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other type) is perhaps the most provocative feature of cerebellar anatomy, and has motivated much of the theorizing. In fact, the function of climbing fibers is the most controversial topic concerning the cerebellum. There are two schools of thought, one following Marr and Albus in holding that climbing fiber input serves primarily as a teaching signal, the other holding that its function is to shape cerebellar output directly. Both views have been defended in great length in numerous publications. In the words of one review, "In trying to synthesize the various hypotheses on the function of the climbing fibers, one has the sense of looking at a drawing by Escher. Each point of view seems to account for a certain collection of findings, but when one attempts to put the different views together, a coherent picture of what the climbing fibers are doing does not appear. For the majority of researchers, the climbing fibers signal errors in motor performance, either in the usual manner of discharge frequency modulation or as a single announcement of an 'unexpected event'. For other investigators, the message lies in the degree of ensemble synchrony and rhythmicity among a population of climbing fibers."
1421:—that is, signals move unidirectionally through the system from input to output, with very little recurrent internal transmission. The small amount of recurrence that does exist consists of mutual inhibition; there are no mutually excitatory circuits. This feedforward mode of operation means that the cerebellum, in contrast to the cerebral cortex, cannot generate self-sustaining patterns of neural activity. Signals enter the circuit, are processed by each stage in sequential order, and then leave. As Eccles, Ito, and Szentágothai wrote, "This elimination in the design of all possibility of reverberatory chains of neuronal excitation is undoubtedly a great advantage in the performance of the cerebellum as a computer, because what the rest of the nervous system requires from the cerebellum is presumably not some output expressing the operation of complex reverberatory circuits in the cerebellum but rather a quick and clear response to the input of any particular set of information."
1747:
2016:
expanded. The size of the human cerebellum, compared to the rest of the brain, has been increasing in size while the cerebrum decreased in size With both the development and implementation of motor tasks, visual-spatial skills and learning taking place in the cerebellum, the growth of the cerebellum is thought to have some form of correlation to greater human cognitive abilities. The lateral hemispheres of the cerebellum are now 2.7 times greater in both humans and apes than they are in monkeys. These changes in the cerebellum size cannot be explained by greater muscle mass. They show that either the development of the cerebellum is tightly linked to that of the rest of the brain or that neural activities taking place in the cerebellum were important during
1454:: The cerebellar system is functionally divided into more or less independent modules, which probably number in the hundreds to thousands. All modules have a similar internal structure, but different inputs and outputs. A module (a multizonal microcompartment in the terminology of Apps and Garwicz) consists of a small cluster of neurons in the inferior olivary nucleus, a set of long narrow strips of Purkinje cells in the cerebellar cortex (microzones), and a small cluster of neurons in one of the deep cerebellar nuclei. Different modules share input from mossy fibers and parallel fibers, but in other respects they appear to function independently—the output of one module does not appear to significantly influence the activity of other modules.
1157:
a single
Purkinje cell. In striking contrast to the 100,000-plus inputs from parallel fibers, each Purkinje cell receives input from exactly one climbing fiber; but this single fiber "climbs" the dendrites of the Purkinje cell, winding around them and making a total of up to 300 synapses as it goes. The net input is so strong that a single action potential from a climbing fiber is capable of producing an extended complex spike in the Purkinje cell: a burst of several spikes in a row, with diminishing amplitude, followed by a pause during which activity is suppressed. The climbing fiber synapses cover the cell body and proximal dendrites; this zone is devoid of parallel fiber inputs.
47:
847:
617:. The input to the pons is from the cerebral cortex and is relayed from the pontine nuclei via transverse pontine fibers to the cerebellum. The middle peduncle is the largest of the three and its afferent fibers are grouped into three separate fascicles taking their inputs to different parts of the cerebellum. The inferior cerebellar peduncle receives input from afferent fibers from the vestibular nuclei, spinal cord and the tegmentum. Output from the inferior peduncle is via efferent fibers to the vestibular nuclei and the reticular formation. The whole of the cerebellum receives modulatory input from the inferior olivary nucleus via the inferior cerebellar peduncle.
2098:
1345:
generate motor activity but lose precision, producing erratic, uncoordinated, or incorrectly timed movements. A standard test of cerebellar function is to reach with the tip of the finger for a target at arm's length: A healthy person will move the fingertip in a rapid straight trajectory, whereas a person with cerebellar damage will reach slowly and erratically, with many mid-course corrections. Deficits in non-motor functions are more difficult to detect. Thus, the general conclusion reached decades ago is that the basic function of the cerebellum is to calibrate the detailed form of a movement, not to initiate movements or to decide which movements to execute.
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therefore all, it is presumed, performing the same computation. If the input and output connections of a module are with motor areas (as many are), then the module will be involved in motor behavior; but, if the connections are with areas involved in non-motor cognition, the module will show other types of behavioral correlates. Thus the cerebellum has been implicated in the regulation of many differing functional traits such as affection, emotion including emotional body language perception and behavior. The cerebellum, Doya proposes, is best understood as predictive action selection based on "internal models" of the environment or a device for
1978:, the cerebellum is barely distinguishable from the brain-stem. Although the spinocerebellum is present in these groups, the primary structures are small, paired-nuclei corresponding to the vestibulocerebellum. The cerebellum is a bit larger in reptiles, considerably larger in birds, and larger still in mammals. The large paired and convoluted lobes found in humans are typical of mammals, but the cerebellum is, in general, a single median lobe in other groups, and is either smooth or only slightly grooved. In mammals, the neocerebellum is the major part of the cerebellum by mass, but, in other vertebrates, it is typically the spinocerebellum.
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conditioned response or CR. Experiments showed that lesions localized either to a specific part of the interposed nucleus (one of the deep cerebellar nuclei) or to a few specific points in the cerebellar cortex would abolish learning of a conditionally timed blink response. If cerebellar outputs are pharmacologically inactivated while leaving the inputs and intracellular circuits intact, learning takes place even while the animal fails to show any response, whereas, if intracerebellar circuits are disrupted, no learning takes place—these facts taken together make a strong case that the learning, indeed, occurs inside the cerebellum.
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2060:. They are glutamatergic, but also resemble Purkinje cells in some respects—they have spiny, flattened superficial dendritic trees that receive parallel fiber input, but they also have basal dendrites that receive input from auditory nerve fibers, which travel across the DCN in a direction at right angles to the parallel fibers. The DCN is most highly developed in rodents and other small animals, and is considerably reduced in primates. Its function is not well understood; the most popular speculations relate it to spatial hearing in one way or another.
1649:(involuntary movement caused by alternating contractions of opposing muscle groups). Damage to the midline portion may disrupt whole-body movements, whereas damage localized more laterally is more likely to disrupt fine movements of the hands or limbs. Damage to the upper part of the cerebellum tends to cause gait impairments and other problems with leg coordination; damage to the lower part is more likely to cause uncoordinated or poorly aimed movements of the arms and hands, as well as difficulties in speed. This complex of motor symptoms is called
2229:
1079:; the splitting of the vertical branch into two horizontal branches gives rise to a distinctive "T" shape. A human parallel fiber runs for an average of 3 mm in each direction from the split, for a total length of about 6 mm (about 1/10 of the total width of the cortical layer). As they run along, the parallel fibers pass through the dendritic trees of Purkinje cells, contacting one of every 3–5 that they pass, making a total of 80–100 synaptic connections with Purkinje cell dendritic spines. Granule cells use
1087:
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1460:: The synapses between parallel fibers and Purkinje cells, and the synapses between mossy fibers and deep nuclear cells, are both susceptible to modification of their strength. In a single cerebellar module, input from as many as a billion parallel fibers converges onto a group of less than 50 deep nuclear cells, and the influence of each parallel fiber on those nuclear cells is adjustable. This arrangement gives tremendous flexibility for fine-tuning the relationship between the cerebellar inputs and outputs.
1551:. The original theory put forth by Braitenberg and Roger Atwood in 1958 proposed that slow propagation of signals along parallel fibers imposes predictable delays that allow the cerebellum to detect time relationships within a certain window. Experimental data did not support the original form of the theory, but Braitenberg continued to argue for modified versions. The hypothesis that the cerebellum functions essentially as a timing system has also been advocated by
1060:, in contrast to Purkinje cells, are among the smallest neurons in the brain. They are also the most numerous neurons in the brain: In humans, estimates of their total number average around 50 billion, which means that about 3/4 of the brain's neurons are cerebellar granule cells. Their cell bodies are packed into a thick layer at the bottom of the cerebellar cortex. A granule cell emits only four to five dendrites, each of which ends in an enlargement called a
1103:, who suggested that they could encode combinations of mossy fiber inputs. The idea is that with each granule cell receiving input from only 4–5 mossy fibers, a granule cell would not respond if only a single one of its inputs were active, but would respond if more than one were active. This combinatorial coding scheme would potentially allow the cerebellum to make much finer distinctions between input patterns than the mossy fibers alone would permit.
1212:). The fastigial and interposed nuclei belong to the spinocerebellum. The dentate nucleus, which in mammals is much larger than the others, is formed as a thin, convoluted layer of gray matter, and communicates exclusively with the lateral parts of the cerebellar cortex. The flocculus of the flocculonodular lobe is the only part of the cerebellar cortex that does not project to the deep nuclei—its output goes to the vestibular nuclei instead.
1589:
1037:
1932:
490:
626:
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59:
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653:(below the posterior fissure). These lobes divide the cerebellum from rostral to caudal (in humans, top to bottom). In terms of function, however, there is a more important distinction along the medial-to-lateral dimension. Leaving out the flocculonodular lobe, which has distinct connections and functions, the cerebellum can be parsed functionally into a medial sector called the
993:, each of which receives synaptic input from a parallel fiber. Purkinje cells receive more synaptic inputs than any other type of cell in the brain—estimates of the number of spines on a single human Purkinje cell run as high as 200,000. The large, spherical cell bodies of Purkinje cells are packed into a narrow layer (one cell thick) of the cerebellar cortex, called the
922:
746:. There is disagreement about the best way to describe the functions of the lateral cerebellum: It is thought to be involved in planning movement that is about to occur, in evaluating sensory information for action, and in a number of purely cognitive functions, such as determining the verb which best fits with a certain noun (as in "sit" for "chair").
2084:, but none have climbing fibers comparable to those of the cerebellum—instead they receive direct input from peripheral sensory organs. None has a demonstrated function, but the most influential speculation is that they serve to transform sensory inputs in some sophisticated way, perhaps to compensate for changes in body posture. In fact,
1503:
spike activity—which is known to reliably indicate activity of the cell's climbing fiber input—during periods when performance was poor. Several studies of motor learning in cats observed complex spike activity when there was a mismatch between an intended movement and the movement that was actually executed. Studies of the
1571:, a neurally inspired abstract learning device. The most basic difference between the Marr and Albus theories is that Marr assumed that climbing fiber activity would cause parallel fiber synapses to be strengthened, whereas Albus proposed that they would be weakened. Albus also formulated his version as a
2158:
the lesion is very extensive. By the beginning of the 20th century, it was widely accepted that the primary function of the cerebellum relates to motor control; the first half of the 20th century produced several detailed descriptions of the clinical symptoms associated with cerebellar disease in humans.
1603:
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Congenital malformation, hereditary disorders, and acquired conditions can affect cerebellar structure and, consequently, cerebellar function. Unless the causative condition is reversible, the only possible treatment is to help people live with their problems. Visualization of the fetal cerebellum by
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The strongest clues to the function of the cerebellum have come from examining the consequences of damage to it. Animals and humans with cerebellar dysfunction show, above all, problems with motor control, on the same side of the body as the damaged part of the cerebellum. They continue to be able to
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with those of the anterior and posterior inferior cerebellar arteries. The AICA supplies the front part of the undersurface of the cerebellum. The PICA arrives at the undersurface, where it divides into a medial branch and a lateral branch. The medial branch continues backward to the cerebellar notch
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the cerebellum for certain types of protein. The best-known of these markers are called "zebrins", because staining for them gives rise to a complex pattern reminiscent of the stripes on a zebra. The stripes generated by zebrins and other compartmentalization markers are oriented perpendicular to the
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Climbing fibers fire at low rates, but a single climbing fiber action potential induces a burst of several action potentials in a target
Purkinje cell (a complex spike). The contrast between parallel fiber and climbing fiber inputs to Purkinje cells (over 100,000 of one type versus exactly one of the
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and receives input from the spinal cord, brainstem and cerebral cortex, its output goes entirely to the cerebellum. A climbing fiber gives off collaterals to the deep cerebellar nuclei before entering the cerebellar cortex, where it splits into about 10 terminal branches, each of which gives input to
1914:
There is a general consensus that the cerebellum is involved in pain processing. The cerebellum receives pain input from both descending cortico-cerebellar pathways and ascending spino-cerebellar pathways, through the pontine nuclei and inferior olives. Some of this information is transferred to the
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There is considerable evidence that the cerebellum plays an essential role in some types of motor learning. The tasks where the cerebellum most clearly comes into play are those in which it is necessary to make fine adjustments to the way an action is performed. There has, however, been much dispute
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infiltrate the structure and make inhibitory synapses onto the granule cell dendrites. The entire assemblage is surrounded by a sheath of glial cells. Each mossy fiber sends collateral branches to several cerebellar folia, generating a total of 20–30 rosettes; thus a single mossy fiber makes contact
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Granule cells receive all of their input from mossy fibers, but outnumber them by 200 to 1 (in humans). Thus, the information in the granule cell population activity state is the same as the information in the mossy fibers, but recoded in a much more expansive way. Because granule cells are so small
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of about 10 ms; a complex spike is a stereotyped sequence of action potentials with very short inter-spike intervals and declining amplitudes. Physiological studies have shown that complex spikes (which occur at baseline rates around 1 Hz and never at rates much higher than 10 Hz) are
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in the first half of the 19th century carried out detailed experimental work, which revealed that animals with cerebellar damage can still move, but with a loss of coordination (strange movements, awkward gait, and muscular weakness), and that recovery after the lesion can be nearly complete unless
2004:
The hallmark of the mammalian cerebellum is an expansion of the lateral lobes, whose main interactions are with the neocortex. As monkeys evolved into great apes, the expansion of the lateral lobes continued, in tandem with the expansion of the frontal lobes of the neocortex. In ancestral hominids,
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signaling stimulates rapid proliferation of cerebellar granule neuron progenitors (CGNPs) in the external granule layer (EGL). Cerebellar development occurs during late embryogenesis and the early postnatal period, with CGNP proliferation in the EGL peaking during early development (postnatal day 7
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studies have shown cerebellar activation in relation to language, attention, and mental imagery; correlation studies have shown interactions between the cerebellum and non-motor areas of the cerebral cortex; and a variety of non-motor symptoms have been recognized in people with damage that appears
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in 1837. They are distinguished by the shape of their dendritic tree: the dendrites branch very profusely, but are severely flattened in a plane perpendicular to the cerebellar folds. Thus, the dendrites of a
Purkinje cell form a dense planar net, through which parallel fibers pass at right angles.
