Supplementary motor area | |
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Identifiers | |
NeuroNames | 3176 |
FMA | 224858 |
Anatomical terms of neuroanatomy |
The supplementary motor area (SMA) is a part of the motor cortex of primates that contributes to the control of movement. It is located on the midline surface of the hemisphere just in front of (anterior to) the primary motor cortex leg representation. In monkeys, the SMA contains a rough map of the body. In humans, the body map is not apparent. Neurons in the SMA project directly to the spinal cord and may play a role in the direct control of movement. Possible functions attributed to the SMA include the postural stabilization of the body, the coordination of both sides of the body such as during bimanual action, the control of movements that are internally generated rather than triggered by sensory events, and the control of sequences of movements. All of these proposed functions remain hypotheses. The precise role or roles of the SMA is not yet known.
For the discovery of the SMA and its relationship to other motor cortical areas, see the main article on the motor cortex.
At least six areas are now recognized within the larger region once defined as the SMA. These subdivisions have been studied most extensively in the monkey brain. The most anterior portion is now commonly termed pre-SMA. [1] [2] [3] It has sparse or no connections to the spinal cord or the primary motor cortex and has extensive connectivity with prefrontal areas. [1] [4] [5] [6] [7]
The supplementary eye field (SEF) is a relatively anterior portion of the SMA that, when stimulated, evokes head and eye movements and perhaps movements of the limbs and torso. [8] [9] [10] [11]
Dum and Strick [5] hypothesized on the basis of cytoarchitecture and connections to the spinal cord that the portion of SMA in the cingulate sulcus, on the medial part of the hemisphere, can be split into three separate areas, the cingulate motor areas. The functions of the cingulate motor areas have not yet been systematically studied, though may be involved in emotionally driven behaviours like the limbic laugh.
SMA proper in monkeys has now been confined to a region on the crown of the hemisphere and extending partly onto the medial wall, just anterior to the primary motor leg representation. SMA proper projects directly to the spinal cord and therefore is one of the primary output areas of the cortical motor system. [5] [12] [13] [14] [15] [16]
Recently, Zhang et al. [17] investigated the functional subdivisions of the medial SFC on the basis of whole-brain connectivity characterized from a large resting-state fMRI data set. Other than replicating the boundaries between SMA and preSMA, the current results support a functional difference between the posterior and anterior pre-SMA. In contrast to the posterior pre-SMA, the anterior pre-SMA is connected with most of the prefrontal but not somatomotor areas. Overall, the SMA is strongly connected to the thalamus and epithalamus, the posterior pre-SMA to putamen, pallidum, and STN and anterior pre-SMA to the caudate nucleus, with the caudate showing significant hemispheric asymmetry.
Penfield and Welch [18] in 1951 first described SMA in the monkey brain and the human brain as a representation of the body on the medial wall of the hemisphere. Woolsey and colleagues [19] in 1952 confirmed SMA in the monkey brain, describing it as a rough somatotopic map with the legs in a posterior location and the face in an anterior location. The representations of different body parts were found to overlap extensively. Stimulation of many sites evoked bilateral movements and sometimes movements of all four limbs. This overlapping somatotopic map in SMA was confirmed by many others. [2] [13] [20] [21] [22]
Four main hypotheses have been proposed for the function of SMA: the control of postural stability during stance or walking, [18] coordinating temporal sequences of actions, [23] [24] [25] [26] [27] [28] [29] [30] bimanual coordination, [31] [32] and the initiation of internally generated as opposed to stimulus driven movement. [3] [29] [30] [33] The data, however, tend not to support an exclusive role of SMA in any one of these functions. Indeed, SMA is demonstrably active during non-sequential, unimanual, and stimulus-cued movements. [34]
In humans, the SMA has been shown to generate the early component of the Bereitschaftspotential (BP) or readiness potential BP1 or BPearly. [35] The role of the SMA was further substantiated by Cunnington et al. 2003, [36] showing that SMA proper and pre-SMA are active prior to volitional movement or action, as well as the cingulate motor area (CMA) and anterior mid-cingulate cortex (aMCC). Recently it has been shown by integrating simultaneously acquired EEG and fMRI that SMA and aMCC have strong reciprocal connections that act to sustain each other’s activity, and that this interaction is mediated during movement preparation according to the Bereitschaftspotential amplitude. [37]
SMA in the monkey brain may emphasize locomotion, especially complex locomotion such as climbing or leaping. [38] [39] [40] This suggestion was based on studies in which stimulation on a behaviorally relevant time scale evoked complex, full body movements that resembled climbing or leaping. This hypothesis is consistent with previous hypotheses, including the involvement of SMA in postural stabilization, in internally generated movements, in bimanual coordination, and in the planning of movement sequences, because all of these functions are heavily recruited in complex locomotion. The locomotion hypothesis is an example of interpreting the motor cortex in terms of the underlying behavioral repertoire from which abstract control functions emerge, an approach emphasized by Graziano and colleagues. [38]
In neuroanatomy, the precuneus is the portion of the superior parietal lobule on the medial surface of each brain hemisphere. It is located in front of the cuneus. The precuneus is bounded in front by the marginal branch of the cingulate sulcus, at the rear by the parieto-occipital sulcus, and underneath by the subparietal sulcus. It is involved with episodic memory, visuospatial processing, reflections upon self, and aspects of consciousness.
