensive action.

The neurons that encode peripersonal space may also provide a neuronal basis for the psychological phenomenon of personal space.[35] Personal space, described by Hall,[36] is the flexible bubble of space around each person that is protected from intrusion by other people.

The peripersonal neurons may also play a central role in the body schema[37] an internally computed model of the body first proposed to exist by Head and Holmes in 1911.[38]

An action map in the motor cortex[edit]

In the 2000s Graziano's lab obtained evidence suggesting that the motor cortex might not contain a simple map of the body's muscles as in classical descriptions such as Penfield's description of a motor homunculus.[39] Instead, the motor cortex may contain a mapping of coordinated, behaviorally useful actions that make up a typical movement repertoire.

In their initial experiments, Graziano and colleagues used electrical microstimulation on the motor cortex of monkeys.[15][19] Most previous protocols in the motor cortex used very brief stimulation, such as for a hundredth of a second. Graziano applied the stimulation for half a second each time, on a behaviorally relevant time scale, in order to match the typical duration of a monkey's reaching and grasping. The longer stimulation train in Graziano's experiments evoked complex movements that included many joints and that resembled movements from the animal's behavioral repertoire.

For example, stimulation of one site always caused the hand to close in a grip, the arm to bring the hand to the mouth, and the mouth to open. Stimulation of another site always caused the grip to open, the palm to face away from the body, and the arm to extend, as if the monkey were reaching to grasp an object. Other sites evoked other complex movements. The behavioral repertoire of the animal seemed to be rendered onto the cortical sheet.

This initial work became controversial because of the method of stimulation on a behavioral time scale. The method was not commonly used in the study of motor cortex[40] although it had been used in the study of other brain regions.[20] That controversy may have partially distracted from the other methods used to study the action map.[20] For example, computational models[12] show that when the complex movement repertoire of a monkey is arranged in a flattened map, with similar movements represented near each other, the map closely resembles the known arrangement of the monkey motor cortex.

In Graziano's proposal, many of the complexities of the motor cortex, such as its overlapping maps of the body and its multiple areas with somewhat different mixtures of properties, may be a result of representing the many parts of the movement repertoire each with its own specialized computational requirements. Graziano[20] suggests that the action-map view does not contradict the more traditional view of motor cortex as a set of fields with differing functions. Instead, the action map may help to explain why motor cortex is divided into functionally distinct fields and why the fields are arranged spatially as they are.

Other researchers have since found a similar, ethological organization to motor cortical regions in monkeys, prosimians, cats, and rats.[21][22][41][42][43] Notwithstanding, direct tests of the idea that the motor cortex contains a movement repertoire have not corroborated this hypothesis.[23] Varying the initial position of the forelimb does not change the muscle synergies evoked by microstimulation of a motor cortical point. Consequently, the evoked movements reach nearly the same final end point and posture, with variability.[23] However, the movement trajectories are quite different depending on the initial limb posture and the starting position of the paw. The evoked movement trajectory is most natural when the forelimb lays pendant ~ perpendicular to the ground (i.e., in equilibrium with the gravitational force). From other starting positions, the movements do not appear natural. The paths of the paw are curved with changes and reversals of direction and the passive influence of the gravitational force on the movements is obvious. These observations demonstrate that while the output of the cortical point evokes a seemingly coordinated limb movement from a rest position, it does not specify a particular movement direction or a controlled trajectory from other initial positions. Thus, in natural conditions a controlled movement must depend on the coordinated activation of a multitude of cortical points, terminating at a final locus of motor cortical activity, which holds the limb at a spatial location.[23]

Basis of consciousness in the brain[edit]

Since 2010 Graziano's lab has studied the brain basis of consciousness. Graziano[44][45] proposed that specialized machinery in the brain computes the feature of awareness and attributes it to other people in a social context. The same machinery, in that hypothesis, also attributes the feature of awareness to oneself. Damage to that machinery disrupts one's own awareness.

The attention schema theory (AST) seeks to explain how an information-processing machine could act the way people do, insisting it has consciousness, describing consciousness in the ways that we do, and claiming that it has an inner magic that transcends mere information-processing, even though it does not. AST is currently being incorporated into artificial intelligence systems through the work of the international Astound project.

The proposed AST was partly motivated by two sets of previous findings.

