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cortex is organized as a map of the natural behavioral repertoire. The complicated, multifaceted nature of the behavioral repertoire results in a complicated, heterogeneous map in cortex, in which different parts of the movement repertoire are emphasized in different cortical subregions. More complex movements such as reaching or climbing require more coordination among body parts, the processing of more complex control variables, the monitoring of objects in the space near the body, and planning several seconds into the future. Other parts of the movement repertoire, such as manipulating an object with the fingers once the object has been acquired, or manipulating an object in the mouth, involve less planning, less computation of spatial trajectory, and more control of individual joint rotations and muscle forces. In this view the more complex movements, especially multi-segmental movements, come to be emphasized in the more anterior part of the motor map because that cortex emphasizes the musculature of the back and neck which serves as the coordinating link between body parts. In contrast the simpler parts of the movement repertoire that tend to focus more on the distal musculature are emphasized in the more
456:
dorsal premotor cortex. When people made internally paced sequences of movements, more blood flow was measured in the supplementary motor area. When people made simple movements that required little planning, such as palpating an object with the hand, the blood flow was more limited to the primary motor cortex. By implication, the primary motor cortex was more involved in execution of simple movement, the premotor cortex was more involved in sensory guided movement, and the supplementary motor area was more involved in internally generated movements.
420:
divided into 6a-alpha (a posterior part adjacent to the primary motor cortex) and 6a-beta (an anterior part adjacent to the prefrontal cortex). These cortical fields formed a hierarchy in which 6a-beta controlled movement at the most complex level, 6a-alpha had intermediate properties, and the primary motor cortex controlled movement at the simplest level. Vogt and Vogt are therefore the original source of the idea of a caudal (6a-alpha) and a rostral (6a-beta) premotor cortex.
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remained active during the few seconds of delay or preparation time before the monkey performed the instructed movement. Neurons in the primary motor cortex showed much less activity during the preparation period and were more likely to be active only during the movement itself. By implication, the dorsal premotor cortex was more involved in planning or preparing for movement and the primary motor cortex more involved in executing movement.
483:. In this alternative view, though movements of lesser complexity are emphasized in the primary motor cortex and movements of greater complexity are emphasized in the caudal premotor cortex, this difference does not necessarily imply a control hierarchy. Instead the regions differ from each other, and contain subregions with differing properties, because the natural movement repertoire itself is heterogeneous.
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the back (caudal premotor cortex). A set of acronyms are commonly used: PMDr (premotor dorsal, rostral), PMDc (premotor dorsal, caudal), PMVr (premotor ventral, rostral), PMVc (premotor ventral, caudal). Some researchers, especially those who study the ventral premotor areas, use a different terminology. Field 7 or F7 denotes PMDr; F2 = PMDc; F5=PMVr; F4=PMVc.
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They are broadly tuned, responding best to one direction of reach and less well to different directions. Electrical stimulation of the PMDc on a behavioral time scale was reported to evoke a complex movement of the shoulder, arm, and hand that resembles reaching with the hand opened in preparation to grasp.
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large-scale meta-analysis of functional imaging studies. A recent prospective fMRI study that was designed to distinguish phonemic and syllable representations in motor codes provided further evidence for this view by demonstrating adaptation effects in the ventral premotor cortex to repeating syllables.
309:(the appearance of the cortex under a microscope), cytohistochemistry (the manner in which the cortex appears when stained by various chemical substances), anatomical connectivity to other brain areas, and physiological properties. These divisions are summarized below in Divisions of the premotor cortex.