781:
Mossy fibers project directly to the deep nuclei, but also give rise to the following pathway: mossy fibers → granule cells → parallel fibers → Purkinje cells → deep nuclei. Climbing fibers project to
Purkinje cells and also send collaterals directly to the deep nuclei. The mossy fiber and climbing
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are embedded in the white matter. Each part of the cortex consists of the same small set of neuronal elements, laid out in a highly stereotyped geometry. At an intermediate level, the cerebellum and its auxiliary structures can be separated into several hundred or thousand independently functioning
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The large base of knowledge about the anatomical structure and behavioral functions of the cerebellum have made it a fertile ground for theorizing—there are perhaps more theories of the function of the cerebellum than of any other part of the brain. The most basic distinction among them is between
1310:
In 2005, Richard Apps and Martin
Garwicz summarized evidence that microzones themselves form part of a larger entity they call a multizonal microcomplex. Such a microcomplex includes several spatially separated cortical microzones, all of which project to the same group of deep cerebellar neurons,
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and others have argued, partly on the basis of these structures and partly on the basis of cerebellar studies, that the cerebellum itself is fundamentally a sensory structure, and that it contributes to motor control by moving the body in a way that controls the resulting sensory signals. Despite
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system that senses pressure waves in water. One of the brain areas that receives primary input from the lateral line organ, the medial octavolateral nucleus, has a cerebellum-like structure, with granule cells and parallel fibers. In electrosensitive fish, the input from the electrosensory system
1918:
These direct pain inputs, as well as indirect inputs, are thought to induce long-term pain avoidance behavior that results in chronic posture changes and consequently, in functional and anatomical remodeling of vestibular and proprioceptive nuclei. As a result, chronic neuropathic pain can induce
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Theories in the "learning" category almost all derive from publications by Marr and Albus. Marr's 1969 paper proposed that the cerebellum is a device for learning to associate elemental movements encoded by climbing fibers with mossy fiber inputs that encode the sensory context. Albus proposed in
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paradigm, in which a neutral conditioned stimulus (CS) such as a tone or a light is repeatedly paired with an unconditioned stimulus (US), such as an air puff, that elicits a blink response. After such repeated presentations of the CS and US, the CS will eventually elicit a blink before the US, a
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Kenji Doya has argued that the cerebellum's function is best understood not in terms of the behaviors it affects, but the neural computations it performs; the cerebellum consists of a large number of more or less independent modules, all with the same geometrically regular internal structure, and
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The first indications of compartmental structure came from studies of the receptive fields of cells in various parts of the cerebellar cortex. Each body part maps to specific points in the cerebellum, but there are numerous repetitions of the basic map, forming an arrangement that has been called
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From the viewpoint of gross anatomy, the cerebellar cortex appears to be a homogeneous sheet of tissue, and, from the viewpoint of microanatomy, all parts of this sheet appear to have the same internal structure. There are, however, a number of respects in which the structure of the cerebellum is
1502:
The idea that climbing fiber activity functions as an error signal has been examined in many experimental studies, with some supporting it but others casting doubt. In a pioneering study by
Gilbert and Thach from 1977, Purkinje cells from monkeys learning a reaching task showed increased complex
1490:
synapses. Marr assumed that climbing fiber input would cause synchronously activated parallel fiber inputs to be strengthened. Most subsequent cerebellar-learning models, however, have followed Albus in assuming that climbing fiber activity would be an error signal, and would cause synchronously
2015:
period, the cerebellum continued to expand, but the frontal lobes expanded more rapidly. The most recent period of human evolution, however, may actually have been associated with an increase in the relative size of the cerebellum, as the neocortex reduced its size somewhat while the cerebellum
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Oscarsson in the late 1970s proposed that these cortical zones can be partitioned into smaller units called microzones. A microzone is defined as a group of
Purkinje cells all having the same somatotopic receptive field. Microzones were found to contain on the order of 1000 Purkinje cells each,
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When a
Purkinje cell axon enters one of the deep nuclei, it branches to make contact with both large and small nuclear cells, but the total number of cells contacted is only about 35 (in cats). Conversely, a single deep nuclear cell receives input from approximately 860 Purkinje cells (again in
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are much longer in the longitudinal direction than in the mediolateral direction, causing them to be confined largely to a single microzone. The consequence of all this structure is that cellular interactions within a microzone are much stronger than interactions between different microzones.
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etc. In the human cerebellum, the total number of mossy fibers has been estimated at 200 million. These fibers form excitatory synapses with the granule cells and the cells of the deep cerebellar nuclei. Within the granular layer, a mossy fiber generates a series of enlargements called
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cerebellar folds—that is, they are narrow in the mediolateral direction, but much more extended in the longitudinal direction. Different markers generate different sets of stripes, the widths and lengths vary as a function of location, but they all have the same general shape.
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reliably associated with climbing fiber activation, while simple spikes are produced by a combination of baseline activity and parallel fiber input. Complex spikes are often followed by a pause of several hundred milliseconds during which simple spike activity is suppressed.
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at a high rate even in the absence of the synaptic input. In awake, behaving animals, mean rates averaging around 40 Hz are typical. The spike trains show a mixture of what are called simple and complex spikes. A simple spike is a single action potential followed by a
1298:, which synchronize their activity, causing Purkinje cells within a microzone to show correlated complex spike activity on a millisecond time scale. Also, the Purkinje cells belonging to a microzone all send their axons to the same small cluster of output cells within the
1176:
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to match the excitatory projection of climbing fibers to the nuclei. There is evidence that each small cluster of nuclear cells projects to the same cluster of olivary cells that send climbing fibers to it; there is strong and matching topography in both directions.
1786:(also called cerebellar granule neurons, CGNs), they migrate to the internal granule layer (IGL), forming the mature cerebellum (by post-natal day 20 in the mouse). Mutations that abnormally activate Sonic hedgehog signaling predispose to cancer of the cerebellum (
3913:
Hernáez-Goñi P, Tirapu-Ustárroz J, Iglesias-Fernández L, Luna-Lario P (November 2010). "Participación del cerebelo en la regulación del afecto, la emoción y la conducta" [The role of the cerebellum in the regulation of affection, emotion and behavior].
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about whether learning takes place within the cerebellum itself, or whether it merely serves to provide signals that promote learning in other brain structures. Most theories that assign learning to the circuitry of the cerebellum are derived from the ideas of
1385:. Three decades of brain research have led to the proposal that the cerebellum generates optimized mental models and interacts closely with the cerebral cortex, where updated internal models are experienced as creative intuition ("a ha") in working memory.
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is equally important. The branches of a climbing fiber (usually numbering about 10) usually activate
Purkinje cells belonging to the same microzone. Moreover, olivary neurons that send climbing fibers to the same microzone tend to be coupled by
1721:
The human cerebellum changes with age. These changes may differ from those of other parts of the brain. The cerebellum is the youngest brain region (and body part) in centenarians according to an epigenetic biomarker of tissue age known as
2182:), which was used in the works of Aristotle, the first known writer to describe the structure. No other name is used in the English-language literature, but historically a variety of Greek or Latin-derived names have been used, including
1443:. Because of the way that they are lined up longitudinally, the 1000 or so Purkinje cells belonging to a microzone may receive input from as many as 100 million parallel fibers, and focus their own output down to a group of less than 50
1410:. Although a full understanding of cerebellar function has remained elusive, at least four principles have been identified as important: (1) feedforward processing, (2) divergence and convergence, (3) modularity, and (4) plasticity.
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trees of
Purkinje cells, along with the huge array of parallel fibers penetrating the Purkinje cell dendritic trees at right angles. This outermost layer of the cerebellar cortex also contains two types of inhibitory interneuron:
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The comparative simplicity and regularity of the cerebellar anatomy led to an early hope that it might imply a similar simplicity of computational function, as expressed in one of the first books on cerebellar electrophysiology,
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of the cerebellum are clusters of gray matter lying within the white matter at the core of the cerebellum. They are, with the minor exception of the nearby vestibular nuclei, the sole sources of output from the cerebellum. These
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within the cerebellum to account for its role in learning, versus theories that account for aspects of ongoing behavior on the basis of cerebellar signal processing. Several theories of both types have been formulated as
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of Purkinje cells, and the huge array of parallel fibers, from the granular layer, that penetrate the Purkinje cell dendritic trees at right angles. The molecular layer also contains two types of inhibitory interneuron:
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cells. Thus, the cerebellar network receives a modest number of inputs, processes them very extensively through its rigorously structured internal network, and sends out the results via a very limited number of output
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arranged in a long, narrow strip, oriented perpendicular to the cortical folds. Thus, as the adjoining diagram illustrates, Purkinje cell dendrites are flattened in the same direction as the microzones extend, while
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and so densely packed, it is difficult to record their spike activity in behaving animals, so there is little data to use as a basis for theorizing. The most popular concept of their function was proposed in 1969 by
568:. High‑resolution MRI finds the adult human cerebellar cortex has an area of 730 square cm, packed within a volume of dimensions 6 cm × 5 cm × 10 cm. Underneath the gray matter of the cortex lies
2079:
All of these cerebellum-like structures appear to be primarily sensory-related rather than motor-related. All of them have granule cells that give rise to parallel fibers that connect to Purkinje-like neurons with
695:
The medial zone of the anterior and posterior lobes constitutes the spinocerebellum, also known as paleocerebellum. This sector of the cerebellum functions mainly to fine-tune body and limb movements. It receives
778:(which are the axons of granule cells). There are two main pathways through the cerebellar circuit, originating from mossy fibers and climbing fibers, both eventually terminating in the deep cerebellar nuclei.
1008:
Purkinje cells form the heart of the cerebellar circuit, and their large size and distinctive activity patterns have made it relatively easy to study their response patterns in behaving animals using
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722:
The lateral zone, which in humans is by far the largest part, constitutes the cerebrocerebellum, also known as neocerebellum. It receives input exclusively from the cerebral cortex (especially the
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between the two hemispheres of the cerebellum; while the lateral branch supplies the under surface of the cerebellum, as far as its lateral border, where it anastomoses with the AICA and the SCA.
3318:
Whitney ER, Kemper TL, Rosene DL, Bauman ML, Blatt GJ (February 2008). "Calbindin-D28k is a more reliable marker of human Purkinje cells than standard Nissl stains: a stereological experiment".
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in the cerebral cortex. The fibers arise from the deep cerebellar nuclei. The middle cerebellar peduncle is connected to the pons and receives all of its input from the pons mainly from the
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in 1664. More anatomical work was done during the 18th century, but it was not until early in the 19th century that the first insights into the function of the cerebellum were obtained.
1919:
macroscopic anatomical remodeling of the hindbrain, including the cerebellum. The magnitude of this remodeling and the induction of neuron progenitor markers suggest the contribution of
1855:, in which tumors elsewhere in the body elicit an autoimmune response that causes neuronal loss in the cerebellum. Cerebellar atrophy can result from an acute deficiency of vitamin B1 (
332:
such as regulating fear and pleasure responses, but its movement-related functions are the most solidly established. The human cerebellum does not initiate movement, but contributes to
1989:
is extraordinarily large and complex. In at least one important respect, it differs in internal structure from the mammalian cerebellum: The fish cerebellum does not contain discrete
6885:
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typically causes problems in skilled voluntary and planned movements which can cause errors in the force, direction, speed and amplitude of movements. Other manifestations include
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1997:(a family of weakly electrosensitive freshwater fish), the cerebellum is considerably larger than the rest of the brain. The largest part of it is a special structure called the
719:
systems. It sends fibers to deep cerebellar nuclei that, in turn, project to both the cerebral cortex and the brain stem, thus providing modulation of descending motor systems.
1563:, which provided an advanced mathematical formulation of the idea that the fundamental computation performed by the cerebellum is to transform sensory into motor coordinates.
1507:(which stabilizes the visual image on the retina when the head turns) found that climbing fiber activity indicated "retinal slip", although not in a very straightforward way.
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than the total from the rest of the brain, but takes up only 10% of the total brain volume. The number of neurons in the cerebellum is related to the number of neurons in the
399:. This complex neural organization gives rise to a massive signal-processing capability, but almost all of the output from the cerebellar cortex passes through a set of small
5776:
Roberts PD, Portfors CV (June 2008). "Design principles of sensory processing in cerebellum-like structures. Early stage processing of electrosensory and auditory objects".
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Baumann O, Borra RJ, Bower JM, Cullen KE, Habas C, Ivry RB, Leggio M, Mattingley JB, Molinari M, Moulton EA, Paulin MG, Pavlova MA, Schmahmann JD, Sokolov AA (April 2015).
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includes assessment of gait (a broad-based gait being indicative of ataxia), finger-pointing tests and assessment of posture. If cerebellar dysfunction is indicated, a
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Prior to the 1990s the function of the cerebellum was almost universally believed to be purely motor-related, but newer findings have brought that view into question.
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Damage to the cerebellum often causes motor-related symptoms, the details of which depend on the part of the cerebellum involved and how it is damaged. Damage to the
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1192:
receive collateral projections from mossy fibers and climbing fibers as well as inhibitory input from the Purkinje cells of the cerebellar cortex. The four nuclei (
4991:
Campbell J, Gilbert WM, Nicolaides KH, Campbell S (August 1987). "Ultrasound screening for spina bifida: cranial and cerebellar signs in a high-risk population".
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as their neurotransmitter. These cells project to a variety of targets outside the cerebellum. Intermixed with them are a lesser number of small cells, which use
430:, based on the observation that each cerebellar Purkinje cell receives two dramatically different types of input: one comprises thousands of weak inputs from the
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1075:
axons of granule cells rise vertically to the upper (molecular) layer of the cortex, where they split in two, with each branch traveling horizontally to form a
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may show up as a loss of equilibrium and in particular an altered, irregular walking gait, with a wide stance caused by difficulty in balancing. Damage to the
253:
629:
Schematic representation of the major anatomical subdivisions of the cerebellum. Superior view of an "unrolled" cerebellum, placing the vermis in one plane.
605:, named by their position relative to the vermis. The superior cerebellar peduncle is mainly an output to the cerebral cortex, carrying efferent fibers via
4661:
1208:) each communicate with different parts of the brain and cerebellar cortex. (The globose and the emboliform nuclei are also referred to as combined in the
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1705:
headache may also produce temporary dysfunction of the cerebellum, of variable severity. Infection can result in cerebellar damage in such conditions as
3186:
Felizola SJ, Nakamura Y, Ono Y, Kitamura K, Kikuchi K, Onodera Y, Ise K, Takase K, Sugawara A, Hattangady N, Rainey WE, Satoh F, Sasano H (April 2014).
442:
is that the climbing fiber serves as a "teaching signal", which induces a long-lasting change in the strength of parallel fiber inputs. Observations of
1993:. Instead, the primary targets of Purkinje cells are a distinct type of cell distributed across the cerebellar cortex, a type not seen in mammals. In
553:. There are about 3.6 times as many neurons in the cerebellum as in the neocortex, a ratio that is conserved across many different mammalian species.