First, certain regions of the cortex are recruited during social perception as people construct models of other people's minds.[46][47][48][49][50][51][52][53] These regions include, among other areas, the superior temporal sulcus (STS) and the temporoparietal junction (TPJ) bilaterally but with a strong emphasis on the right hemisphere.

Second, when these same regions of cortex are damaged, people suffer from a catastrophic disruption of their own awareness of events and objects around them. The clinical syndrome of hemispatial neglect, or loss of awareness of one side of space, is particularly profound after damage to the TPJ or STS in the right hemisphere.[54][55]

The conjunction of these two previous findings led to the suggestion that awareness may be a computed feature constructed by an expert system in the brain, that at least partly overlaps the TPJ and STS. In that proposal, the feature of awareness can be attributed to other people in the context of social perception. It can also be attributed to oneself, in effect creating one's own awareness.

Why construct the feature of awareness and attribute it to other people? In order to understand and predict the behavior of other people, it is useful to monitor other people's attentional state. Attention is a data handling method by which some signals in the brain are enhanced at the expense of others. According to the AST,[45] when the brain computes that person X is aware of thing Y, it is in effect modeling the state in which person X is applying an attentional enhancement to signal Y. Awareness is an attention schema. In that theory, the same process can be applied to oneself. One's own awareness is a schematized model of one's own attention.


Graziano writes literary novels under his own name and children's novels under the pseudonym B. B. Wurge. His stated reason for the pseudonym is to ensure that children do not accidentally read the wrong category of book.[56] His novels have been praised for their originality, vividness, and surreal imagination.[26][27][28] His book for children, The Last Notebook of Leonardo, won the 2011 Moonbeam Award.

His books include:

Literary Novels:

The Love Song of Monkey (2008)
Cretaceous Dawn (2008)
The Divine Farce (2009)
Death My Own Way (2012)

Children's Novels (written under the name B. B. Wurge):

Billy and the Birdfrogs (2008)
Squiggle (2009)
The Last Notebook of Leonardo (2010)

Books on Neuroscience:

The Intelligent Movement Machine (2008)
God, Soul, Mind, Brain (2010)
Consciousness and the Social Brain (2013)
The Spaces Between Us: A Story of Neuroscience, Evolution, and Human Nature (2018)
Rethinking Consciousness: A Scientific Theory of Subjective Experience (2019)

Books of music:

Three Modern Symphonies (2011)
Symphonies 4, 5, and 6 (2012)
Five String Quartets (2012)