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At least three representations of the hand were reported in the motor cortex, one in the primary motor cortex, one in the ventral premotor cortex, and one in the dorsal premotor cortex. By implication, at least three different cortical fields may exist, each one performing its own special function in
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Rizzolatti and colleagues divided the premotor cortex into four parts or fields based on cytoarchitectonics, two dorsal fields and two ventral fields. They then studied the properties of the ventral premotor fields, establishing tactile, visual, and motor properties of a complex nature (summarized in
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Wise and his colleagues studied the dorsal premotor cortex of monkeys. The monkeys were trained to perform a delayed response task, making a movement in response to a sensory instruction cue. During the task, neurons in the dorsal premotor cortex became active in response to the sensory cue and often
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A caveat about the premotor cortex, noted early in its study, is that the hierarchy between the premotor cortex and the primary motor cortex is not absolute. Instead both the premotor cortex and primary motor cortex project directly to the spinal cord, and each has some capability to control movement
423:
Fulton in 1935 helped to solidify the distinction between a primary motor map of the body in area 4 and a higher-order premotor cortex in area 6. His main evidence came from lesion studies in monkeys. It is not clear where the term "premotor" came from or who used it first, but Fulton popularized the
419:
Vogt and Vogt in 1919 also suggested that motor cortex was divided into a primary motor cortex (area 4) and a higher-order motor cortex (area 6) adjacent to it. Furthermore, in their account, area 6 could be divided into 6a (the dorsal part) and 6b (the ventral part). The dorsal part could be further
370:
PMDr may participate in learning to associate arbitrary sensory stimuli with specific movements or learning arbitrary response rules. In this sense, it may resemble the prefrontal cortex more than other motor cortex fields. It may also have some relation to eye movement. Electrical stimulation in the
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The premotor cortex is now generally divided into four sections. First it is divided into an upper (or dorsal) premotor cortex and a lower (or ventral) premotor cortex. Each of these is further divided into a region more toward the front of the brain (rostral premotor cortex) and a region more toward
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of the body. It may also play a role in planning movement, in the spatial guidance of movement, in the sensory guidance of movement, in understanding the actions of others, and in using abstract rules to perform specific tasks. Different subregions of the premotor cortex have different properties and
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and colleagues suggested an alternative principle of organization for the primary motor cortex and the caudal part of the premotor cortex, all regions that project directly to the spinal cord and that were included in the
Penfield and Woolsey definition of M1. In this alternative proposal, the motor
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Roland and colleagues studied the dorsal premotor cortex and the supplementary motor area in humans while blood flow in the brain was monitored in a positron emission scanner. When people made complex sensory-guided movements such as following verbal instructions, more blood flow was measured in the
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were first discovered in area F5 in the monkey brain by
Rizzolatti and colleagues. These neurons are active when the monkey grasps an object. Yet the same neurons become active when the monkey watches an experimenter grasp an object in the same way. The neurons are therefore both sensory and motor.
442:
Given this work by
Penfield on the human brain and by Woolsey on the monkey brain, by the 1960s the idea of a lateral premotor cortex as separate from the primary motor cortex had mainly disappeared from the literature. Instead M1 was considered to be a single map of the body, perhaps with complex
379:
PMVc or F4 is often studied with respect to its role in the sensory guidance of movement. Neurons here are responsive to tactile stimuli, visual stimuli, and auditory stimuli. These neurons are especially sensitive to objects in the space immediately surrounding the body, in so-called peripersonal
471:
For these and other reasons, a consensus has now emerged that the lateral motor cortex does not consist of a single, simple map of the body, but instead contains multiple subregions including the primary motor cortex and several premotor fields. These premotor fields have diverse properties. Some
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PMVr or F5 is often studied with respect to its role in shaping the hand during grasping and in interactions between the hand and the mouth. Electrical stimulation of at least some parts of F5, when the stimulation is applied on a behavioral time scale, evokes a complex movement in which the hand
361:
PMDc is often studied with respect to its role in guiding reaching. Neurons in PMDc are active during reaching. When monkeys are trained to reach from a central location to a set of target locations, neurons in PMDc are active during the preparation for the reach and also during the reach itself.
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The premotor cortex can be distinguished from
Brodmann area 46 of the prefrontal cortex, just anterior to it, by the presence of a fully formed granular layer IV in area 46. The premotor cortex is therefore anatomically a transition between the agranular motor cortex and the granular, six-layered
434:
in 1937 notably disagreed with the idea of a premotor cortex. He suggested that there was no functional distinction between a primary motor and a premotor area. In his view both were part of the same map. The most posterior part of the map, in area 4, emphasized the hand and fingers and the most
438:
Woolsey who studied the motor map in monkeys in 1956 also believed there was no distinction between primary motor and premotor cortex. He used the term M1 for the proposed single map that encompassed both the primary motor cortex and the premotor cortex. He used the term M2 for the medial motor
404:
In the earliest work on the motor cortex, researchers recognized only one cortical field involved in motor control. Campbell in 1905 was the first to suggest that there might be two fields, a "primary" motor cortex and an "intermediate precentral" motor cortex. His reasons were largely based on
344:
In the study of neurolinguistics, the ventral premotor cortex has been implicated in motor vocabularies in both speech and manual gestures. A mental syllabary — a repository of gestural scores for the most highly used syllables in a language — has been linked to the ventral premotor cortex in a
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space. Electrical stimulation of these neurons causes an apparent defensive movement as if protecting the body surface. This premotor region may be part of a larger circuit for maintaining a margin of safety around the body and guiding movement with respect to nearby objects.
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in layer V, whereas giant pyramidal cells are less common and smaller in the premotor cortex. Second, the primary motor cortex is agranular: it lacks a layer IV marked by the presence of granule cells. The premotor cortex is dysgranular: it contains a faint layer IV.