1738:. Some studies have reported reductions in numbers of cells or volume of tissue, but the amount of data relating to this question is not very large.
984:
are among the most distinctive neurons in the brain, and one of the earliest types to be recognized—they were first described by the Czech anatomist
5314:
4359:
Braitenberg V, Heck D, Sultan F (June 1997). "The detection and generation of sequences as a key to cerebellar function: experiments and theory".
2134:) or brain proper. Galen's extensive description is the earliest that survives. He speculated that the cerebellum was the source of motor nerves.
5056:
Polkinghorn WR, Tarbell NJ (May 2007). "Medulloblastoma: tumorigenesis, current clinical paradigm, and efforts to improve risk stratification".
3777:"Neuropsychological consequences of cerebellar tumour resection in children: cerebellar cognitive affective syndrome in a paediatric population"
1331:(PICA). The SCA supplies the upper region of the cerebellum. It divides at the upper surface and branches into the pia mater where the branches
6158:
801:
The cerebellar cortex is divided into three layers. At the bottom lies the thick granular layer, densely packed with granule cells, along with
541:). A set of large folds is, by convention, used to divide the overall structure into 10 smaller "lobules". Because of its large number of tiny
1417:: The cerebellum differs from most other parts of the brain (especially the cerebral cortex) in that the signal processing is almost entirely
977:
Purkinje cells in the human cerebellum (in orange, from top to bottom 40X, 100X and 200X magnification) stained according to published methods
5819:
Bower JM (1997). "Chapter 27 is the cerebellum sensory for motor's sake, or motor for sensory's sake: The view from the whiskers of a rat?".
229:
6183:
5262:
5184:
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The majority of neurons in the deep nuclei have large cell bodies and spherical dendritic trees with a radius of about 400 μm, and use
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Most vertebrate species have a cerebellum and one or more cerebellum-like structures, brain areas that resemble the cerebellum in terms of
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5240:
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cause progressive neurodegeneration linked to cerebellar loss. Congenital brain malformations outside the cerebellum can, in turn, cause
1001:, where they make on the order of 1,000 contacts each with several types of nuclear cells, all within a small domain. Purkinje cells use
367:
Anatomically, the human cerebellum has the appearance of a separate structure attached to the bottom of the brain, tucked underneath the
4139:
2967:
Petersen SE, Fox PT, Posner MI, Mintun M, Raichle ME (1989). "Positron emission tomographic studies of the processing of single words".
2020:
evolution. Due to the cerebellum's role in cognitive functions, the increase in its size may have played a role in cognitive expansion.
556:
The unusual surface appearance of the cerebellum conceals the fact that most of its volume is made up of a very tightly folded layer of
6895:
2048:
that rise to the superficial layer and travel across it horizontally. The superficial layer contains a set of GABAergic neurons called
4975:
2052:
that resemble Purkinje cells anatomically and chemically—they receive parallel fiber input, but do not have any inputs that resemble
5158:
2044:. The DCN is a layered structure, with the bottom layer containing granule cells similar to those of the cerebellum, giving rise to
6232:
5985:
Ito M (December 2002). "Historical review of the significance of the cerebellum and the role of Purkinje cells in motor learning".
1131:. Each glomerulus has a mossy fiber rosette at its center, and up to 20 granule cell dendritic claws contacting it. Terminals from
4638:
4459:
Pellionisz A, Llinás R (1982). "Space-time representation in the brain. The cerebellum as a predictive space-time metric tensor".
1357:
or Schmahmann's syndrome has been described in adults and children. Estimates based on functional mapping of the cerebellum using
1053:
Granule cells (GR, bottom), parallel fibers (horizontal lines, top), and Purkinje cells (P, middle) with flattened dendritic trees
525:; all of its connections with other parts of the brain travel through the pons. Anatomists classify the cerebellum as part of the
1750:
Ultrasound image of the fetal head at 19 weeks of pregnancy in a modified axial section, showing the normal fetal cerebellum and
1706:
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1354:
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Horvath S, Mah V, Lu AT, Woo JS, Choi OW, Jasinska AJ, Riancho JA, Tung S, Coles NS, Braun J, Vinters HV, Coles LS (May 2015).
2346:
Wolf U, Rapoport MJ, Schweizer TA (2009). "Evaluating the affective component of the cerebellar cognitive affective syndrome".
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Other conditions that are closely linked to cerebellar degeneration include the idiopathic progressive neurological disorders
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plus a group of coupled olivary neurons that project to all of the included microzones as well as to the deep nuclear area.
6437:
5911:"Strength and timing of motor responses mediated by rebound firing in the cerebellar nuclei after Purkinje cell activation"
4950:
3143:
Schweighofer N, Doya K, Kuroda S (March 2004). "Cerebellar aminergic neuromodulation: towards a functional understanding".
1328:
147:
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1324:
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Sagittal cross-section of human cerebellum, showing the dentate nucleus, as well as the pons and inferior olivary nucleus
446:
in parallel fiber inputs have provided some support for theories of this type, but their validity remains controversial.
143:
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MacLeod CE, Zilles K, Schleicher A, Rilling JK, Gibson KR (April 2003). "Expansion of the neocerebellum in Hominoidea".
375:
surface is covered with finely spaced parallel grooves, in striking contrast to the broad irregular convolutions of the
1361:
suggest that more than half of the cerebellar cortex is interconnected with association zones of the cerebral cortex.
1029:. Calbindin staining of rat brain after unilateral chronic sciatic nerve injury suggests that Purkinje neurons may be
6816:
5214:
5124:
4555:
4042:
1645:(inability to perform rapid alternating movements such as walking), impaired check reflex or rebound phenomenon, and
4092:
Boyden ES, Katoh A, Raymond JL (2004). "Cerebellum-dependent learning: the role of multiple plasticity mechanisms".
2250:
6481:
260:
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Doya K (October 1999). "What are the computations of the cerebellum, the basal ganglia and the cerebral cortex?".
2089:
Bower's viewpoint, there is also strong evidence that the cerebellum directly influences motor output in mammals.
6191:. Gruol, D.L., Koibuchi, N., Manto, M., Molinari, M., Schmahmann, J.D., Shen, Y. (Eds.). Springer, New York, 2016
4873:
Andersen BB, Gundersen HJ, Pakkenberg B (November 2003). "Aging of the human cerebellum: a stereological study".
3491:
Apps R, Garwicz M (April 2005). "Anatomical and physiological foundations of cerebellar information processing".
1864:
1399:
5471:
Bell CC, Han V, Sawtell NB (2008). "Cerebellum-like structures and their implications for cerebellar function".
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he called a CMAC (Cerebellar Model Articulation Controller), which has been tested in a number of applications.
30:
This article is about the smaller region in the lower part of the brain. For the large region of the brain, see
6395:
5344:
1887:
1834:
1018:
17:
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Hatten ME, Heintz N (1995). "Mechanisms of neural patterning and specification in the developing cerebellum".
2604:
Handbook of Neurosurgery, Neurology, and Spinal Medicine for Nurses and Advanced Practice Health Professionals
1710:
6853:
6505:
3640:
Doya K (December 2000). "Complementary roles of basal ganglia and cerebellum in learning and motor control".
2001:, which has an unusually regular architecture and receives much of its input from the electrosensory system.
1920:
1795:
1009:
642:
248:
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Fine EJ, Ionita CC, Lohr L (December 2002). "The history of the development of the cerebellar examination".
1966:
There is considerable variation in the size and shape of the cerebellum in different vertebrate species. In
948:
The top, outermost layer of the cerebellar cortex is the molecular layer. This layer contains the flattened
576:
nerve fibers running to and from the cortex. Embedded within the white matter—which is sometimes called the
379:. These parallel grooves conceal the fact that the cerebellar cortex is actually a continuous thin layer of
6849:
6806:
6802:
6476:
6225:
5587:"The neuronal organization of a unique cerebellar specialization: the valvula cerebelli of a mormyrid fish"
2632:
1698:
817:. In the middle lies the Purkinje layer, a narrow zone that contains the cell bodies of Purkinje cells and
646:
602:
594:
4918:"Age and sex differences in the cerebellum and the ventral pons: a prospective MR study of healthy adults"
46:
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Manni E, Petrosini L (March 2004). "A century of cerebellar somatotopy: a debated representation".
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The lower trace shows an attempt by a patient with cerebellar disease to reproduce the upper trace.
1428:
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1145:
1127:. The contacts between mossy fibers and granule cell dendrites take place within structures called
1111:
874:
864:
767:
348:, and integrates these inputs to fine-tune motor activity. Cerebellar damage produces disorders in
187:
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Turner BM, Paradiso S, Marvel CL, Pierson R, Boles Ponto LL, Hichwa RD, Robinson RG (March 2007).
3012:"Cerebellar contributions to cognitive functions: a progress report after two decades of research"
2114:
Even the earliest anatomists were able to recognize the cerebellum by its distinctive appearance.
410:
In addition to its direct role in motor control, the cerebellum is necessary for several types of
6569:
6218:
5266:
5188:
4316:
Braitenberg V, Atwood RP (February 1958). "Morphological observations on the cerebellar cortex".
2153:
in 1809 established the key finding that damage to the cerebellum results in motor disturbances.
2037:
1959:. This has been taken as evidence that the cerebellum performs functions important to all animal
1875:
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1432:
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997:. After emitting collaterals that affect nearby parts of the cortex, their axons travel into the
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498:
418:. Several theoretical models have been developed to explain sensorimotor calibration in terms of
396:
357:
3291:
Simpson JI, Wylie DR, De Zeeuw CI (1996). "On climbing fiber signals and their consequence(s)".
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6400:
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3870:. Michael Adamaszek, Mario Manto, Dennis J. L. G. Schutter. Cham, Switzerland: Springer. 2022.
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1990:
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motor system inducing a conscious motor avoidance of pain, graded according to pain intensity.
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1064:. These enlargements are sites of excitatory input from mossy fibers and inhibitory input from
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6166:– Manto, M., Gruol, D.L., Schmahmann, J., Koibuchi, N., Rossi, F. (Eds.) – Springer – New York
5236:
4980:(18th ed.). Whitehouse Station, New Jersey: Merck Research Libraries. pp. 1886–1887.
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1951:, including fish, reptiles, birds, and mammals. There is also an analogous brain structure in
312:
it may be as large as it or even larger. In humans, the cerebellum plays an important role in
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5862:"Precise control of movement kinematics by optogenetic inhibition of Purkinje cell activity"
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scan can be used to obtain a detailed picture of any structural alterations that may exist.
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6137:
5994:
5742:
5647:
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3237:"The excitatory synaptic action of climbing fibres on the Purkinje cells of the cerebellum"
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2154:
1763:
676:. It is the oldest part in evolutionary terms (archicerebellum) and participates mainly in
650:
522:
443:
3104:"The cerebellum and cognitive function: 25 years of insight from anatomy and neuroimaging"
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730:(forming cortico-ponto-cerebellar pathways), and sends output mainly to the ventrolateral
8:
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3818:"The organization of the human cerebellum estimated by intrinsic functional connectivity"
2606:, Abingdon, Oxon ; New York, NY : Routledge, 2018.: Routledge, pp. 53–62,
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goes to the dorsal octavolateral nucleus, which also has a cerebellum-like structure. In
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684:, although it also receives visual and other sensory input. Damage to this region causes
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590:
419:
368:
6141:
5998:
5746:
5651:
5034:
4419:
3685:"Schmahmann's syndrome - identification of the third cornerstone of clinical ataxiology"
3577:
Oscarsson O (1979). "Functional units of the cerebellum-sagittal zones and microzones".
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2819:
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activated parallel fiber inputs to be weakened. Some of these later models, such as the
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Schematic illustration of the structure of zones and microzones in the cerebellar cortex
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4538:
Schmitz TJ (2007). "Examination of Coordination". In O'Sullivan SB, Schmitz TJ (eds.).
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582:(tree of life) because of its branched, tree-like appearance in cross-section—are four
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Discoveries in the human brain. Neuroscience prehistory, brain structure, and function
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Illustration from 1912 of the altered walking gait of a woman with cerebellar disease
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The Handbook of Cognitive Neuropsychology: What Deficits Reveal about the Human Mind
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Cerebellar atrophy has been observed in many other neurological disorders including
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1269:"fractured somatotopy". A clearer indication of compartmentalization is obtained by
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as their neurotransmitter, and therefore exert excitatory effects on their targets.
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as their neurotransmitter, and therefore exert inhibitory effects on their targets.
589:
Connecting the cerebellum to different parts of the nervous system are three paired
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5514:
Woodhams PL (July 1977). "The ultrastructure of a cerebellar analogue in octopus".
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discussed the cerebellum briefly, and the anatomy was described more thoroughly by
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The list of medical problems that can produce cerebellar damage is long, including
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349:
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Gilbert PF, Thach WT (June 1977). "Purkinje cell activity during motor learning".
2040:(DCN), one of the two primary sensory nuclei that receive input directly from the
1535:"learning theories" and "performance theories"—that is, theories that make use of
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fiber inputs each carry fiber-specific information; the cerebellum also receives
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4917:
4662:"Veterans of Iraq, Afghanistan Show Brain Changes Related to Explosion Exposure"
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2903:"Receiving Areas of the Tactile, Auditory, and Visual Systems in the Cerebellum"
1894:. Cerebellar atrophy can also occur as a result of exposure to toxins including
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Proceedings of the National Academy of Sciences of the United States of America
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4635:"NINDS Ataxias and Cerebellar or Spinocerebellar Degeneration Information Page"
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Purves D, Augustine GJ, Fitzpatrick D, Hall WC, LaMantia AS, White LE (2007).
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Purves D, Augustine GJ, Fitzpatrick D, Hall WC, LaMantia AS, White LE (2011).
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Lyu, Wenjiao; Wu, Ye; Huynh, Khoi Minh; Ahmad, Sahar; Yap, Pew-Thian (2024).
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Perhaps the earliest "performance" theory was the "delay line" hypothesis of
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or "hindbrain". Like the cerebral cortex, the cerebellum is divided into two
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313:
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5660:
3188:"PCP4: a regulator of aldosterone synthesis in human adrenocortical tissues"
2919:
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2178:. The Latin name is a direct translation of the Ancient Greek παρεγκεφαλίς (
1495:
model of Fujita made attempts to understand cerebellar function in terms of
1319:
The cerebellum is provided with blood from three paired major arteries: the
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compartmentalized. There are large compartments that are generally known as
1114:
enter the granular layer from their points of origin, many arising from the
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6038:
A Latin dictionary founded on Andrews' edition of Freund's Latin dictionary
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3221:
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Ghez C, Fahn S (1985). "The cerebellum". In Kandel ER, Schwartz JH (eds.).
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2007:
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in the adult brain, initiating the organization of new cerebellar lobules.