  1. ^ As stated in the catalog of the National Library of Israel.
  2. ^ Princeton University, Department of Psychology, Faculty Information Archived 2010-04-29 at the Wayback Machine
  3. ^ Graziano, Michael S. A. (2013). Consciousness and the Social Brain. Oxford University Press. ISBN 978-0-19-992864-4.
  4. ^ Graziano, MSA; Kastner, S (2011). "Human consciousness and its relationship to social neuroscience: A novel hypothesis". Cognitive Neuroscience. 2 (2): 98–113. doi:10.1080/17588928.2011.565121. PMC 3223025. PMID 22121395.
  5. ^ Aflalo, TN; Graziano, MSA (2008). "Four dimensional spatial reasoning in humans". Journal of Experimental Psychology: Human Perception and Performance. 34 (5): 1066–1077. CiteSeerX doi:10.1037/0096-1523.34.5.1066. PMID 18823195.
  6. ^ Graziano, MSA; Cooke, DF; Taylor, CSR (2000). "Coding the location of the arm by sight". Science. 290 (5497): 1782–1786. Bibcode:2000Sci...290.1782G. doi:10.1126/science.290.5497.1782. PMID 11099420. S2CID 13104435.
  7. ^ a b c Graziano, MSA; Reiss, LAJ; Gross, CG (1999). "A neuronal representation of the location of nearby sounds". Nature. 397 (6718): 428–430. Bibcode:1999Natur.397..428G. doi:10.1038/17115. PMID 9989407. S2CID 4415358.
  8. ^ a b c Graziano, MSA; Hu, XT; Gross, CG (1997). "Coding the locations of objects in the dark". Science. 277 (5323): 239–241. doi:10.1126/science.277.5323.239. PMID 9211852. S2CID 13648729.
  9. ^ a b Graziano, MSA; Hu, XT; Gross, CG (1997). "Visuo-spatial properties of ventral premotor cortex". Journal of Neurophysiology. 77 (5): 2268–2292. doi:10.1152/jn.1997.77.5.2268. PMID 9163357. S2CID 1730131.
  10. ^ a b Graziano, MSA; Yap, GS; Gross, CG (1994). "Coding of visual space by pre-motor neurons". Science. 266 (5187): 1054–1057. Bibcode:1994Sci...266.1054G. CiteSeerX doi:10.1126/science.7973661. PMID 7973661.
  11. ^ a b Graziano, MSA; Gross, CG (1993). "A bimodal map of space: somatosensory receptive fields in the macaque putamen with corresponding visual receptive fields". Experimental Brain Research. 97 (1): 96–109. doi:10.1007/BF00228820. PMID 8131835. S2CID 9387557.
  12. ^ a b Graziano, MSA; Aflalo, TN (2007). "Mapping behavioral repertoire onto the cortex". Neuron. 56 (2): 239–251. doi:10.1016/j.neuron.2007.09.013. PMID 17964243.
  13. ^ Aflalo, TN; Graziano, MSA (2007). "Relationship between unconstrained arm movement and single neuron firing in the macaque motor cortex". Journal of Neuroscience. 27 (11): 2760–2780. doi:10.1523/JNEUROSCI.3147-06.2007. PMC 6672576. PMID 17360898.
  14. ^ Graziano, MSA (2006). "The organization of behavioral repertoire in motor cortex". Annual Review of Neuroscience. 29 (1): 105–134. doi:10.1146/annurev.neuro.29.051605.112924. PMID 16776581. S2CID 7510738.
  15. ^ a b Graziano, MSA; Aflalo, TN; Cooke, DF (2005). "Arm movements evoked by electrical stimulation in the motor cortex of monkeys". Journal of Neurophysiology. 94 (6): 4209–4223. doi:10.1152/jn.01303.2004. PMID 16120657. S2CID 9521256.
  16. ^ a b c d Cooke, DF; Graziano, MSA (2004). "Super-flinchers and nerves of steel: Defensive movements altered by chemical manipulation of a cortical motor area". Neuron. 43 (4): 585–593. doi:10.1016/j.neuron.2004.07.029. PMID 15312656.
  17. ^ a b Cooke, DF; Graziano, MSA (2004). "Sensorimotor integration in the precentral gyrus: Polysensory neurons and defensive movements". Journal of Neurophysiology. 91 (4): 1648–1660. doi:10.1152/jn.00955.2003. PMID 14586035. S2CID 13627099.
  18. ^ a b Cooke, DF; Taylor, CSR; Moore, T; Graziano, MSA (2003). "Complex movements evoked by microstimulation of Area VIP". Proceedings of the National Academy of Sciences of the United States of America. 100 (10): 6163–6168. doi:10.1073/pnas.1031751100. PMC 156343. PMID 12719522.
  19. ^ a b c Graziano, MSA; Taylor, CSR; Moore, T (2002). "Complex movements evoked by microstimulation of precentral cortex". Neuron. 34 (5): 841–851. doi:10.1016/S0896-6273(02)00698-0. PMID 12062029.