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The connectivity of the premotor cortex is diverse, partly because the premotor cortex itself is heterogenous and different subregions have different connectivity. Generally the premotor cortex has strong afferent (input) and efferent (output) connectivity to the
352:
These subdivisions of premotor cortex were originally described and remain primarily studied in the monkey brain. Exactly how they may correspond to areas of the human brain, or whether the organization in the human brain is somewhat different, is not yet clear.
143:
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The hypothesis of a separate premotor cortex re-emerged and gained ground in the 1980s. Several key lines of research helped to establish the premotor cortex by showing that it had properties distinct from those of the adjacent primary motor cortex.
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even in the absence of the other. Therefore, the two cortical fields operate at least partly in parallel rather than in a strict hierarchy. This parallel relationship was noted as early as 1919 by Vogt and Vogt and also emphasized by Fulton.
1980:
Woolsey, C.N., Settlage, P.H., Meyer, D.R., Sencer, W., Hamuy, T.P. and Travis, A.M. (1952). "Pattern of localization in precentral and "supplementary" motor areas and their relation to the concept of a premotor area".
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project to the spinal cord and may play a direct role in movement control, whereas others do not. Whether these cortical areas are arranged in a hierarchy or share some other more complex relationship is still debated.
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The premotor cortex can be distinguished from the primary motor cortex, Brodmann area 4, just posterior to it, based on two main anatomical markers. First, the primary motor cortex contains giant pyramidal cells called
264:. It occupies part of Brodmann's area 6. It has been studied mainly in primates, including monkeys and humans. The functions of the premotor cortex are diverse and not fully understood. It projects directly to the
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Rizzolatti, G., Camarda, R., Fogassi, L., Gentilucci, M., Luppino, G. and
Matelli, M (1988). "Functional organization of inferior area 6 in the macaque monkey. II. Area F5 and the control of distal movements".
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Gentilucci M, Fogassi L, Luppino G, Matelli M, Camarda R, Rizzolatti G (1988). "Functional organization of inferior area 6 in the macaque monkey. I. Somatotopy and the control of proximal movements".
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presumably emphasize different functions. Nerve signals generated in the premotor cortex cause much more complex patterns of movement than the discrete patterns generated in the primary motor cortex.
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On similar criteria
Brodmann in 1909 also distinguished between his area 4 (coextensive with the primary motor cortex) and his area 6 (coextensive with the premotor cortex).
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Roland, P.E., Skinhoj, E., Lassen, N.A. and Larsen, B. (1980). "Different cortical areas in man in organization of voluntary movements in extrapersonal space".
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Fogassi, L., Gallese, V., Fadiga, L., Luppino, G., Matelli, M. and
Rizzolatti, G (1996). "Coding of peripersonal space in inferior premotor cortex (area F4)".
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Mirror neurons are proposed to be a basis for understanding the actions of others by internally imitating the actions using one's own motor control circuits.
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Rizzolatti, G., Scandolara, C., Matelli, M. and
Gentilucci, J (1981). "Afferent properties of periarcuate neurons in macaque monkeys, II. Visual responses".
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that lies on the lateral surface of the cerebral hemisphere. The medial extension of area 6, onto the midline surface of the hemisphere, is the site of the
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moves to the mouth, closes in a grip, orients such that the grip faces the mouth, the neck turns to align the mouth to the hand, and the mouth opens.
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Matelli, M., Luppino, G. and Rizzolati, G (1985). "Patterns of cytochrome oxidase activity in the frontal agranular cortex of the macaque monkey".
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cortex now commonly known as the supplementary motor area. (Sometimes in modern reviews M1 is incorrectly equated with the primary motor cortex.)
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857:"Neural correlates of reaching decisions in dorsal premotor cortex: specification of multiple direction choices and final selection of action"
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di Pellegrino, G., Fadiga, L., Fogassi, L., Gallese, V. and Rizzolatti, G (1992). "Understanding motor events: a neurophysiological study".
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686:"Topographic organization of corticospinal projections from the frontal lobe: motor areas on the medial surface of the hemisphere"
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Weinrich, M., Wise, S.P. and Mauritz, K.H (1984). "A neurophyiological study of the premotor cortex in the rhesus monkey".
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1516:"Super-flinchers and nerves of steel: Defensive movements altered by chemical manipulation of a cortical motor area"
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751:"Movement representation in the dorsal and ventral premotor areas of owl monkeys: a microstimulation study"
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Boussaoud D (1985). "Primate premotor cortex: modulation of preparatory neuronal activity by gaze angle".
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PMDr can evoke eye movements and neuronal activity in the PMDr can be modulated by eye movement.