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529:, which also includes the pons; the metencephalon is the upper part of the
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of the granule cells; the other is an extremely strong input from a single
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3366:"Peripheral nerve injury induces adult brain neurogenesis and remodelling"
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A specific, recognizable feature of Purkinje neurons is the expression of
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A Man's Incomplete Brain Reveals Cerebellum's Role In Thought And Emotion
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2012:
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It is not only receptive fields that define the microzone structure: The
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Buckner RL, Krienen FM, Castellanos A, Diaz JC, Yeo BT (November 2011).
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2244:). The updated content was reintegrated into the Knowledge page under a
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1726:: it is about 15 years younger than expected in a centenarian. Further,
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5636:"Reciprocal evolution of the cerebellum and neocortex in fossil humans"
5069:
4277:"Neural substrates of eyeblink conditioning: acquisition and retention"
4202:
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Strick PL, Dum RP, Fiez JA (2009). "Cerebellum and nonmotor function".
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518:
298:
199:
5698:
Schoenemann PT (December 1, 2009). "Evolution of Brain and Language".
5602:
4886:
661:. A narrow strip of protruding tissue along the midline is called the
266:
6299:
6272:
6173:
5417:"Consensus paper: the role of the cerebellum in perceptual processes"
5368:"The cerebellum and pain: passive integrator or active participator?"
4293:
4276:
2115:
2017:
1986:
1967:
1868:
1638:
1630:
1510:
One of the most extensively studied cerebellar learning tasks is the
1403:
1026:
787:
550:
384:
321:
317:
294:
127:
3547:
3504:
1602:
821:. At the top lies the molecular layer, which contains the flattened
422:
within the cerebellum. These models derive from those formulated by
6321:
6294:
6257:
6093:. Göttingen: Verlag der Deuerlich- und Dieterichschen Buchhandlung.
5909:
Witter L, Canto CB, Hoogland TM, de Gruijl JR, De Zeeuw CI (2013).
2753:"Coordinated scaling of cortical and cerebellar numbers of neurons"
1860:
1702:
1682:
1572:
949:
822:
783:
754:
Two types of neuron play dominant roles in the cerebellar circuit:
731:
672:
The smallest region, the flocculonodular lobe, is often called the
514:
513:
are in front of the cerebellum. It is separated from the overlying
325:
306:
302:
211:
31:
27:
Structure at the rear of the vertebrate brain, beneath the cerebrum
5213:. National Institutes of Health. 29 September 2011. Archived from
1049:
1036:
6210:
5343:. National Institutes of Health. 14 February 2007. Archived from
5265:. National Institutes of Health. 14 February 2011. Archived from
5187:. National Institutes of Health. 14 February 2014. Archived from
5161:. National Institutes of Health. 23 December 2013. Archived from
4990:
4746:. National Institutes of Health. 28 February 2014. Archived from
4596:
Gilman S (March 1998). "Imaging the brain. Second of two parts".
4402:
Ivry RB, Spencer RM, Zelaznik HN, Diedrichsen J (December 2002).
1975:
1971:
1960:
1956:
329:
5366:
Moulton EA, Schmahmann JD, Becerra L, Borsook D (October 2010).
2646:
Standring S, Borley NR, et al., eds. (2008). "Chapter 20".
1482:
provide a teaching signal that induces synaptic modification in
625:
489:
6309:
5365:
4916:
Raz N, Gunning-Dixon F, Head D, Williamson A, Acker JD (2001).
3774:
2648:
Gray's anatomy : the anatomical basis of clinical practice
1670:
1651:
1072:
573:
546:
388:
134:
6901:
Knowledge articles published in peer-reviewed literature (W2J)
5823:. Progress in Brain Research. Vol. 114. pp. 463–96.
5732:
5317:. National Institutes of Health. 12 March 2009. Archived from
5291:. National Institutes of Health. 16 April 2014. Archived from
4915:
4771:"The cerebellum ages slowly according to the epigenetic clock"
4401:
2804:"A multimodal submillimeter MRI atlas of the human cerebellum"
2572:
2510:(5th ed.). Sunderland, Mass.: Sinauer. pp. 417–423.
2505:
680:
and spatial orientation; its primary connections are with the
6267:
3815:
2650:(40th ed.). London: Churchill Livingstone. p. 297.
2123:
2036:. The only cerebellum-like structure found in mammals is the
1231:. Thus, the nucleo-olivary projection provides an inhibitory
345:
175:
58:
5908:
5263:"NINDS Dyssynergia Cerebellaris Myoclonica Information Page"
4817:
4189:
Fujita M (1982). "Adaptive filter model of the cerebellum".
1248:
391:
with a highly regular arrangement, the most important being
6282:
6106:
Synonymia anatomica. Synonymik der anatomischen Nomenclatur
5860:
Heiney SA, Kim J, Augustine GJ, Medina JF (February 2014).
4872:
3948:
3185:
2577:(4th ed.). New York: W. H. Freeman. pp. 197–200.
1741:
1002:
763:
506:
309:
154:
109:
103:
94:
88:
5564:. Philadelphia: Holt-Saunders International. p. 531.
2795:
1261:; these can be divided into smaller compartments known as
1175:
921:
6170:
Stained brain slice images which include the "cerebellum"
3317:
1090:
Diagram of the layers of the cerebellar cortex showing a
336:, precision, and accurate timing: it receives input from
6886:
Knowledge articles published in peer-reviewed literature
5859:
2137:
Further significant developments did not come until the
2076:
has a layer—the marginal layer—that is cerebellum-like.
1427:: In the human cerebellum, information from 200 million
6891:
Knowledge articles published in WikiJournal of Medicine
5414:
3775:
Levisohn L, Cronin-Golomb A, Schmahmann JD (May 2000).
2966:
1805:) of the cerebellar vermis is a characteristic of both
464:, the cerebellum consists of a tightly folded layer of
5585:
Shi Z, Zhang Y, Meek J, Qiao J, Han VZ (August 2008).
4035:
The new revolution in psychology and the neurosciences
3142:
1943:
The circuits in the cerebellum are similar across all
1782:
in the mouse). As CGNPs terminally differentiate into
293:; Latin for "little brain") is a major feature of the
5289:"NINDS Olivopontocerebellar Atrophy Information Page"
4358:
4057:
4032:
2348:
Journal of Neuropsychiatry and Clinical Neurosciences
2345:
1353:
to be confined to the cerebellum. In particular, the
1223:
as a neurotransmitter and project exclusively to the
85:
82:
5960:
Clarke E, O'Malley CD (1996). "Ch. 11: Cerebellum".
5693:
5691:
5689:
5341:"NINDS Wernicke-Korsakoff Syndrome Information Page"
3290:
2536:
Albus JS (1971). "A theory of cerebellar function".
2317:. Berlin, Heidelberg: Springer. pp. 1240–1243.
1926:
1641:(problems judging distances or ranges of movement),
1567:
1971 that a cerebellar Purkinje cell functions as a
1518:
481:
modules called "microzones" or "microcompartments".
106:
100:
91:
6164:
Handbook of the Cerebellum and Cerebellar Disorders
5239:. National Institutes of Health. 10 December 2014.
4137:
2946:(2nd ed.). Lippincott Williams & Wilkins.
1527:Model of a cerebellar perceptron, as formulated by
1040:
A mouse Purkinje cell injected with fluorescent dye
1012:recording techniques. Purkinje cells normally emit
112:
97:
5959:
5055:
4969:
4967:
4965:
4963:
4091:
4059:
3234:
2645:
1693:such as gunshot wounds or explosives, and chronic
1555:. Another influential "performance" theory is the
938:
798:inputs that presumably perform global modulation.
6189:Essentials of Cerebellum and Cerebellar Disorders
5686:
5315:"NINDS Paraneoplastic Syndromes Information Page"
4690:
4511:Intelligent Control Systems and Signal Processing
4315:
4274:
3733:
2260:; Finn Årup Nielsen; et al. (30 June 2016).
6877:
5964:(2nd ed.). Norman Publishing. p. 629.
5584:
4824:"Aging and gene expression in the primate brain"
4744:"NINDS Cerebellar Degeneration Information Page"
4637:. National Institutes of Health. 16 April 2014.
4629:
4627:
4575:. London, UK: Academic Press. pp. 337–351.
4458:
2750:
2721:
2380:
883: • CFC: Climbing fiber collateral
637:can be distinguished within the cerebellum: the
564:. Each ridge or gyrus in this layer is called a
414:, most notably learning to adjust to changes in
51:Position of the human cerebellum (sagittal view)
5775:
4960:
4768:
3486:
2801:
1148:on the contralateral side of the brainstem via
774:(which enter the cerebellum from outside), and
6121:
6050:
5470:
5211:"NINDS Cerebellar Hypoplasia Information Page"
4738:
4736:
3533:
3484:
3482:
3480:
3478:
3476:
3474:
3472:
3470:
3468:
3466:
3235:Eccles JC, Llinás R, Sasaki K (January 1966).
3179:
2459:
2063:Most species of fish and amphibians possess a
1762:at 18 to 20 weeks of pregnancy can be used to
931:, showing principal cell types and connections
537:; it also contains a narrow midline zone (the
387:. Within this thin layer are several types of
6445:
6431:
6226:
6088:Kritisch-etymologisches medicinisches Lexikon
6066:
6064:
6062:
5098:
4624:
4140:"Models of the cerebellum and motor learning"
3736:"The cerebellar cognitive affective syndrome"
3359:
3357:
3058:
2900:
2722:Llinas RR, Walton KD, Lang EJ (2004). "Ch. 7
2023:
1801:Congenital malformation or underdevelopment (
5559:
5101:"Cerebellum development and medulloblastoma"
5020:
4973:
4231:
3363:
1813:. In very rare cases, the entire cerebellum
861: • (-): Inhibitory connection
859: • (+): Excitatory connection
493:View of the cerebellum from above and behind
5697:
4733:
4570:
4504:"CMAC: Reconsidering an old neural network"
4133:
4131:
3682:
3490:
3463:
3009:
2598:Dididze, Marine; Levi, Allan (2017-12-15),
2597:
2313:Hodos W (2009). "Evolution of Cerebellum".
2174:(brain); it can be translated literally as
1730:patterns in the human cerebellum show less
1144:Purkinje cells also receive input from the
6707:Bergmann glia cell = Golgi epithelial cell
6438:
6424:
6233:
6219:
6200:The Cerebellum – Journal (Springer Nature)
6059:
6035:
5987:Annals of the New York Academy of Sciences
4408:Annals of the New York Academy of Sciences
3370:Journal of Cellular and Molecular Medicine
3354:
3286:
3284:
3282:
3280:
2421:
2254:). The version of record as reviewed is:
1152:. Although the inferior olive lies in the
57:
45:
6149:
5936:
5926:
5885:
5821:The Cerebellum: From Structure to Control
5669:
5659:
5610:
5440:
5391:
5159:"NINDS Joubert Syndrome Information Page"
5134:
4977:The Merck Manual of Diagnosis and Therapy
4936:
4849:
4839:
4794:
4716:
4427:
4292:
4158:
4138:Houk JC, Buckingham JT, Barto AG (1996).
4087:
4085:
4058:Eccles JC, Ito M, Szentágothai J (1967).
3974:
3951:"The cerebellum and emotional experience"
3841:
3792:
3751:
3710:
3700:
3576:
3446:
3389:
3260:
3211:
3119:
3027:
2918:
2888:Principles of Neural Science, 2nd edition
2843:
2778:
2768:
2549:
2398:
2381:Schmahmann JD, Caplan D (February 2006).
2280:
1939:, with the cerebellum highlighted in blue
1709:and Miller Fisher syndrome, a variant of
1136:with an estimated 400–600 granule cells.
734:(in turn connected to motor areas of the
6103:
5513:
4128:
4033:Manto M, Marvel C, Vandervert L (2022).
3734:Schmahmann JD, Sherman JC (April 1998).
2941:
2901:Snider RS, Stowell A (1 November 1944).
2885:
2501:
2499:
2096:
1930:
1745:
1742:Developmental and degenerative disorders
1578:
1522:
1247:
1174:
1085:
1048:
1035:
972:
624:
488:
5466:
5464:
5462:
5460:
5105:Current Topics in Developmental Biology
4537:
4501:
3635:
3633:
3413:
3411:
3409:
3277:
3101:
2890:. New York: Elsevier. pp. 502–522.
2881:
2879:
2877:
2875:
2873:
2871:
2869:
2867:
2865:
2863:
2717:
2715:
2713:
2711:
2709:
2707:
2705:
2703:
2701:
2699:
2697:
2695:
2693:
2691:
2689:
2687:
2531:
2529:
2527:
2383:"Cognition, emotion and the cerebellum"
1817:. The inherited neurological disorders
1355:cerebellar cognitive affective syndrome
965:synapses onto Purkinje cell dendrites.
838:synapses onto Purkinje cell dendrites.
657:and a larger lateral sector called the
633:Based on the surface appearance, three
14:
6878:
6070:
5633:
5485:10.1146/annurev.neuro.30.051606.094225
4595:
4546:. Philadelphia: F. A. Davis. pp.
4188:
4106:10.1146/annurev.neuro.27.070203.144238
4082:
3967:10.1016/j.neuropsychologia.2006.09.023
3145:Brain Research. Brain Research Reviews
3073:10.1146/annurev.neuro.31.060407.125606
2728:The Synaptic Organization of the Brain
2685:
2683:
2681:
2679:
2677:
2675:
2673:
2671:
2669:
2667:
2455:
2453:
2451:
2449:
1853:paraneoplastic cerebellar degeneration
1439:outputs then converge onto 15 million
961:. Both stellate and basket cells form
834:. Both stellate and basket cells form
6419:
6214:
6085:
5818:
5555:
5553:
5185:"NINDS Dandy-Walker Information Page"
5016:
5014:
4818:Fraser HB, Khaitovich P, Plotkin JB,
4691:Vincent M, Hadjikhani N (June 2007).
4672:from the original on January 20, 2016
4659:
2730:. New York: Oxford University Press.
2535:
2496:
2312:
2101:Base of the human brain, as drawn by
1637:(problems with speech articulation),
700:input from the dorsal columns of the
649:(below the primary fissure), and the
6075:. New York: D. Appleton and Company.
5457:
5243:from the original on 27 October 2011
4641:from the original on 9 February 2015
4077:The Cerebellum as a Neuronal Machine
4062:The Cerebellum as a Neuronal Machine
3997:
3639:
3630:
3611:
3417:
3406:
3364:Rusanescu G, Mao J (February 2017).
2860:
2639:
2524:
2442:– via Elsevier Science Direct.
2056:. The output neurons of the DCN are
1955:with well-developed brains, such as
1396:The Cerebellum as a Neuronal Machine
1329:posterior inferior cerebellar artery
301:. Although usually smaller than the
5984:
5035:10.1146/annurev.ne.18.030195.002125
4974:Albert RK, Porter RS, eds. (2006).