Brodmann area 10 is the anterior-most portion of the prefrontal cortex in the human brain. BA10 was originally defined broadly in terms of its cytoarchitectonic traits as they were observed in the brains of cadavers, but because modern functional imaging cannot precisely identify these boundaries, the terms anterior prefrontal cortex, rostral prefrontal cortex and frontopolar prefrontal cortex are used to refer to the area in the most anterior part of the frontal cortex that approximately covers BA10—simply to emphasize the fact that BA10 does not include all parts of the prefrontal cortex.
In neuroanatomy, the superior colliculus is a structure lying on the roof of the mammalian midbrain. In non-mammalian vertebrates, the homologous structure is known as the optic tectum or optic lobe. The adjective form tectal is commonly used for both structures.
The motor cortex is the region of the cerebral cortex involved in the planning, control, and execution of voluntary movements. The motor cortex is an area of the frontal lobe located in the posterior precentral gyrus immediately anterior to the central sulcus.
The insular cortex is a portion of the cerebral cortex folded deep within the lateral sulcus within each hemisphere of the mammalian brain.
In neurology, the Bereitschaftspotential or BP, also called the pre-motor potential or readiness potential (RP), is a measure of activity in the motor cortex and supplementary motor area of the brain leading up to voluntary muscle movement. The BP is a manifestation of cortical contribution to the pre-motor planning of volitional movement. It was first recorded and reported in 1964 by Hans Helmut Kornhuber and Lüder Deecke at the University of Freiburg in Germany. In 1965 the full publication appeared after many control experiments.
In animals, including humans, the startle response is a largely unconscious defensive response to sudden or threatening stimuli, such as sudden noise or sharp movement, and is associated with negative affect. Usually the onset of the startle response is a startle reflex reaction. The startle reflex is a brainstem reflectory reaction (reflex) that serves to protect vulnerable parts, such as the back of the neck and the eyes (eyeblink) and facilitates escape from sudden stimuli. It is found across many different species, throughout all stages of life. A variety of responses may occur depending on the affected individual's emotional state, body posture, preparation for execution of a motor task, or other activities. The startle response is implicated in the formation of specific phobias.
In physiology, motor coordination is the orchestrated movement of multiple body parts as required to accomplish intended actions, like walking. This coordination is achieved by adjusting kinematic and kinetic parameters associated with each body part involved in the intended movement. The modifications of these parameters typically relies on sensory feedback from one or more sensory modalities, such as proprioception and vision.
Supplementary eye field (SEF) is the name for the anatomical area of the dorsal medial frontal lobe of the primate cerebral cortex that is indirectly involved in the control of saccadic eye movements. Evidence for a supplementary eye field was first shown by Schlag, and Schlag-Rey. Current research strives to explore the SEF's contribution to visual search and its role in visual salience. The SEF constitutes together with the frontal eye fields (FEF), the intraparietal sulcus (IPS), and the superior colliculus (SC) one of the most important brain areas involved in the generation and control of eye movements, particularly in the direction contralateral to their location. Its precise function is not yet fully known. Neural recordings in the SEF show signals related to both vision and saccades somewhat like the frontal eye fields and superior colliculus, but currently most investigators think that the SEF has a special role in high level aspects of saccade control, like complex spatial transformations, learned transformations, and executive cognitive functions.