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and therefore may play a role in the direct control of behavior, with a relative emphasis on the
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and Cooke, D.F. (2006). "Parieto-frontal interactions, personal space, and defensive behavior".
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cytoarchitectonics. The primary motor cortex contains cells with giant cell bodies known as "
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Fulton, J (1935). "A note on the definition of the "motor" and "premotor" areas".
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Churchland, M.M., Yu, B.M., Ryu, S.I., Santhanam, G. and Shenoy, K.V (2006).
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10.1002/(SICI)1096-9861(19960805)371:4<649::AID-CNE12>3.0.CO;2-E
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The premotor cortex has been divided into finer subregions on the basis of
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Some commonly accepted divisions of the cortical motor system of the monkey
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anterior part, in area 6, emphasized the muscles of the back and neck.
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Histological Studies on the Localization of Cerebral Function
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greater detail above in Divisions of the premotor cortex).
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1875:. Cambridge, Massachusetts: Cambridge University Press.
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Preuss, T.M., Stepniewska, I. and Kaas, J.H (1996).
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515:(1919). "Ergebnisse unserer Hirnforschung".
1809:Rizzolatti, G. & Sinigaglia, C (2010).
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468:relation to the fingers and wrist.
329:. Subcortically it projects to the
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1052:10.1111/j.0953-816X.2003.03181.x
1040:European Journal of Neuroscience
969:The Intelligent Movement Machine
220:Anatomical terms of neuroanatomy
23:
3285:Posterior parahippocampal gyrus
3227:Collateral (temporal+occipital)
34:needs additional citations for
2873:Secondary somatosensory cortex
2565:Ventromedial prefrontal cortex
914:10.1523/JNEUROSCI.3762-05.2006
1:
3488:Poles of cerebral hemispheres
3222:Cingulate (frontal+cingulate)
1474:10.1016/S0896-6273(02)00698-0
2861:Primary somatosensory cortex
2226:10.1016/j.neuron.2007.09.013
1533:10.1016/j.neuron.2004.07.029
1222:10.1016/0166-4328(81)90053-X
874:10.1016/j.neuron.2005.01.027
638:10.1016/0166-4328(85)90068-3
321:, the superior and inferior
2953:Transverse occipital sulcus
1818:Nature Reviews Neuroscience
3538:
3365:Isthmus of cingulate gyrus
3186:Central (frontal+parietal)
2921:Occipital pole of cerebrum
1095:10.1162/jocn.2006.18.6.974
3496:
3022:Transverse temporal gyrus
2885:Posterior parietal cortex
2207:and Aflalo, T.N. (2007).
1706:10.1152/jn.1997.78.4.2226
218:
140:
128:
3143:Inferior temporal sulcus
3064:Superior temporal sulcus
2747:Inferior parietal lobule
2730:Superior parietal lobule
2670:Supplementary motor area
2077:10.1152/jn.1980.43.1.137
2028:10.1152/jn.1980.43.1.118
1869:Campbell, A. W. (1905).
1279:10.1152/jn.1996.76.1.141
1187:10.1152/jn.1995.73.2.886
1152:10.1152/jn.1985.54.3.714
341:among other structures.
319:supplementary motor area
287:supplementary motor area
3148:Inferior temporal gyrus
3092:Occipitotemporal sulcus
3038:Superior temporal gyrus
2931:Lateral occipital gyrus
2682:Supplementary eye field
2423:Inferior frontal sulcus
2418:Superior frontal sulcus
1346:10.1126/science.7973661
999:10.1093/brain/107.2.385
16:Part of the human brain
3409:Fimbria of hippocampus
2475:Superior frontal gyrus
2373:Inferior frontal gyrus
2327:Superior frontal gyrus
1953:10.1093/brain/60.4.389
1923:10.1093/brain/58.2.311
1895:. Leipzig: J.A. Barth.
241:
173:Middle cerebral artery
3396:Hippocampal formation
3255:Parahippocampal gyrus
3069:Middle temporal gyrus
260:just anterior to the
239:
3369:Retrosplenial cortex
3217:Longitudinal fissure
3109:Medial temporal lobe
2786:Intraparietal sulcus
2646:Primary motor cortex
2543:Orbitofrontal cortex
2492:Medial frontal gyrus
2349:Middle frontal gyrus
570:(50–51): 1190–1194.
315:primary motor cortex
262:primary motor cortex
43:improve this article
3346:Posterior cingulate
2756:Supramarginal gyrus
1398:1999Natur.397..428G
1338:1994Sci...266.1054G
1332:(5187): 1054–1057.
576:1926NW.....14.1190V
564:Naturwissenschaften
302:prefrontal cortex.