2664:
2446:
1325:anterior inferior cerebellar artery
927:Transverse section of a cerebellar
742:of the cerebral cortex) and to the
282:
24:
6240:
6007:10.1111/j.1749-6632.2002.tb07574.x
5550:
5117:10.1016/B978-0-12-380916-2.00008-5
5058:Nature Clinical Practice. Oncology
5011:
4925:American Journal of Neuroradiology
4429:10.1111/j.1749-6632.2002.tb07576.x
4275:Christian KM, Thompson RF (2003).
3192:Journal of Molecular Endocrinology
1777:In normal development, endogenous
1139:
943:
383:tightly folded in the style of an
25:
6922:
6896:Externally peer reviewed articles
6159:Cerebellum–Cell Centered Database
6115:
6073:An illustrated medical dictionary
5384:10.1016/j.brainresrev.2010.05.005
4404:"The cerebellum and event timing"
3616:. Psychology Press. p. 481.
3157:10.1016/j.brainresrev.2003.10.004
2969:Journal of Cognitive Neuroscience
1970:, it is little developed, and in
1927:Comparative anatomy and evolution
1658:To identify cerebellar problems,
1519:Theories and computational models
1431:inputs is expanded to 40 billion
1044:
968:
856:Abbreviations and representations
497:The cerebellum is located in the
5712:10.1111/j.1467-9922.2009.00539.x
5591:Journal of Comparative Neurology
5516:Journal of Comparative Neurology
5237:"Chiari Malformation Fact Sheet"
4956:from the original on 2008-12-17.
4875:Journal of Comparative Neurology
4709:10.1111/j.1526-4610.2006.00715.x
4367:(2): 229–45, discussion 245–77.
4318:Journal of Comparative Neurology
4037:. Switzerland: Springer Nature.
2227:
2130:), as opposed to the ἐγκέφαλος (
2072:(by far the largest group), the
1935:Cross-section of the brain of a
1601:
1587:
920:
852:Microcircuitry of the cerebellum
845:
484:
261:Anatomical terms of neuroanatomy
78:
6184:Woman living without cerebellum
6097:
6079:
6051:Marshall LH, Magoun HW (1998).
6044:
6029:
5978:
5962:The Human Brain and Spinal Cord
5953:
5902:
5853:
5812:
5769:
5726:
5627:
5578:
5507:
5408:
5359:
5333:
5307:
5281:
5255:
5229:
5203:
5177:
5151:
5092:
5049:
4984:
4909:
4866:
4811:
4762:
4684:
4653:
4598:New England Journal of Medicine
4589:
4564:
4531:
4495:
4452:
4395:
4352:
4309:
4268:
4225:
4182:
4070:
4051:
4026:
3991:
3942:
3906:
3858:
3809:
3768:
3727:
3676:
3642:Current Opinion in Neurobiology
3605:
3570:
3527:
3423:"A theory of cerebellar cortex"
3320:Journal of Neuroscience Methods
3311:
3228:
3136:
3095:
3052:
3003:
2960:
2935:
2894:
2744:
2109:
1831:herniation of cerebellar tissue
1544:and simulated using computers.
1314:
1243:
1118:, others from the spinal cord,
1106:
989:The dendrites are covered with
939:Layers of the cerebellar cortex
749:
620:
545:, the cerebellum contains more
6122:Llinas R, Negrello MN (2015).
5878:10.1523/JNEUROSCI.4547-13.2014
5099:Roussel MF, Hatten ME (2011).
3439:10.1113/jphysiol.1969.sp008820
3332:10.1016/j.jneumeth.2007.09.009
3253:10.1113/jphysiol.1966.sp007824
2591:
2566:
2424:"The cerebellum and cognition"
2415:
2374:
2339:
2323:10.1007/978-3-540-29678-2_3124
2306:
2232:This article was submitted to
2126:) called it the παρεγκεφαλίς (
1888:progressive myoclonus epilepsy
1794:and in genetically engineered
1164:
472:underneath and a fluid-filled
305:, in some animals such as the
13:
1:
6854:Ventral spinocerebellar tract
5829:10.1016/S0079-6123(08)63381-6
5755:10.1016/S0047-2484(03)00028-9
5560:Romer AS, Parsons TS (1977).
5473:Annual Review of Neuroscience
5023:Annual Review of Neuroscience
4822:, Eisen MB (September 2005).
4693:"The cerebellum and migraine"
4361:Behavioral and Brain Sciences
4094:Annual Review of Neuroscience
4012:10.1016/S0893-6080(99)00046-5
3654:10.1016/S0959-4388(00)00153-7
3061:Annual Review of Neuroscience
2422:Schmahmann, Jeremy D (2019).
2222:
1388:
715:, as well as from visual and
6807:Dorsal spinocerebellar tract
5915:Frontiers in Neural Circuits
4841:10.1371/journal.pbio.0030274
4660:Yuhas D (January 15, 2016).
4473:10.1016/0306-4522(82)90224-X
4246:10.1016/0006-8993(77)90997-0
3591:10.1016/0166-2236(79)90057-2
3536:Nature Reviews. Neuroscience
3493:Nature Reviews. Neuroscience
3121:10.1016/j.neuron.2013.10.044
2944:Concise Text of Neuroscience
2560:10.1016/0025-5564(71)90051-4
2440:10.1016/j.neulet.2018.07.005
2315:Encyclopedia of Neuroscience
2161:
1734:alteration than that in the
1699:olivopontocerebellar atrophy
1282:cross them at right angles.
603:inferior cerebellar peduncle
595:superior cerebellar peduncle
449:
407:interior of the cerebellum.
344:and from other parts of the
7:
6195:Cerebellum histology images
4610:10.1056/NEJM199803263381307
3102:Buckner RL (October 2013).
2751:Herculano-Houzel S (2010).
2205:
1865:Wernicke–Korsakoff syndrome
1846:Ramsay Hunt syndrome type I
1833:, as seen in some forms of
1464:
1339:
586:, composed of gray matter.
438:. The basic concept of the
63:Animation of the cerebellum
10:
6927:
6832:Trigeminocerebellar fibers
6207:– Journal (BioMed Central)
6040:. Oxford: Clarendon Press.
6036:Lewis CT, Short C (1879).
5735:Journal of Human Evolution
4571:Mariën P, Manto M (2016).
3822:Journal of Neurophysiology
3683:Manto M, Mariën P (2015).
3010:Timmann D, Daum I (2007).
2907:Journal of Neurophysiology
2828:10.1038/s41598-024-55412-y
2092:
2024:Cerebellum-like structures
1835:Arnold–Chiari malformation
1664:magnetic resonance imaging
1614:
1425:Divergence and convergence
1321:superior cerebellar artery
1168:
766:also play dominant roles:
599:middle cerebellar peduncle
453:
416:sensorimotor relationships
29:
6826:Vestibulocerebellar tract
6793:
6777:
6768:
6668:
6626:
6617:
6514:
6464:
6457:
6381:
6348:
6341:Peripheral nervous system
6339:
6248:
6151:10.4249/scholarpedia.4606
5790:10.1007/s00422-008-0217-1
5433:10.1007/s12311-014-0627-7
4993:Obstetrics and Gynecology
4573:The linguistic cerebellum
4373:10.1017/s0140525x9700143x
4169:10.1017/S0140525X00081474
3702:10.1186/s40673-015-0023-1
3305:10.1017/S0140525X00081486
3029:10.1080/14734220701496448
2981:10.1162/jocn.1989.1.2.153
2757:Frontiers in Neuroanatomy
2726:". In Shepherd GM (ed.).
2360:10.1176/jnp.2009.21.3.245
1677:, swelling of the brain (
1633:(decreased muscle tone),
456:Anatomy of the cerebellum
259:
247:
235:
223:
210:
198:
186:
174:
169:
153:
133:
123:
73:
68:
56:
44:
39:
5928:10.3389/fncir.2013.00133
5634:Weaver AH (March 2005).
3928:10.33588/rn.5110.2010394
3867:The emotional cerebellum
3794:10.1093/brain/123.5.1041
3689:Cerebellum & Ataxias
2770:10.3389/fnana.2010.00012
2631:: CS1 maint: location (
1784:cerebellar granule cells
1716:
1660:neurological examination
1302:. Finally, the axons of
1291:inferior olivary nucleus
1225:inferior olivary nucleus
1146:inferior olivary nucleus
1058:Cerebellar granule cells
6570:Vallecula of cerebellum
6055:. Totowa: Humana Press.
5866:Journal of Neuroscience
5661:10.1073/pnas.0500692102
4542:Physical Rehabilitation
3753:10.1093/brain/121.4.561
2920:10.1152/jn.1944.7.6.331
2600:"Neuroanatomy overview"
2269:WikiJournal of Medicine
2234:WikiJournal of Medicine
2212:Vestibulo–ocular reflex
2038:dorsal cochlear nucleus
1909:
1842:multiple system atrophy
1711:Guillain–Barré syndrome
1505:vestibulo–ocular reflex
986:Jan Evangelista Purkyně
900: • GgC:
868: • DCN:
686:disturbances of balance
517:by a layer of leathery
499:posterior cranial fossa
6843:Pontocerebellar fibers
6628:Deep cerebellar nuclei
6250:Central nervous system
6205:Cerebellum and Ataxias
5778:Biological Cybernetics
5372:Brain Research Reviews
4191:Biological Cybernetics
3898:: CS1 maint: others (
2106:
1991:deep cerebellar nuclei
1940:
1819:Machado–Joseph disease
1754:
1531:
1478:, who postulated that
1415:Feedforward processing
1375:reinforcement learning
1300:deep cerebellar nuclei
1253:
1180:
1171:Deep cerebellar nuclei
1095:
1094:in the granular layer.
1054:
1041:
999:deep cerebellar nuclei
978:
910: • BC:
905: • SC:
895: • PC:
890: • PF:
885: • GC:
878: • CF:
873: • IO:
870:Deep cerebellar nuclei
863: • MF:
669:is Latin for "worm".)
630:
584:deep cerebellar nuclei
535:cerebellar hemispheres
494:
478:deep cerebellar nuclei
6859:Dentatothalamic tract
6817:Cuneocerebellar tract
6812:Olivocerebellar tract
5528:10.1002/cne.901740209
4787:10.18632/aging.100742
4330:10.1002/cne.901090102
4281:Learning & Memory
3916:Revista de Neurología
3834:10.1152/jn.00339.2011
3427:Journal of Physiology
3241:Journal of Physiology
2462:Seminars in Neurology
2282:10.15347/WJM/2016.001
2217:Eyeblink conditioning
2100:
1934:
1823:ataxia telangiectasia
1807:Dandy–Walker syndrome
1749:
1579:Clinical significance
1557:Tensor network theory
1549:Valentino Braitenberg
1526:
1512:eyeblink conditioning
1383:unsupervised learning
1369:, in contrast to the
1251:
1178:
1089:
1052:
1039:
976:
706:spinocerebellar tract
628:
572:, made up largely of
492:
6489:Flocculonodular lobe
6104:Schreger CH (1805).
2942:Kingsley RE (2000).
2474:10.1055/s-2002-36759
2428:Neuroscience Letters
2400:10.1093/brain/awh729
2238:academic peer review
2155:Jean Pierre Flourens
1892:Niemann–Pick disease
1876:Huntington's disease
1623:flocculonodular lobe
819:Bergmann glial cells
815:unipolar brush cells
651:flocculonodular lobe
591:cerebellar peduncles
523:cerebellar tentorium
444:long-term depression
369:cerebral hemispheres
6822:Juxtarestiform body
6736:Unipolar brush cell
6697:Purkinje cell layer
6586:Alar central lobule
6142:2015SchpJ..10.4606L
5999:2002NYASA.978..273I
5747:2003JHumE..44..401M
5652:2005PNAS..102.3576W
5562:The Vertebrate Body
5269:on 16 February 2015
4750:on 18 February 2015
4666:Scientific American
4420:2002NYASA.978..302I
3204:10.1530/JME-13-0248
2820:2024NatSR..14.5622L
2170:is a diminutive of
2082:modifiable synapses
1850:autoimmune disorder
1827:Friedreich's ataxia
1774:rate of up to 99%.
1768:neural tube defects
1697:conditions such as
1542:mathematical models
1537:synaptic plasticity
1367:supervised learning
809:but also including
674:vestibulocerebellum
611:upper motor neurons
420:synaptic plasticity
318:cognitive functions
6721:Granule cell layer
6482:Horizontal fissure
6071:Foster FD (1891).
5295:on 27 January 2012
5070:10.1038/ncponc0794
4502:Horváth G (2003).
4203:10.1007/BF00336192
4066:. Springer-Verlag.
3382:10.1111/jcmm.12965
2808:Scientific Reports
2196:cerebrum posterius
2107:
1981:The cerebellum of
1941:
1923:to these changes.
1921:adult neurogenesis
1904:recreational drugs
1880:multiple sclerosis
1755:
1707:the prion diseases
1643:dysdiadochokinesia
1573:software algorithm
1559:of Pellionisz and
1532:
1408:János Szentágothai
1350:Functional imaging
1254:
1210:interposed nucleus
1181:
1096:
1055:
1042:
979:
740:primary motor area
631:
495:
476:at the base. Four
6871:
6870:
6867:
6866:
6764:
6763:
6670:Cerebellar cortex
6613:
6612:
6603:Cerebellar tonsil
6413:
6412:
6409:
6408:
6174:BrainMaps project
6086:Kraus LA (1844).