  20. ^ a b c d Graziano, M. S. A. (2008). The Intelligent Movement Machine. Oxford University Press. ISBN 978-0195326703.[page needed]
  21. ^ a b Ethier, C; Brizzi, L; Darling, WG; Capaday, C (2006). "Linear summation of cat motor cortex outputs". Journal of Neuroscience. 26 (20): 5574–5581. doi:10.1523/JNEUROSCI.5332-05.2006. PMC 6675291. PMID 16707808.
  22. ^ a b Ramanathan, D; Conner, JM; Tuszynski, MH (2006). "A form of motor cortical plasticity that correlates with recovery of function after brain injury". Proceedings of the National Academy of Sciences of the United States of America. 103 (30): 11370–11375. Bibcode:2006PNAS..10311370R. doi:10.1073/pnas.0601065103. PMC 1544093. PMID 16837575.
  23. ^ a b c d Capaday, Charles (2022-08-11). "Motor cortex outputs evoked by long-duration microstimulation encode synergistic muscle activation patterns not controlled movement trajectories". Frontiers in Computational Neuroscience. 16. doi:10.3389/fncom.2022.851485. ISSN 1662-5188. PMC 9434634. PMID 36062251.
  24. ^ Graziano, M. S. A. (2008). The Love Song of Monkey. Leapfrog Press. ISBN 9780981514802.
  25. ^ Graziano, M. S. A. (2009). The Divine Farce. Leapfrog Press. ISBN 978-1935248040.
  26. ^ a b [1] Archived 2012-03-02 at the Wayback Machine Geekscribe.com, book review from Oct. 18, 2009.
  27. ^ a b "book review from Sept. 14, 2009". Publishers Weekly. Archived from the original on October 29, 2013.
  28. ^ a b "book review from Dec. 3, 2009". Barnstable Patriot. 2009-12-03. Archived from the original on 2015-10-16. Retrieved 2010-01-04.
  29. ^ Graziano, MSA (2011). Three modern symphonies and five character sketches in full score. Teaticket, Massachusetts: Quercus Press. ISBN 978-0982747018.
  30. ^ "Sensory Motor Lab".
  31. ^ Hyvarinen, J. (1981). "Regional distribution of functions in parietal association area 7 of the monkey". Brain Research. 206 (2): 287–303. doi:10.1016/0006-8993(81)90533-3. PMID 7214136. S2CID 53178684.
  32. ^ Hyvarinen, J; Poranen, A (1974). "Function of the parietal associative area 7 as revealed from cellular discharges in alert monkeys". Brain. 97 (4): 673–692. doi:10.1093/brain/97.1.673. PMID 4434188.
  33. ^ Rizzolatti, G; Scandolara, C; Matelli, M; Gentilucci, M. (1981). "Afferent properties of periarcuate neurons in macaque monkeys. II. Visual responses". Behavioural Brain Research. 2 (2): 147–163. doi:10.1016/0166-4328(81)90053-X. PMID 7248055. S2CID 4028658.
  34. ^ Fogassi, L; Gallese, V.; di Pellegrino, G; Fadiga, L; Gentilucci, M; Luppino, M; Pedotti, A; Rizzolatti, G (1992). "Space coding by premotor cortex". Experimental Brain Research. 89 (3): 686–690. doi:10.1007/bf00229894. PMID 1644132. S2CID 38313581.
  35. ^ a b Graziano, MSA; Cooke, DF (2006). "Parieto-frontal interactions, personal space, and defensive behavior". Neuropsychologia. 44 (6): 845–859. doi:10.1016/j.neuropsychologia.2005.09.009. PMID 16277998. S2CID 11368801.
  36. ^ Hall, ET (1966). The Hidden Dimension. Garden City, New York: Anchor Books. ISBN 978-0385084765.[page needed]
  37. ^ Graziano, MSA; Botvinick, MM (2002). "How the brain represents the body: insights from neurophysiology and psychology". {{cite journal}}: Cite journal requires |journal= (help) In: Prinz, W; Hommel, B, eds. (2002). Common Mechanisms in Perception and Action. Attention and Performance. Oxford, UK: Oxford University Press. pp. 136–157. ISBN 978-0198510697.
  38. ^ Head, H; Holmes, G (1911). "Sensory disturbances from cerebral lesions". Brain. 34 (2–3): 102–254. doi:10.1093/brain/34.2-3.102.
  39. ^ Penfield, W; Boldrey, E (1937). "Somatic motor and sensory representation in the cerebral cortex of man as studied by electrical stimulation". Brain. 60 (4): 389–443. doi:10.1093/brain/60.4.389.
  40. ^ Strick, PL (2002). "Stimulating Research on Motor Cortex". Nature Neuroscience. 5 (8): 714–715. doi:10.1038/nn0802-714. PMID 12149622. S2CID 6146202.