189:cortex praemotorius
3404:Hippocampal sulcus
3324:Anterior cingulate
3201:Preoccipital notch
2802:Paracentral lobule
2772:Parietal operculum
2694:Frontal eye fields
2626:Paracentral sulcus
2614:Paracentral lobule
2515:Paraolfactory area
2511:Paraterminal gyrus
2175:10.1007/bf00248741
1766:10.1007/bf00230027
1649:10.1007/bf00248742
826:10.1007/bf00231153
584:10.1007/bf01451766
248:is an area of the
242:
3509:
3508:
3453:
3452:
3278:Postrhinal cortex
3273:Perirhinal cortex
3268:Entorhinal cortex
3240:
3239:
3196:Parieto-occipital
3163:
3162:
3001:
3000:
2900:
2899:
2857:Postcentral gyrus
2709:
2708:
2634:
2633:
2454:
2453:
2446:Precentral sulcus
2409:Pars triangularis
1700:(4): 2226–22230.
1392:(6718): 428–430.
1083:J. Cogn. Neurosci
908:(14): 3697–3712.
327:prefrontal cortex
252:lying within the
234:
233:
229:
119:
118:
111:
93:
58:"Premotor cortex"
3529:
3435:Indusium griseum
3300:Cingulate cortex
3290:Prepyriform area
3251:
3176:
3056:Planum temporale
3012:
2993:Calcarine sulcus
2911:
2720:
2591:Olfactory sulcus
2577:Subcallosal area
2463:
2441:Precentral gyrus
2400:Pars opercularis
2315:
2308:
2283:
2276:
2269:
2260:
2253:
2252:
2246:
2238:
2228:
2205:Graziano, M.S.A.
2201:
2195:
2194:
2158:
2152:
2151:
2141:
2124:(9): 1329–1345.
2109:
2103:
2102:
2096:
2088:
2060:
2054:
2053:
2047:
2039:
2011:
2005:
2004:
1998:
1990:
1977:
1971:
1970:
1964:
1956:
1933:
1927:
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1725:
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1682:
1676:
1668:
1631:
1620:
1619:
1613:
1605:
1574:Neuropsychologia
1570:Graziano, M.S.A.
1566:
1560:
1559:
1553:
1545:
1535:
1510:Cooke, D.F. and
1507:
1501:
1500:
1494:
1486:
1476:
1453:Graziano, M.S.A.
1449:
1440:
1439:
1433:
1425:
1382:Graziano, M.S.A.
1378:
1372:
1371:
1365:
1357:
1323:
1315:Graziano, M.S.A.
1311:
1305:
1304:
1298:
1290:
1262:
1256:
1255:
1249:
1241:
1210:Behav. Brain Res
1205:
1199:
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1128:
1122:
1114:
1078:
1072:
1071:
1031:
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1024:
1018:
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982:
973:
972:
965:Graziano, M.S.A.
961:
950:
949:
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893:
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886:
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845:
809:
803:
802:
796:
788:
770:
746:
740:
739:
733:
725:
715:
705:
696:(5): 3284–3306.
681:
672:
671:
665:
657:
626:Behav. Brain Res
621:
610:
609:
603:
595:
552:
539:
538:
532:
524:
505:
481:posterior cortex
337:, and the motor
307:cytoarchitecture
226:edit on Wikidata
145:
133:
121:
114:
107:
103:
100:
94:
92:
51:
27:
19:
3537:
3536:
3532:
3531:
3530:
3528:
3527:
3526:
3522:Cerebral cortex
3512:
3511:
3510:
3505:
3492:
3471:
3449:
3423:
3390:
3294:
3236:
3232:Callosal sulcus
3210:Medial/inferior
3205:
3170:
3159:
3085:Medial/inferior
3080:
3052:Wernicke's area
3026:Auditory cortex
2997:
2962:Medial/inferior
2957:
2896:
2845:
2841:Marginal sulcus
2795:Medial/inferior
2790:
2705:
2658:Premotor cortex
2630:
2600:
2459:Medial/inferior
2450:
2427:
2297:
2291:cerebral cortex
2289:Anatomy of the
2287:
2257:
2256:
2239:
2203:
2202:
2198:
2160:
2159:
2155:
2111:
2110:
2106:
2089:
2065:J. Neurophysiol
2062:
2061:
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2016:J. Neurophysiol
2013:
2012:
2008:
1991:
1979:
1978:
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1957:
1935:
1934:
1930:
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1900:
1885:
1884:
1880:
1868:
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1830:10.1038/nrn2805
1813:
1808:
1807:
1803:
1786:
1751:
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1743:
1726:
1694:J. Neurophysiol
1691:
1690:
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1669:
1633:
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1623:
1606:
1568:
1567:
1563:
1546:
1512:Graziano, M.S.A
1509:
1508:
1504:
1487:
1451:
1450:
1443:
1426:
1380:
1379:
1375:
1358:
1321:
1313:
1312:
1308:
1291:
1267:J. Neurophysiol
1264:
1263:
1259:
1242:
1207:
1206:
1202:
1175:J. Neurophysiol
1172:
1171:
1167:
1140:J. Neurophysiol
1137:
1136:
1132:
1115:
1080:
1079:
1075:
1033:
1032:
1028:
1011:
984:
983:
976:
963:
962:
953:
936:
895:
894:
890:
854:
853:
849:
811:
810:
806:
789:
755:J. Comp. Neurol
748:
747:
743:
726:
683:
682:
675:
658:
623:
622:
613:
596:
554:
553:
542:
525:
507:
506:
497:
492:
486:
449:
402:
386:
377:
368:
359:
323:parietal cortex
283:Brodmann area 6
279:
246:premotor cortex
230:
150:
148:Brodmann area 6
136:
124:Premotor cortex
115:
104:
98:
95:
52:
50:
40:
28:
17:
12:
11:
5:
3535:
3533:
3525:
3524:
3514:
3513:
3507:
3506:
3501:Brodmann areas
3497:
3494:
3493:
3491:
3490:
3485:
3479:
3477:
3473:
3472:
3470:
3469:
3467:Insular cortex
3463:
3461:
3459:Insular cortex
3455:
3454:
3451:
3450:
3448:
3447:
3442:
3437:
3431:
3429:
3425:
3424:
3422:
3421:
3416:
3411:
3406:
3400:
3398:
3392:
3391:
3389:
3388:
3387:
3386:
3381:
3376:
3361:
3360:
3359:
3358:
3353:
3343:
3342:
3341:
3336:
3331:
3321:
3320:
3319:
3312:Subgenual area
3308:
3306:
3296:
3295:
3293:
3292:
3287:
3282:
3281:
3280:
3275:
3270:
3259:
3257:
3248:
3242:
3241:
3238:
3237:
3235:
3234:
3229:
3224:
3219:
3213:
3211:
3207:
3206:
3204:
3203:
3198:
3193:
3188:
3182:
3180:
3173:
3171:sulci/fissures
3165:
3164:
3161:
3160:
3158:
3157:
3156:
3155:
3145:
3139:
3138:
3137:
3136:
3131:
3126:
3121:
3116:
3106:
3105:
3104:
3097:Fusiform gyrus
3094:
3088:
3086:
3082:
3081:
3079:
3078:
3077:
3076:
3066:
3061:
3060:
3059:
3045:
3035:
3034:
3033:
3018:
3016:
3009:
3003:
3002:
2999:
2998:
2996:
2995:
2989:
2988:
2983:
2978:
2977:
2976:
2965:
2963:
2959:
2958:
2956:
2955:
2950:
2945:
2944:
2943:
2938:
2928:
2926:Occipital gyri
2923:
2917:
2915:
2908:
2906:Occipital lobe
2902:
2901:
2898:
2897:
2895:
2894:
2893:
2892:
2882:
2881:
2880:
2870:
2869:
2868:
2853:
2851:
2847:
2846:
2844:
2843:
2838:
2837:
2836:
2826:
2825:
2824:
2819:
2814:
2809:
2798:
2796:
2792:
2791:
2789:
2788:
2782:
2781:
2780:
2779:
2769:
2768:
2767:
2758:
2744:
2743:
2742:
2737:
2726:
2724:
2717:
2711:
2710:
2707:
2706:
2704:
2703:
2702:
2701:
2691:
2690:
2689:
2679:
2678:
2677:
2667:
2666:
2665:
2655:
2654:
2653:
2642:
2640:
2636:
2635:
2632:
2631:
2629:
2628:
2623:
2622:
2621:
2610:
2608:
2602:
2601:
2599:
2598:
2596:Orbital sulcus
2593:
2587:
2586:
2585:
2584:
2574:
2573:
2572:
2562:
2561:
2560:
2555:
2550:
2536:
2535:
2534:
2527:Straight gyrus
2524:
2523:
2522:
2507:
2506:
2505:
2504:
2499:
2489:
2488:
2487:
2482:
2471:
2469:
2460:
2456:
2455:
2452:
2451:
2449:
2448:
2443:
2437:
2435:
2429:
2428:
2426:
2425:
2420:
2414:
2413:
2412:
2411:
2402:
2388:
2386:Pars orbitalis
2379:
2369:
2368:
2367:
2366:
2361:
2356:
2346:
2345:
2344:
2339:
2334:
2323:
2321:
2312:
2305:
2299:
2298:
2288:
2286:
2285:
2278:
2271:
2263:
2255:
2254:
2219:(2): 239–251.