5971:978-0-930405-25-0
5838:978-0-444-82313-7
5700:Language Learning
5603:10.1002/cne.21735
5571:978-0-03-910284-5
5347:on 4 January 2015
5321:on 4 January 2015
5217:on 4 January 2015
5191:on 4 January 2015
5165:on 4 January 2015
4887:10.1002/cne.10884
4582:978-0-12-801608-4
3877:978-3-030-99550-8
3623:978-1-84169-044-5
2953:978-0-683-30460-2
2737:978-0-19-515955-4
2657:978-0-8089-2371-8
2613:978-1-315-38276-0
2584:978-0-87893-697-7
2538:Math. Biosciences
2517:978-0-87893-695-3
2332:978-3-540-23735-8
2070:ray-finned fishes
2011:until the middle
1790:) in humans with
1617:Cerebellar ataxia
1381:, which performs
1154:medulla oblongata
1120:vestibular nuclei
1019:refractory period
1014:action potentials
762:. Three types of
682:vestibular nuclei
663:cerebellar vermis
659:cerebrocerebellum
562:cerebellar cortex
440:Marr–Albus theory
330:emotional control
275:
274:
270:
16:(Redirected from
6918:
6775:
6774:
6624:
6623:
6462:
6461:
6440:
6433:
6426:
6417:
6416:
6346:
6345:
6235:
6228:
6221:
6212:
6211:
6155:
6153:
6110:
6109:
6101:
6095:
6094:
6090:(Dritte Auflage)
6083:
6077:
6076:
6068:
6057:
6056:
6048:
6042:
6041:
6033:
6027:
6026:
5982:
5976:
5975:
5957:
5951:
5950:
5940:
5930:
5906:
5900:
5899:
5889:
5857:
5851:
5850:
5816:
5810:
5809:
5773:
5767:
5766:
5730:
5724:
5723:
5695:
5684:
5683:
5673:
5663:
5631:
5625:
5624:
5614:
5582:
5576:
5575:
5557:
5548:
5547:
5511:
5505:
5504:
5468:
5455:
5454:
5444:
5412:
5406:
5405:
5395:
5363:
5357:
5356:
5354:
5352:
5337:
5331:
5330:
5328:
5326:
5311:
5305:
5304:
5302:
5300:
5285:
5279:
5278:
5276:
5274:
5259:
5253:
5252:
5250:
5248:
5233:
5227:
5226:
5224:
5222:
5207:
5201:
5200:
5198:
5196:
5181:
5175:
5174:
5172:
5170:
5155:
5149:
5148:
5138:
5096:
5090:
5089:
5053:
5047:
5046:
5018:
5009:
5008:
4988:
4982:
4981:
4971:
4958:
4957:
4955:
4940:
4922:
4913:
4907:
4906:
4870:
4864:
4863:
4853:
4843:
4815:
4809:
4808:
4798:
4766:
4760:
4759:
4757:
4755:
4740:
4731:
4730:
4720:
4688:
4682:
4681:
4679:
4677:
4657:
4651:
4650:
4648:
4646:
4631:
4622:
4621:
4593:
4587:
4586:
4568:
4562:
4561:
4545:
4535:
4529:
4528:
4526:
4525:
4519:
4513:. Archived from
4508:
4499:
4493:
4492:
4456:
4450:
4449:
4431:
4399:
4393:
4392:
4356:
4350:
4349:
4313:
4307:
4306:
4296:
4294:10.1101/lm.59603
4272:
4266:
4265:
4229:
4223:
4222:
4186:
4180:
4179:
4177:
4171:. Archived from
4162:
4147:Behav. Brain Sci
4144:
4135:
4126:
4125:
4089:
4080:
4074:
4068:
4067:
4065:
4055:
4049:
4048:
4030:
4024:
4023:
4006:(7–8): 961–974.
3995:
3989:
3988:
3978:
3955:Neuropsychologia
3946:
3940:
3939:
3910:
3904:
3903:
3897:
3889:
3862:
3856:
3855:
3845:
3813:
3807:
3806:
3796:
3772:
3766:
3765:
3755:
3731:
3725:
3724:
3714:
3704:
3680:
3674:
3673:
3637:
3628:
3627:
3609:
3603:
3602:
3574:
3568:
3567:
3531:
3525:
3524:
3488:
3461:
3460:
3450:
3415:
3404:
3403:
3393:
3361:
3352:
3351:
3315:
3309:
3308:
3293:Behav. Brain Sci
3288:
3275:
3274:
3264:
3232:
3226:
3225:
3215:
3183:
3177:
3176:
3140:
3134:
3133:
3123:
3099:
3093:
3092:
3056:
3050:
3049:
3031:
3007:
3001:
3000:
2964:
2958:
2957:
2939:
2933:
2932:
2922:
2898:
2892:
2891:
2883:
2858:
2857:
2847:
2799:
2793:
2792:
2782:
2772:
2748:
2742:
2741:
2719:
2662:
2661:
2643:
2637:
2636:
2630:
2622:
2621:
2620:
2595:
2589:
2588:
2570:
2564:
2563:
2553:
2533:
2522:
2521:
2503:
2494:
2493:
2457:
2444:
2443:
2419:
2413:
2412:
2402:
2378:
2372:
2371:
2343:
2337:
2336:
2310:
2302:
2284:
2266:
2262:"The Cerebellum"
2253:
2242:reviewer reports
2231:
2103:Andreas Vesalius
2030:cytoarchitecture
1884:essential tremor
1811:Joubert syndrome
1724:epigenetic clock
1701:. Some forms of
1647:intention tremor
1605:
1591:
1373:, which perform
1327:(AICA), and the
1227:, the source of
991:dendritic spines
924:
849:
713:trigeminal nerve
593:. These are the
503:fourth ventricle
460:At the level of
284:
267:edit on Wikidata
264:
119:
118:
115:
114:
111:
108:
105:
102:
99:
96:
93:
90:
87:
84:
61:
49:
37:
36:
21:
6926:
6925:
6921:
6920:
6919:
6917:
6916:
6915:
6876:
6875:
6872:
6863:
6789:
6760:
6678:Molecular layer
6664:
6609:
6598:Biventer lobule
6510:
6506:Primary fissure
6453:
6444:
6414:
6405:
6396:Parasympathetic
6377:
6335:
6244:
6239:
6118:
6113:
6102:
6098:
6084:
6080:
6069:
6060:
6049:
6045:
6034:
6030:
5983:
5979:
5972:
5958:
5954:
5907:
5903:
5858:
5854:
5839:
5817:
5813:
5774:
5770:
5731:
5727:
5696:
5687:
5646:(10): 3576–80.
5632:
5628:
5583:
5579:
5572:
5558:
5551:
5512:
5508:
5469:
5458:
5413:
5409:
5364:
5360:
5350:
5348:
5339:
5338:
5334:
5324:
5322:
5313:
5312:
5308:
5298:
5296:
5287:
5286:
5282:
5272:
5270:
5261:
5260:
5256:
5246:
5244:
5235:
5234:
5230:
5220:
5218:
5209:
5208:
5204:
5194:
5192:
5183:
5182:
5178:
5168:
5166:
5157:
5156:
5152:
5127:
5097:
5093:
5054:
5050:
5019:
5012:
4989:
4985:
4972:
4961:
4953:
4920:
4914:
4910:
4871:
4867:
4816:
4812:
4767:
4763:
4753:
4751:
4742:
4741:
4734:
4689:
4685:
4675:
4673:
4658:
4654:
4644:
4642:
4633:
4632:
4625:
4594:
4590:
4583:
4569:
4565:
4558:
4536:
4532:
4523:
4521:
4517:
4506:
4500:
4496:
4467:(12): 2949–70.
4457:
4453:
4400:
4396:
4357:
4353:
4314:
4310:
4273:
4269:
4230:
4226:
4187:
4183:
4175:
4160:10.1.1.118.2997
4142:
4136:
4129:
4090:
4083:
4075:
4071:
4056:
4052:
4045:
4031:
4027:
4000:Neural Networks
3996:
3992:
3947:
3943:
3922:(10): 597–609.
3911:
3907:
3891:
3890:
3878:
3864:
3863:
3859:
3814:
3810:
3773:
3769:
3732:
3728:
3681:
3677:
3638:
3631:
3624:
3612:Rapp B (2001).
3610:
3606:
3579:Trends Neurosci
3575:
3571:
3548:10.1038/nrn1347
3532:
3528:
3505:10.1038/nrn1646
3489:
3464:
3416:
3407:
3362:
3355:
3316:
3312:
3289:
3278:
3233:
3229:
3184:
3180:
3151:(2–3): 103–16.
3141:
3137:
3100:
3096:
3057:
3053:
3008:
3004:
2965:
2961:
2954:
2940:
2936:
2899:
2895:
2884:
2861:
2800:
2796:
2749:
2745:
2738:
2720:
2665:
2658:
2644:
2640:
2624:
2623:
2618:
2616:
2614:
2596:
2592:
2585:
2571:
2567:
2534:
2525:
2518:
2504:
2497:
2458:
2447:
2420:
2416:
2393:(Pt 2): 290–2.
2379:
2375:
2344:
2340:
2333:
2311:
2307:
2264:
2256:Marion Wright;
2255:
2249:
2225:
2208:
2184:cerebrum parvum
2164:
2112:
2095:
2058:pyramidal cells
2054:climbing fibers
2050:cartwheel cells
2046:parallel fibers
2026:
1937:porbeagle shark
1929:
1912:
1792:Gorlin Syndrome
1788:medulloblastoma
1760:ultrasound scan
1744:
1736:cerebral cortex
1728:gene expression
1719:
1619:
1613:
1612:
1611:
1610:
1609:
1606:
1597:
1596:
1595:
1592:
1581:
1521:
1497:optimal control
1493:Adaptive Filter
1480:climbing fibers
1467:
1391:
1379:cerebral cortex
1342:
1317:
1289:input from the
1280:parallel fibers
1246:
1229:climbing fibers
1173:
1167:
1150:climbing fibers
1142:
1140:Climbing fibers
1109:
1047:
1031:newly generated
971:
950:dendritic trees
946:
944:Molecular layer
941:
936:
935:
934:
933:
932:
925:
917:
916:
915:
914:
909:
904:
899:
894:
889:
884:
882:
877:
872:
867:
862:
860:
857:
853:
850:
776:parallel fibers
772:climbing fibers
752:
736:premotor cortex
708:) and from the
704:(including the
655:spinocerebellum
643:primary fissure
623:
607:thalamic nuclei
531:rhombencephalon
487:
458:
452:
432:parallel fibers
377:cerebral cortex
338:sensory systems
271:
81:
77:
64:
52:
35:
28:
23:
22:
15:
12:
11:
5:
6924:
6914:
6913:
6908:
6903:
6898:
6893:
6888:
6869:
6868:
6865:
6864:
6862:
6861:
6856:
6846:
6845:
6835:
6834:
6829:
6819:
6814:
6809:
6799:
6797:
6791:
6790:
6788:
6787:
6781:
6779:
6772:
6766:
6765:
6762:
6761:
6759:
6758:
6756:Parallel fiber
6753:
6751:Climbing fiber
6748:
6741:
6740:
6739:
6738:
6733:
6728:
6717:
6716:
6715:
6714:
6709:
6704:
6693:
6692:
6691:
6690:
6685:
6674:
6672:
6666:
6665:
6663:
6662:
6657:
6656:
6655:
6650:
6638:
6632:
6630:
6621:
6615:
6614:
6611:
6610:
6608:
6607:
6606:
6605:
6600:
6590:
6589:
6588:
6573:
6572:
6567:
6566:
6565:
6560:
6555:
6545:
6544:
6543:
6538:
6533:
6531:Central lobule
6518:
6516:
6515:Medial/lateral
6512:
6511:
6509:
6508:
6503:
6502:
6501:
6496:
6486:
6485:
6484:
6477:Posterior lobe
6474:
6468:
6466:
6459:
6455:
6454:
6443:
6442:
6435:
6428:
6420:
6411:
6410:
6407:
6406:
6404:
6403:
6398:
6393:
6387:
6385:
6379:
6378:
6376:
6375:
6370:
6368:Cranial nerves
6365:
6360:
6354:
6352:
6343:
6337:
6336:
6334:
6333:
6332:
6331:
6326:
6325:
6324:
6319:
6318:
6317:
6312:
6297:
6292:
6291:
6290:
6285:
6280:
6265:
6260:
6254:
6252:
6246:
6245:
6242:Nervous system
6238:
6237:
6230:
6223:
6215:
6209:
6208:
6202:
6197:
6192:
6186:
6181:
6176:
6167:
6161:
6156:
6117:
6116:External links
6114:
6112:
6111:
6096:
6078:
6058:
6043:
6028:
5977:
5970:
5952:
5901:
5872:(6): 2321–30.
5852:
5837:
5811:
5784:(6): 491–507.
5768:
5725:
5685:
5626:
5577:
5570:
5549:
5506:
5456:
5427:(2): 197–220.
5407:
5358:
5332:
5306:
5280:
5254:
5228:
5202:
5176:
5150:
5125:
5091:
5064:(5): 295–304.
5048:
5010:
4983:
4959:
4908:
4865:
4810:
4781:(5): 294–306.
4761:
4732:
4683:
4652:
4623:
4604:(13): 889–96.
4588:
4581:
4563:
4556:
4530:
4494:
4451:
4394:
4351:
4308:
4267:
4234:Brain Research
4224:
4197:(3): 195–206.
4181:
4178:on 2017-08-09.
4153:(3): 368–383.
4127:
4081:
4069:
4050:
4043:
4025:
3990:
3961:(6): 1331–41.
3941:
3918:(in Spanish).
3905:
3876:
3857:
3828:(5): 2322–45.
3808:
3787:(5): 1041–50.
3767:
3726:
3675:
3629:
3622:
3604:
3569:
3526:
3499:(4): 297–311.
3462:
3405:
3376:(2): 299–314.
3353:
3310:
3299:(3): 384–398.
3276:
3227:
3178:
3135:
3094:
3051:
3002:
2959:
2952:
2934:
2913:(6): 331–357.
2893:
2859:
2794:
2743:
2736:
2663:
2656:
2638:
2612:
2590:
2583:
2565:
2551:10.1.1.14.7524
2544:(1–2): 25–61.
2523:
2516:
2495:
2445:
2434:(688): 62–75.
2414:
2373:
2338:
2331:
2304:
2258:William Skaggs
2224:
2221:
2220:
2219:
2214:
2207:
2204:
2163:
2160:
2111:
2108:
2094:
2091:
2086:James M. Bower
2042:auditory nerve
2034:neurochemistry
2025:
2022:
1963:with a brain.