  41. ^ Stepniewska, I; Fang, PC; Kaas, JH (2005). "Microstimulation reveals specialized subregions for different complex movements in posterior parietal cortex of prosimian galagos". Proceedings of the National Academy of Sciences of the United States of America. 102 (13): 4878–4883. Bibcode:2005PNAS..102.4878S. doi:10.1073/pnas.0501048102. PMC 555725. PMID 15772167.
  42. ^ Gharbawie, OA; Stepniewska, I; Qi, H; Kaas, JH (2011). "Multiple parietal-frontal pathways mediate grasping in macaque monkeys". Journal of Neuroscience. 31 (32): 11660–11677. doi:10.1523/JNEUROSCI.1777-11.2011. PMC 3166522. PMID 21832196.
  43. ^ Haiss, F; Schwartz, C (2005). "Spatial segregation of different modes of movement control in the whisker representation of rat primary motor cortex". Journal of Neuroscience. 25 (6): 1579–1587. doi:10.1523/JNEUROSCI.3760-04.2005. PMC 6726007. PMID 15703412.
  44. ^ Graziano, MSA (2010). God, Soul, Mind, Brain: A Neuroscientist's Reflections on the Spirit World. Teaticket MA: Leapfrog Press. ISBN 9781935248118.[page needed]
  45. ^ a b Graziano, MSA; Kastner, S (2011). "Human consciousness and its relationship to social neuroscience: A novel hypothesis". Cog Neurosci. 2 (2): 98–113. doi:10.1080/17588928.2011.565121. PMC 3223025. PMID 22121395.
  46. ^ Brunet, E; Sarfati, Y; Hardy-Baylé, MC; Decety, J (2000). "A PET investigation of the attribution of intentions with a nonverbal task". NeuroImage. 11 (2): 157–166. doi:10.1006/nimg.1999.0525. PMID 10679187. S2CID 11846982.
  47. ^ Ciaramidaro, A; Adenzato, M; Enrici, I; Erk, S; Pia, L; Bara, BG; Walter, H (2007). "The intentional network: how the brain reads varieties of intentions". Neuropsychologia. 45 (13): 3105–3113. doi:10.1016/j.neuropsychologia.2007.05.011. PMID 17669444. S2CID 16379004.
  48. ^ Fletcher, PC; Happé, F; Frith, U; Baker, SC; Dolan, RJ; Frackowiak, RS; Frith, CD (1995). "Other minds in the brain: a functional imaging study of "theory of mind" in story comprehension". Cognition. 57 (2): 109–128. doi:10.1016/0010-0277(95)00692-R. hdl:21.11116/0000-0001-A1FA-F. PMID 8556839. S2CID 16321133.
  49. ^ Gallagher, HL; Happé, F; Brunswick, N; Fletcher, PC; Frith, U; Frith, CD (2000). "Reading the mind in cartoons and stories: an fMRI study of 'theory of mind' in verbal and nonverbal tasks". Neuropsychologia. 38 (1): 11–21. doi:10.1016/S0028-3932(99)00053-6. PMID 10617288. S2CID 8933444.
  50. ^ Goel, V; Grafman, J; Sadato, N; Hallett, M (1995). "Modeling other minds". NeuroReport. 6 (13): 1741–1746. doi:10.1097/00001756-199509000-00009. PMID 8541472. S2CID 8560791.
  51. ^ Saxe, R; Kanwisher, N (2003). "People thinking about thinking people: fMRI investigations of theory of mind". NeuroImage. 19 (4): 1835–1842. doi:10.1016/S1053-8119(03)00230-1. PMID 12948738. S2CID 206118958.
  52. ^ Saxe, R; Wexler, A (2005). "Making sense of another mind: the role of the right temporo-parietal junction". Neuropsychologia. 43 (10): 1391–1399. doi:10.1016/j.neuropsychologia.2005.02.013. PMID 15936784. S2CID 2714027.
  53. ^ Vogeley, K; Bussfeld, P; Newen, A; Herrmann, S; Happé, F; Falkai, P; Maier, W; Shah, NJ; et al. (2001). "Mind reading: neural mechanisms of theory of mind and self-perspective". NeuroImage. 14 (1 Pt 1): 170–81. doi:10.1006/nimg.2001.0789. PMID 11525326. S2CID 7053366.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  54. ^ Karnath, HO; Ferber, S; Himmelbach, M (2001). "Spatial awareness is a function of the temporal not the posterior parietal lobe". Nature. 411 (6840): 950–953. Bibcode:2001Natur.411..950K. doi:10.1038/35082075. PMID 11418859. S2CID 4312553.
  55. ^ Vallar, G; Perani, D (1986). "The anatomy of unilateral neglect after right-hemisphere stroke lesions. A clinical/CT-scan correlation study in man". Neuropsychologia. 24 (5): 609–622. doi:10.1016/0028-3932(86)90001-1. PMID 3785649. S2CID 40805768.
  56. ^ "Book Review". Princeton Packet. November 25, 2008.