2196:
2169:(3): 475–490.
2163:Exp. Brain Res
2153:
2104:
2071:(1): 137–150.
2055:
2022:(1): 118–136.
2006:
1972:
1947:(4): 389–443.
1928:
1917:(2): 311–316.
1898:
1878:
1861:
1824:(4): 264–274.
1801:
1760:(1): 176–180.
1754:Exp. Brain Res
1741:
1684:
1643:(3): 491–507.
1637:Exp. Brain Res
1621:
1580:(6): 845–859.
1561:
1526:(4): 585–593.
1502:
1467:(5): 841–851.
1441:
1373:
1306:
1273:(1): 141–157.
1257:
1216:(2): 147–163.
1200:
1181:(2): 886–890.
1165:
1146:(3): 714–734.
1130:
1089:(6): 974–989.
1073:
1046:(3): 721–740.
1026:
993:(2): 385–414.
974:
951:
888:
867:(5): 801–814.
847:
820:(2): 285–302.
814:Exp. Brain Res
804:
761:(4): 649–676.
741:
673:
632:(2): 125–136.
611:
540:
494:
493:
491:
488:
448:
445:
401:
398:
393:Mirror neurons
385:
382:
376:
373:
367:
364:
358:
355:
278:
275:
232:
231:
222:
216:
215:
210:
204:
203:
198:
192:
191:
186:
180:
179:
175:
174:
171:
165:
164:
161:
157:
156:
152:
151:
146:
138:
137:
134:
126:
125:
117:
116:
31:
29:
22:
15:
13:
10:
9:
6:
4:
3:
2:
3534:
3523:
3520:
3519:
3517:
3504:
3502:
3495:
3489:
3486:
3484:
3481:
3480:
3478:
3474:
3468:
3465:
3464:
3462:
3460:
3456:
3446:
3443:
3441:
3438:
3436:
3433:
3432:
3430:
3426:
3420:
3419:Rhinal sulcus
3417:
3415:
3414:Dentate gyrus
3412:
3410:
3407:
3405:
3402:
3401:
3399:
3397:
3393:
3385:
3382:
3380:
3377:
3375:
3372:
3371:
3370:
3366:
3363:
3362:
3357:
3354:
3352:
3349:
3348:
3347:
3344:
3340:
3337:
3335:
3332:
3330:
3327:
3326:
3325:
3322:
3318:
3315:
3314:
3313:
3310:
3309:
3307:
3305:
3301:
3297:
3291:
3288:
3286:
3283:
3279:
3276:
3274:
3271:
3269:
3266:
3265:
3264:
3261:
3260:
3258:
3256:
3252:
3249:
3247:
3243:
3233:
3230:
3228:
3225:
3223:
3220:
3218:
3215:
3214:
3212:
3208:
3202:
3199:
3197:
3194:
3192:
3189:
3187:
3184:
3183:
3181:
3179:Superolateral
3177:
3174:
3172:
3166:
3154:
3151:
3150:
3149:
3146:
3144:
3141:
3140:
3135:
3132:
3130:
3127:
3125:
3122:
3120:
3117:
3115:
3112:
3111:
3110:
3107:
3103:
3100:
3099:
3098:
3095:
3093:
3090:
3089:
3087:
3083:
3075:
3072:
3071:
3070:
3067:
3065:
3062:
3057:
3053:
3049:
3046:
3044:
3041:
3040:
3039:
3036:
3032:
3029:
3028:
3027:
3023:
3020:
3019:
3017:
3015:Superolateral
3013:
3010:
3008:
3007:Temporal lobe
3004:
2994:
2991:
2990:
2987:
2986:Lingual gyrus
2984:
2982:
2979:
2975:
2972:
2971:
2970:
2969:Visual cortex
2967:
2966:
2964:
2960:
2954:
2951:
2949:
2948:Lunate sulcus
2946:
2942:
2939:
2937:
2934:
2933:
2932:
2929:
2927:
2924:
2922:
2919:
2918:
2916:
2914:Superolateral
2912:
2909:
2907:
2903:
2891:
2888:
2887:
2886:
2883:
2879:
2876:
2875:
2874:
2871:
2867:
2864:
2863:
2862:
2858:
2855:
2854:
2852:
2848:
2842:
2839:
2835:
2832:
2831:
2830:
2827:
2823:
2820:
2818:
2815:
2813:
2810:
2808:
2805:
2804:
2803:
2800:
2799:
2797:
2793:
2787:
2784:
2783:
2778:
2775:
2774:
2773:
2770:
2766:
2765:Angular gyrus
2762:
2759:
2757:
2753:
2750:
2749:
2748:
2745:
2741:
2738:
2736:
2733:
2732:
2731:
2728:
2727:
2725:
2723:Superolateral
2721:
2718:
2716:
2715:Parietal lobe
2712:
2700:
2697:
2696:
2695:
2692:
2688:
2685:
2684:
2683:
2680:
2676:
2673:
2672:
2671:
2668:
2664:
2661:
2660:
2659:
2656:
2652:
2649:
2648:
2647:
2644:
2643:
2641:
2637:
2627:
2624:
2620:
2617:
2616:
2615:
2612:
2611:
2609:
2607:
2603:
2597:
2594:
2592:
2589:
2588:
2583:
2580:
2579:
2578:
2575:
2571:
2568:
2567:
2566:
2563:
2559:
2556:
2554:
2551:
2549:
2546:
2545:
2544:
2540:
2537:
2533:
2530:
2529:
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99:December 2009
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60: –
59:
55:
54:Find sources:
48:
44:
38:
37:
32:This article
30:
26:
21:
20:
3498:
3262:
2657:
2539:Orbital gyri
2391:Broca's area
2303:Frontal lobe
2243:cite journal
2216:
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2199:
2166:
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2156:
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2117:
2107:
2093:cite journal
2068:
2064:
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2044:cite journal
2019:
2015:
2009:
1995:cite journal
1986:
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1961:cite journal
1944:
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1937:Penfield, W.
1931:
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1757:
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447:Re-emergence
441:
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254:frontal lobe
250:motor cortex
245:
243:
188:
105:
96:
86:
79:
72:
65:
53:
41:Please help
36:verification
33:
3246:Limbic lobe
2295:human brain
2118:J. Neurosci
1887:Brodmann, K
1839:2434/147582
902:J. Neurosci
690:J. Neurosci
331:spinal cord
266:spinal cord
178:Identifiers
3503:span gyri.
3169:Interlobar
2866:3, 1 and 2
2606:Precentral
2467:Prefrontal
2433:Precentral
2319:Prefrontal
523:: 277–462.
490:References
411:Betz cells
407:Betz cells
295:Betz cells
289:, or SMA.
196:NeuroNames
69:newspapers
3483:Operculum
3031:41 and 42
2829:Precuneus
1782:206772150
357:PMDc (F2)
277:Structure
163:The brain
3516:Category
3445:Amygdala
3263:anterior
2235:17964243
2191:11642213
1889:(1909).
1848:20216547
1665:26064832
1602:11368801
1594:16277998
1542:15312656
1514:(2004).
1483:12062029
1111:10212467
1103:16839304
1068:30681663
1060:14984423
967:(2008).
932:16597724
883:15748854
842:38010957
785:37009687
654:41391502
592:42912356
560:Vogt, O.
556:Vogt, C.
513:Vogt, O.
509:Vogt, C.
476:Graziano
432:Penfield
384:PMVr(F5)
375:PMVc(F4)
366:PMDr(F7)
339:thalamus
335:striatum
3476:General
2293:of the
2183:3416964
2148:7119878
2139:6564318
2085:7351548
2036:7351547
1774:1301372
1722:7679990
1714:9325390
1657:3416965
1422:4415358
1414:9989407
1394:Bibcode
1354:7973661
1334:Bibcode
1326:Science
1287:8836215
1238:4028658
1230:7248055
1195:7760145
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572:Bibcode
409:". The
400:History
256:of the
160:Part of
155:Details
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2981:Cuneus
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333:, the
325:, and
317:, the
213:224852
169:Artery
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3440:Uncus
3428:Other
3304:gyrus
2187:S2CID
1941:Brain
1911:Brain
1852:S2CID
1814:(PDF)
1778:S2CID
1718:S2CID
1661:S2CID
1598:S2CID
1418:S2CID
1322:(PDF)
1234:S2CID
1107:S2CID
1064:S2CID
987:Brain
838:S2CID
781:S2CID
650:S2CID
588:S2CID
258:brain
224:[
184:Latin
90:JSTOR
76:books
2850:Both
2639:Both
2249:link
2231:PMID
2179:PMID
2144:PMID
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2032:PMID
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1056:PMID
1021:link
1003:PMID
946:link
928:PMID
879:PMID
830:PMID
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773:PMID
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718:PMID
668:link
642:PMID
606:link
558:and
535:link
511:and
244:The
201:2331
62:news
2221:doi
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2134:PMC
2126:doi
2073:doi
2024:doi
1949:doi
1919:doi
1834:hdl
1826:doi
1762:doi
1702:doi
1645:doi
1582:doi
1528:doi
1469:doi
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1390:397
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