The premotor cortex is an area of the motor cortex lying within the frontal lobe of the brain just anterior to the primary motor cortex. 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 spinal cord and therefore may play a role in the direct control of behavior, with a relative emphasis on the trunk muscles 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 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.
The basal ganglia form a major brain system in all species of vertebrates, but in primates there are special features that justify a separate consideration. As in other vertebrates, the primate basal ganglia can be divided into striatal, pallidal, nigral, and subthalamic components. In primates, however, there are two pallidal subdivisions called the external globus pallidus (GPe) and internal globus pallidus (GPi). Also in primates, the dorsal striatum is divided by a large tract called the internal capsule into two masses named the caudate nucleus and the putamen—in most other species no such division exists, and only the striatum as a whole is recognized. Beyond this, there is a complex circuitry of connections between the striatum and cortex that is specific to primates. This complexity reflects the difference in functioning of different cortical areas in the primate brain.
Beevor's Axiom is the idea that the brain does not know muscles, only movements. In other words, the brain registers the movements that muscles combine to make, not the individual muscles that are making the movements. Hence, this is why one can sign their name with their foot. Beevor's Axiom was coined by Dr. Charles Edward Beevor, an English neurologist.
Premovement neuronal activity in neurophysiological literature refers to neuronal modulations that alter the rate at which neurons fire before a subject produces movement. Through experimentation with multiple animals, predominantly monkeys, it has been shown that several regions of the brain are particularly active and involved in initiation and preparation of movement. Two specific membrane potentials, the bereitschaftspotential, or the BP, and contingent negative variation, or the CNV, play a pivotal role in premovement neuronal activity. Both have been shown to be directly involved in planning and initiating movement. Multiple factors are involved with premovement neuronal activity including motor preparation, inhibition of motor response, programming of the target of movement, closed-looped and open-looped tasks, instructed delay periods, short-lead and long-lead changes, and mirror motor neurons.
Michael Matthias Merzenich is an American neuroscientist and professor emeritus at the University of California, San Francisco. He took the sensory cortex maps developed by his predecessors and refined them using dense micro-electrode mapping techniques. Using this, he definitively showed there to be multiple somatotopic maps of the body in the postcentral sulcus, and multiple tonotopic maps of the acoustic inputs in the superior temporal plane.
Kurt A. Thoroughman is an Associate Professor in the Department of Biomedical Engineering at Washington University in St. Louis. He is known for his work in the study of motor control, motor learning, and computational neuroscience.
The isothalamus is a division used by some researchers in describing the thalamus.
The primary motor cortex is a brain region that in humans is located in the dorsal portion of the frontal lobe. It is the primary region of the motor system and works in association with other motor areas including premotor cortex, the supplementary motor area, posterior parietal cortex, and several subcortical brain regions, to plan and execute voluntary movements. Primary motor cortex is defined anatomically as the region of cortex that contains large neurons known as Betz cells, which, along with other cortical neurons, send long axons down the spinal cord to synapse onto the interneuron circuitry of the spinal cord and also directly onto the alpha motor neurons in the spinal cord which connect to the muscles.
Michael Steven Anthony Graziano is an American scientist and novelist who is currently a professor of Psychology and Neuroscience at Princeton University. 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. His previous work focused on how the cerebral cortex monitors the space around the body and controls movement within that space. 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. His publications on this topic have had a widespread impact among neuroscientists but have also generated controversy. His novels rely partly on his background in psychology and are known for surrealism or magic realism. Graziano also composes music including symphonies and string quartets.
A motor program is an abstract metaphor of the central organization of movement and control of the many degrees of freedom involved in performing an action.p. 182 Signals transmitted through efferent and afferent pathways allow the central nervous system to anticipate, plan or guide movement. Evidence for the concept of motor programs include the following:p. 182
Nonprimary motor cortex is a functionally defined portion of the frontal lobe. It includes two subdivisions, the premotor cortex and the supplementary motor cortex. Largely coincident with the cytoarchitecturally defined area 6 of Brodmann (human), it is located primarily in the rostral portion of the precentral gyrus and caudal portions of the superior frontal gyrus and the middle frontal gyrus, It aids in cerebral control of movement. Anatomically speaking, several nonmprimary areas exist, and make direct connections with the spinal cord.
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