1928:
1925:
1911:
1908:
1900:pharmaceutical
1779:sonic hedgehog
1752:cisterna magna
1743:
1740:
1718:
1715:
1679:cerebral edema
1615:Main article:
1607:
1600:
1599:
1598:
1593:
1586:
1585:
1584:
1583:
1582:
1580:
1577:
1520:
1517:
1484:parallel fiber
1466:
1463:
1462:
1461:
1455:
1449:
1441:Purkinje cells
1437:parallel fiber
1422:
1400:John C. Eccles
1390:
1387:
1359:functional MRI
1341:
1338:
1316:
1313:
1287:climbing fiber
1271:immunostaining
1245:
1242:
1169:Main article:
1166:
1163:
1141:
1138:
1116:pontine nuclei
1108:
1105:
1077:parallel fiber
1062:dendritic claw
1046:
1045:Granular layer
1043:
995:Purkinje layer
982:Purkinje cells
970:
969:Purkinje layer
967:
955:stellate cells
945:
942:
940:
937:
926:
919:
918:
892:Parallel fiber
880:Climbing fiber
875:Inferior olive
858:
855:
854:
851:
844:
843:
842:
841:
840:
828:stellate cells
756:Purkinje cells
751:
748:
728:pontine nuclei
698:proprioceptive
647:posterior lobe
622:
619:
615:pontine nuclei
486:
483:
454:Main article:
451:
448:
436:climbing fiber
412:motor learning
393:Purkinje cells
362:motor learning
273:
272:
263:
257:
256:
251:
245:
244:
239:
233:
232:
227:
221:
220:
215:
208:
207:
202:
196:
195:
190:
184:
183:
178:
172:
171:
167:
166:
157:
151:
150:
137:
131:
130:
125:
121:
120:
75:
71:
70:
66:
65:
62:
54:
53:
50:
42:
41:
26:
18:Folium (brain)
9:
6:
4:
3:
2:
6923:
6912:
6909:
6907:
6904:
6902:
6899:
6897:
6894:
6892:
6889:
6887:
6884:
6883:
6881:
6874:
6860:
6857:
6855:
6851:
6848:
6847:
6844:
6840:
6837:
6836:
6833:
6830:
6827:
6823:
6820:
6818:
6815:
6813:
6810:
6808:
6804:
6801:
6800:
6798:
6796:
6792:
6786:
6783:
6782:
6780:
6776:
6773:
6771:
6767:
6757:
6754:
6752:
6749:
6747:
6743:
6742:
6737:
6734:
6732:
6729:
6727:
6724:
6723:
6722:
6719:
6718:
6713:
6710:
6708:
6705:
6703:
6702:Purkinje cell
6700:
6699:
6698:
6695:
6694:
6689:
6686:
6684:
6683:Stellate cell
6681:
6680:
6679:
6676:
6675:
6673:
6671:
6667:
6661:
6658:
6654:
6651:
6649:
6646:
6645:
6644:
6643:
6639:
6637:
6634:
6633:
6631:
6629:
6625:
6622:
6620:
6616:
6604:
6601:
6599:
6596:
6595:
6594:
6591:
6587:
6584:
6583:
6582:
6578:
6575:
6574:
6571:
6568:
6564:
6561:
6559:
6556:
6554:
6551:
6550:
6549:
6546:
6542:
6539:
6537:
6534:
6532:
6529:
6528:
6527:
6523:
6520:
6519:
6517:
6513:
6507:
6504:
6500:
6497:
6495:
6492:
6491:
6490:
6487:
6483:
6480:
6479:
6478:
6475:
6473:
6472:Anterior lobe
6470:
6469:
6467:
6463:
6460:
6456:
6452:
6448:
6441:
6436:
6434:
6429:
6427:
6422:
6421:
6418:
6402:
6399:
6397:
6394:
6392:
6389:
6388:
6386:
6384:
6380:
6374:
6371:
6369:
6366:
6364:
6361:
6359:
6358:Sensory nerve
6356:
6355:
6353:
6351:
6347:
6344:
6342:
6338:
6330:
6329:Limbic system
6327:
6323:
6320:
6316:
6313:
6311:
6308:
6307:
6306:
6303:
6302:
6301:
6298:
6296:
6293:
6289:
6286:
6284:
6281:
6279:
6276:
6275:
6274:
6271:
6270:
6269:
6266:
6264:
6261:
6259:
6256:
6255:
6253:
6251:
6247:
6243:
6236:
6231:
6229:
6224:
6222:
6217:
6216:
6213:
6206:
6203:
6201:
6198:
6196:
6193:
6190:
6187:
6185:
6182:
6180:
6177:
6175:
6171:
6168:
6165:
6162:
6160:
6157:
6152:
6147:
6143:
6139:
6135:
6131:
6130:
6125:
6120:
6119:
6107:
6100:
6092:
6089:
6082:
6074:
6067:
6065:
6063:
6054:
6047:
6039:
6032:
6024:
6020:
6016:
6012:
6008:
6004:
6000:
5996:
5993:(1): 273–88.
5992:
5988:
5981:
5973:
5967:
5963:
5956:
5948:
5944:
5939:
5934:
5929:
5924:
5920:
5916:
5912:
5905:
5897:
5893:
5888:
5883:
5879:
5875:
5871:
5867:
5863:
5856:
5848:
5844:
5840:
5834:
5830:
5826:
5822:
5815:
5807:
5803:
5799:
5795:
5791:
5787:
5783:
5779:
5772:
5764:
5760:
5756:
5752:
5748:
5744:
5741:(4): 401–29.
5740:
5736:
5729:
5721:
5717:
5713:
5709:
5705:
5701:
5694:
5692:
5690:
5681:
5677:
5672:
5667:
5662:
5657:
5653:
5649:
5645:
5641:
5637:
5630:
5622:
5618:
5613:
5608:
5604:
5600:
5597:(5): 449–73.
5596:
5592:
5588:
5581:
5573:
5567:
5563:
5556:
5554:
5545:
5541:
5537:
5533:
5529:
5525:
5522:(2): 329–45.
5521:
5517:
5510:
5502:
5498:
5494:
5490:
5486:
5482:
5478:
5474:
5467:
5465:
5463:
5461:
5452:
5448:
5443:
5438:
5434:
5430:
5426:
5422:
5418:
5411:
5403:
5399:
5394:
5389:
5385:
5381:
5377:
5373:
5369:
5362:
5346:
5342:
5336:
5320:
5316:
5310:
5294:
5290:
5284:
5268:
5264:
5258:
5242:
5238:
5232:
5216:
5212:
5206:
5190:
5186:
5180:
5164:
5160:
5154:
5146:
5142:
5137:
5132:
5128:
5126:9780123809162
5122:
5118:
5114:
5110:
5106:
5102:
5095:
5087:
5083:
5079:
5075:
5071:
5067:
5063:
5059:
5052:
5044:
5040:
5036:
5032:
5028:
5024:
5017:
5015:
5006:
5002:
4999:(2): 247–50.
4998:
4994:
4987:
4979:
4978:
4970:
4968:
4966:
4964:
4952:
4948:
4944:
4939:
4934:
4931:(6): 1161–7.
4930:
4926:
4919:
4912:
4904:
4900:
4896:
4892:
4888:
4884:
4881:(3): 356–65.
4880:
4876:
4869:
4861:
4857:
4852:
4847:
4842:
4837:
4833:
4829:
4825:
4821:
4814:
4806:
4802:
4797:
4792:
4788:
4784:
4780:
4776:
4772:
4765:
4749:
4745:
4739:
4737:
4728:
4724:
4719:
4714:
4710:
4706:
4703:(6): 820–33.
4702:
4698:
4694:
4687:
4671:
4667:
4663:
4656:
4640:
4636:
4630:
4628:
4619:
4615:
4611:
4607:
4603:
4599:
4592:
4584:
4578:
4574:
4567:
4559:
4557:9780803612471
4553:
4549:
4544:
4543:
4534:
4520:on 2020-05-20
4516:
4512:
4505:
4498:
4490:
4486:
4482:
4478:
4474:
4470:
4466:
4462:
4455:
4447:
4443:
4439:
4435:
4430:
4425:
4421:
4417:
4414:(1): 302–17.
4413:
4409:
4405:
4398:
4390:
4386:
4382:
4378:
4374:
4370:
4366:
4362:
4355:
4347:
4343:
4339:
4335:
4331:
4327:
4323:
4319:
4312:
4304:
4300:
4295:
4290:
4287:(6): 427–55.
4286:
4282:
4278:
4271:
4263:
4259:
4255:
4251:
4247:
4243:
4240:(2): 309–28.
4239:
4235:
4228:
4220:
4216:
4212:
4208:
4204:
4200:
4196:
4192:
4185:
4174:
4170:
4166:
4161:
4156:
4152:
4148:
4141:
4134:
4132:
4123:
4119:
4115:
4111:
4107:
4103:
4099:
4095:
4088:
4086:
4078:
4073:
4064:
4063:
4054:
4046:
4044:9783031060922
4040:
4036:
4029:
4021:
4017:
4013:
4009:
4005:
4001:
3994:
3986:
3982:
3977:
3972:
3968:
3964:
3960:
3956:
3952:
3945:
3937:
3933:
3929:
3925:
3921:
3917:
3909:
3901:
3895:
3887:
3883:
3879:
3873:
3869:
3868:
3861:
3853:
3849:
3844:
3839:
3835:
3831:
3827:
3823:
3819:
3812:
3804:
3800:
3795:
3790:
3786:
3782:
3778:
3771:
3763:
3759:
3754:
3749:
3746:(4): 561–79.
3745:
3741:
3737:
3730:
3722:
3718:
3713:
3708:
3703:
3698:
3694:
3690:
3686:
3679:
3671:
3667:
3663:
3659:
3655:
3651:
3647:
3643:
3636:
3634:
3625:
3619:
3615:
3608:
3600:
3596:
3592:
3588:
3584:
3580:
3573:
3565:
3561:
3557:
3553:
3549:
3545:
3541:
3537:
3530:
3522:
3518:
3514:
3510:
3506:
3502:
3498:
3494:
3487:
3485:
3483:
3481:
3479:
3477:
3475:
3473:
3471:
3469:
3467:
3458:
3454:
3449:
3444:
3440:
3436:
3433:(2): 437–70.
3432:
3428:
3424:
3421:(June 1969).
3420:
3414:
3412:
3410:
3401:
3397:
3392:
3387:
3383:
3379:
3375:
3371:
3367:
3360:
3358:
3349:
3345:
3341:
3337:
3333:
3329:
3325:
3321:
3314:
3306:
3302:
3298:
3294:
3287:
3285:
3283:
3281:
3272:
3268:
3263:
3258:
3254:
3250:
3247:(2): 268–96.
3246:
3242:
3238:
3231:
3223:
3219:
3214:
3209:
3205:
3201:
3198:(2): 159–67.
3197:
3193:
3189:
3182:
3174:
3170:
3166:
3162:
3158:
3154:
3150:
3146:
3139:
3131:
3127:
3122:
3117:
3114:(3): 807–15.
3113:
3109:
3105:
3098:
3090:
3086:
3082:
3078:
3074:
3070:
3066:
3062:
3055:
3047:
3043:
3039:
3035:
3030:
3025:
3022:(3): 159–62.
3021:
3017:
3013:
3006:
2998:
2994:
2990:
2986:
2982:
2978:
2975:(2): 153–70.
2974:
2970:
2963:
2955:
2949:
2945:
2938:
2930:
2926:
2921:
2916:
2912:
2908:
2904:
2897:
2889:
2882:
2880:
2878:
2876:
2874:
2872:
2870:
2868:
2866:
2864:
2855:
2851:
2846:
2841:
2837:
2833:
2829:
2825:
2821:
2817:
2813:
2809:
2805:
2798:
2790:
2786:
2781:
2776:
2771:
2766:
2762:
2758:
2754:
2747:
2739:
2733:
2729:
2725:
2718:
2716:
2714:
2712:
2710:
2708:
2706:
2704:
2702:
2700:
2698:
2696:
2694:
2692:
2690:
2688:
2686:
2684:
2682:
2680:
2678:
2676:
2674:
2672:
2670:
2668:
2659:
2653:
2649:
2642:
2634:
2628:
2615:
2609:
2605:
2601:
2594:
2586:
2580:
2576:
2569:
2561:
2557:
2552:
2547:
2543:
2539:
2532:
2530:
2528:
2519:
2513:
2509:
2502:
2500:
2491:
2487:
2483:
2479:
2475:
2471:
2468:(4): 375–84.
2467:
2463:
2456:
2454:
2452:
2450:
2441:
2437:
2433:
2429:
2425:
2418:
2410:
2406:
2401:
2396:
2392:
2388:
2384:
2377:
2369:
2365:
2361:
2357:
2354:(3): 245–53.
2353:
2349:
2342:
2334:
2328:
2324:
2320:
2316:
2309:
2305:
2303:
2300:
2296:
2292:
2288:
2283:
2278:
2274:
2270:
2263:
2259:
2252:
2247:
2243:
2239:
2236:for external
2235:
2230:
2218:
2215:
2213:
2210:
2209:
2203:
2201:
2200:parencephalis
2197:
2193:
2189:
2185:
2181:
2180:parenkephalis
2177:
2173:
2169:
2159:
2156:
2152:
2151:Luigi Rolando
2148:
2147:Thomas Willis
2144:
2140:
2135:
2133:
2129:
2128:parenkephalis
2125:
2121:
2117:
2104:
2099:
2090:
2087:
2083:
2077:
2075:
2071:
2066:
2061:
2059:
2055:
2051:
2047:
2043:
2039:
2035:
2031:
2021:
2019:
2014:
2010:
2009:
2002:
2000:
1996:
1995:mormyrid fish
1992:
1988:
1984:
1983:cartilaginous
1979:
1977:
1973:
1969:
1964:
1962:
1958:
1954:
1950:
1946:
1938:
1933:
1924:
1922:
1916:
1907:
1905:
1901:
1897:
1893:
1889:
1885:
1881:
1877:
1872:
1870:
1866:
1862:
1859:) as seen in
1858:
1854:
1851:
1847:
1843:
1838:
1836:
1832:
1828:
1824:
1820:
1816:
1815:may be absent
1812:
1808:
1804:
1799:
1797:
1793:
1789:
1785:
1780:
1775:
1773:
1769:
1765:
1761:
1753:
1748:
1739:
1737:
1733:
1729:
1725:
1714:
1712:
1708:
1704:
1700:
1696:
1692:
1688:
1684:
1680:
1676:
1672:
1667:
1665:
1661:
1656:
1654:
1653:
1648:
1644:
1640:
1636:
1632:
1628:
1624:
1618:
1604:
1590:
1576:
1574:
1570:
1564:
1562:
1558:
1554:
1550:
1545:
1543:
1538:
1530:
1525:
1516:
1513:
1508:
1506:
1500:
1498:
1494:
1489:
1488:Purkinje cell
1485:
1481:
1477:
1473:
1459:
1456:
1453:
1450:
1446:
1442:
1438:
1434:
1433:granule cells
1430:
1426:
1423:
1420:
1416:
1413:
1412:
1411:
1409:
1405:
1401:
1397:
1386:
1384:
1380:
1376:
1372:
1371:basal ganglia
1368:
1362:
1360:
1356:
1351:
1346:
1337:
1334:
1330:
1326:
1322:
1312:
1308:
1305:
1301:
1297:
1296:gap junctions
1292:
1288:
1283:
1281:
1275:
1272:
1266:
1264:
1260:
1250:
1241:
1237:
1234:
1230:
1226:
1222:
1218:
1213:
1211:
1207:
1203:
1199:
1195:
1191:
1186:
1177:
1172:
1162:
1158:
1155:
1151:
1147:
1137:
1134:
1130:
1126:
1121:
1117:
1113:
1104:
1102:
1093:
1088:
1084:
1082:
1078:
1074:
1069:
1067:
1063:
1059:
1051:
1038:
1034:
1032:
1028:
1023:
1020:
1015:
1011:
1010:extracellular
1006:
1004:
1000:
996:
992:
987:
983:
975:
966:
964:
960:
956:
951:
930:
923:
913:
908:
907:Stellate cell
903:
898:
897:Purkinje cell
893:
888:
881:
876:
871:
866:
848:
839:
837:
833:
829:
824:
820:
816:
812:
808:
804:
799:
797:
793:
792:noradrenergic
789:
785:
779:
777:
773:
769:
765:
761:
760:granule cells
757:
747:
745:
741:
737:
733:
729:
725:
724:parietal lobe
720:
718:
714:
711:
707:
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699:
693:
691:
687:
683:
679:
675:
670:
668:
664:
660:
656:
652:
648:
644:
640:
639:anterior lobe
636:
627:
618:
616:
612:
608:
604:
600:
596:
592:
587:
585:
581:
580:
575:
571:
567:
563:
559:
554:
552:
548:
544:
543:granule cells
540:
536:
532:
528:
527:metencephalon
524:
520:
516:
512:
508:
504:
500:
491:
485:Gross anatomy
482:
479:
475:
471:
467:
463:
462:gross anatomy
457:
447:
445:
441:
437:
433:
429:
425:
421:
417:
413:
408:
406:
403:lying in the
402:
398:
397:granule cells
394:
390:
386:
382:
378:
374:
370:
365:
363:
359:
355:
351:
350:fine movement
347:
343:
339:
335:
331:
327:
323:
319:
315:
314:motor control
311:
308:
304:
300:
296:
292:
288:
280:
268:
262:
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255:
252:
250:
246:
243:
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74:Pronunciation
72:
67:
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6911:Motor system
6873:
6852:(midbrain):
6770:White matter
6746:Mossy fibers
6731:Granule cell
6720:
6712:Fañanas cell
6696:
6677:
6669:
6640:
6592:
6580:
6547:
6525:
6450:
6373:Spinal nerve
6305:Diencephalon
6287:
6133:
6129:Scholarpedia
6127:
6124:"Cerebellum"
6105:
6099:
6091:
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5375:
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5345:the original
5335:
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5319:the original
5309:
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5293:the original
5283:
5271:. Retrieved
5267:the original
5257:
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5231:
5219:. Retrieved
5215:the original
5205:
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5189:the original
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4827:
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4700:
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4515:the original
4510:
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4461:Neuroscience
4460:
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2641:
2617:, retrieved
2603:
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2575:Neuroscience
2574:
2568:
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2508:Neuroscience
2507:
2465:
2461:
2431:
2427:
2417:
2390:
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2376:
2351:
2347:
2341:
2314:
2308:
2272:
2268:
2246:CC-BY-SA-3.0
2233:
2226:
2199:
2195:
2191:
2187:
2183:
2179:
2176:little brain
2175:
2171:
2167:
2165:
2136:
2131:
2127:
2113:
2110:Descriptions
2078:
2074:optic tectum
2065:lateral line
2062:
2027:
2008:Homo sapiens
2006:
2003:
1998:
1980:
1965:
1942:
1917:
1913:
1896:heavy metals
1873:
1871:deficiency.
1839:
1800:
1796:mouse models
1776:
1756:
1720:
1695:degenerative
1668:
1657:
1650:
1627:lateral zone
1620:
1565:
1553:Richard Ivry
1546:
1533:
1509:
1501:
1492:
1468:
1457:
1451:
1445:deep nuclear
1424:
1414:
1395:
1392:
1363:
1347:
1343:
1318:
1315:Blood supply
1309:
1304:basket cells
1284:
1276:
1267:
1262:
1258:
1255:
1244:Compartments
1238:
1214:
1182:
1159:
1143:
1124:
1112:Mossy fibers
1110:
1107:Mossy fibers
1097:
1076:
1073:unmyelinated
1070:
1061:
1056:
1024:
1007:
994:
980:
959:basket cells
947:
887:Granule cell
832:basket cells
811:Lugaro cells
803:interneurons
800:
788:serotonergic
784:dopaminergic
780:
768:mossy fibers
753:
750:Microanatomy
721:
694:
671:
666:
632:
621:Subdivisions
588:
577:
570:white matter
565:
561:
555:
496:
470:white matter
459:
409:
405:white matter
366:
334:coordination
290:
286:
278:
276:
230:A14.1.07.001
218:birnlex_1489
180:
6805:(medulla):
6785:Arbor vitae
6688:Basket cell
6619:Grey matter
6391:Sympathetic
6363:Motor nerve
6315:Optic nerve
6263:Spinal cord
6136:(1): 4606.
5706:: 162–186.
5029:: 385–408.
4834:(9): e274.
4676:January 21,
4324:(1): 1–33.
4100:: 581–609.
3585:: 143–145.
3326:(1): 42–7.
2814:(1): 5622.
2188:encephalion
2139:Renaissance
2122:(quoted in
2013:Pleistocene
1987:bony fishes
1953:cephalopods
1949:vertebrates
1772:sensitivity
1732:age-related
1689:, physical
1529:James Albus
1476:James Albus
1429:mossy fiber
1419:feedforward
1323:(SCA), the
1185:deep nuclei
1165:Deep nuclei
1133:Golgi cells
1066:Golgi cells
912:Basket cell
865:Mossy fiber
807:Golgi cells
796:cholinergic
744:red nucleus
702:spinal cord
641:(above the
579:arbor vitae
558:gray matter
428:James Albus
401:deep nuclei
364:in humans.
354:equilibrium
342:spinal cord
328:as well as
299:vertebrates
291:cerebellums
170:Identifiers
6906:Cerebellum
6880:Categories
6726:Golgi cell
6648:Emboliform
6642:interposed
6577:Hemisphere
6451:cerebellum
6288:Cerebellum
5421:Cerebellum
5273:1 February
5111:: 235–82.
4754:2 February
4645:2 February
4524:2009-12-24
3886:1338132789
3067:: 413–34.
3016:Cerebellum
2724:Cerebellum
2619:2024-08-23
2223:References
2168:cerebellum
2132:enkephalos
2120:Herophilus
1968:amphibians
1848:, and the
1803:hypoplasia
1766:for fetal
1687:alcoholism
1675:hemorrhage
1635:dysarthria
1569:perceptron
1472:David Marr
1458:Plasticity
1452:Modularity
1389:Principles
1377:, and the
1333:anastomose
1263:microzones
1202:emboliform
1101:David Marr
1092:glomerulus
1071:The thin,
902:Golgi cell
726:) via the
574:myelinated
519:dura mater
424:David Marr
279:cerebellum
200:NeuroNames
181:cerebellum
40:Cerebellum
6795:Peduncles
6660:Fastigial
6593:posterior
6548:posterior
6494:Flocculus
6383:Autonomic
6300:Forebrain
6273:Hindbrain
5351:9 January
5325:9 January
5299:9 January
5247:9 January
5221:9 January
5195:9 January
5169:9 January
4155:CiteSeerX
3894:cite book
2929:146700933
2836:2045-2322
2546:CiteSeerX
2490:260317107
2299:Q44001486
2291:2002-4436
2248:license (
2240:in 2016 (
2192:encranion
2166:The name
2162:Etymology
2116:Aristotle
2018:Hominidae
1957:octopuses
1869:vitamin E
1639:dysmetria
1631:hypotonia
1404:Masao Ito
1217:glutamate
1206:fastigial
1129:glomeruli
1081:glutamate
1027:calbindin
963:GABAergic
836:GABAergic
823:dendritic
805:, mainly
551:neocortex
474:ventricle
450:Structure
385:accordion
322:attention
295:hindbrain
287:cerebella
128:Hindbrain
6850:Superior
6841:(pons):
6803:Inferior
6778:Internal
6744:Fibers:
6581:anterior
6526:anterior
6322:Cerebrum
6295:Midbrain
6258:Meninges
6108:. Fürth.
6023:22860609
6015:12582060
5947:23970855
5896:24501371
5806:14393814
5798:18491162
5763:12727461
5720:22230641
5680:15731345
5621:18537139
5544:43112389
5501:14536411
5493:18275284
5479:: 1–24.
5451:25479821
5402:20553761
5241:Archived
5145:21295689
5086:24461280
5078:17464337
4951:Archived
4947:11415913
4895:14556293
4860:16048372
4805:26000617
4727:17578530
4697:Headache
4670:Archived
4639:Archived
4489:20520737
4446:27237058
4438:12582062
4389:36802745
4381:10096998
4338:13563670
4303:14657256
4262:40799652
4114:15217344
4079:, p. 311
4020:12662639
3985:17123557
3936:21069639
3852:21795627
3803:10775548
3721:26331045
3670:10962570
3662:11240282
3599:53272245
3564:30232749
3556:14976523
3521:10769826
3513:15803161
3400:27665307
3348:10505177
3340:17961663
3222:24403568
3165:15003388
3130:24183029
3081:19555291
3046:25671398
3038:17786810
2997:35159122
2989:23968463
2854:38453991
2845:10920891
2789:20300467
2627:citation
2482:12539058
2409:16434422
2368:19776302
2295:Wikidata
2206:See also
2172:cerebrum
2143:Vesalius
1972:lampreys
1861:beriberi
1857:thiamine
1703:migraine
1499:theory.
1465:Learning
1435:, whose
1340:Function
1233:feedback
1125:rosettes
732:thalamus
717:auditory
601:and the
515:cerebrum
373:cortical
326:language
320:such as
307:mormyrid
303:cerebrum
212:NeuroLex
164:inferior
160:Superior
32:Cerebrum
6653:Globose
6636:Dentate
6541:Lingula
6458:Surface
6449:of the
6447:Anatomy
6401:Enteric
6350:Somatic
6278:Medulla
6172:at the
6138:Bibcode
5995:Bibcode
5938:3748751
5921:: 133.
5887:3913874
5847:9193161
5743:Bibcode
5648:Bibcode
5612:5884697
5442:4346664
5393:2943015
5136:3213765
5043:7605067
5005:3299184
4938:7974784
4903:7091227
4851:1181540
4820:Pääbo S
4796:4468311
4718:3761082
4618:9516225
4481:7162624
4416:Bibcode
4346:8989536
4219:3695770
4211:7171642
4122:1310007
3976:1868674
3843:3214121
3762:9577385
3712:4552302
3457:5784296
3448:1351491
3391:5264155
3271:5944665
3262:1357472
3213:4103644
3173:7352039
3089:1066141
2816:Bibcode
2780:2839851
2105:in 1543
2093:History
2005:and in
1999:valvula
1976:hagfish
1961:species
1945:classes
1863:and in
1770:with a
1240:cats).
1198:globose
1194:dentate
710:cranial
678:balance
645:), the
547:neurons
511:medulla
468:, with
389:neurons
358:posture
340:of the
297:of all
193:D002531
124:Part of
69:Details
6839:Middle
6553:Folium
6536:Culmen
6522:Vermis
6499:Nodule
6310:Retina
6021:
6013:
5968:
5945:
5935:
5894:
5884:
5845:
5835:
5804:
5796:
5761:
5718:
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5671:553338
5668:
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4803:
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4554:
4550:–225.
4487:
4479:
4444:
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4387:
4379:
4344:
4336:
4301:
4260:
4254:194656
4252:
4217:
4209:
4157:
4120:
4112:
4041:
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3934:
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3562:
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3519:
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3455:
3445:
3419:Marr D
3398:
3388:
3346:
3338:
3269:
3259:
3220:
3210:
3171:
3163:
3128:
3108:Neuron
3087:
3079:
3044:
3036:
2995:
2987:
2950:
2927:
2852:
2842:
2834:
2787:
2777:
2763:: 12.
2734:
2654:
2610:
2581:
2548:
2514:
2488:
2480:
2407:
2366:
2329:
2297:
2289:
2198:, and
1974:, and
1890:, and
1825:, and
1764:screen
1691:trauma
1683:tumors
1671:stroke
1652:ataxia
1561:Llinás
1448:cells.
1406:, and
1204:, and
1190:nuclei
929:folium
794:, and
667:Vermis
597:, the
566:folium
560:: the
539:vermis
521:, the
501:. The
466:cortex
381:tissue
371:. Its
360:, and
310:fishes
135:Artery
6563:Uvula
6558:Tuber
6465:Lobes
6268:Brain
6019:S2CID
5802:S2CID
5540:S2CID
5497:S2CID
5082:S2CID
4954:(PDF)
4921:(PDF)
4899:S2CID
4775:Aging
4518:(PDF)
4507:(PDF)
4485:S2CID
4442:S2CID
4385:S2CID
4342:S2CID
4258:S2CID
4215:S2CID
4176:(PDF)
4143:(PDF)
4118:S2CID
3781:Brain
3740:Brain
3695:: 2.
3666:S2CID
3595:S2CID
3560:S2CID
3517:S2CID
3344:S2CID
3169:S2CID
3085:S2CID
3042:S2CID
2993:S2CID
2925:S2CID
2486:S2CID
2387:Brain
2275:(1).
2265:(PDF)
2124:Galen
1867:, or
1717:Aging
1259:zones
764:axons
635:lobes
346:brain
265:[
254:67944
176:Latin
6283:Pons
6011:PMID
5966:ISBN
5943:PMID
5892:PMID
5843:PMID
5833:ISBN
5794:PMID
5759:PMID
5716:PMID
5676:PMID
5617:PMID
5566:ISBN
5532:PMID
5489:PMID
5447:PMID
5398:PMID
5353:2015
5327:2015
5301:2015
5275:2015
5249:2015
5223:2015
5197:2015
5171:2015
5141:PMID
5121:ISBN
5074:PMID
5039:PMID
5001:PMID
4943:PMID
4891:PMID
4856:PMID
4801:PMID
4756:2015
4723:PMID
4678:2016
4647:2015
4614:PMID
4577:ISBN
4552:ISBN
4477:PMID
4434:PMID
4377:PMID
4334:PMID
4299:PMID
4250:PMID
4207:PMID
4110:PMID
4039:ISBN
4016:PMID
3981:PMID
3932:PMID
3900:link
3882:OCLC
3872:ISBN
3848:PMID
3799:PMID
3758:PMID
3717:PMID
3658:PMID
3618:ISBN
3552:PMID
3509:PMID
3453:PMID
3396:PMID
3336:PMID
3267:PMID
3218:PMID
3161:PMID
3126:PMID
3077:PMID
3034:PMID
2985:PMID
2948:ISBN
2850:PMID
2832:ISSN
2785:PMID
2732:ISBN
2652:ISBN
2633:link
2608:ISBN
2579:ISBN
2512:ISBN
2478:PMID
2405:PMID
2364:PMID
2327:ISBN
2287:ISSN
2251:2016
2118:and
2032:and
1985:and
1910:Pain
1844:and
1809:and
1474:and
1221:GABA
1183:The
1003:GABA
957:and
830:and
813:and
770:and
758:and
738:and
690:gait
688:and
509:and
507:pons
426:and
395:and
324:and
316:and
277:The
242:5788
225:TA98
188:MeSH
155:Vein
148:PICA
144:AICA
6146:doi
6003:doi
5991:978
5933:PMC
5923:doi
5882:PMC
5874:doi
5825:doi
5786:doi
5751:doi
5708:doi
5666:PMC
5656:doi
5644:102
5607:PMC
5599:doi
5595:509
5524:doi
5520:174
5481:doi
5437:PMC
5429:doi
5388:PMC
5380:doi
5131:PMC
5113:doi
5066:doi
5031:doi
4933:PMC
4883:doi
4879:466
4846:PMC
4836:doi
4791:PMC
4783:doi
4713:PMC
4705:doi
4606:doi
4602:338
4548:193
4469:doi
4424:doi
4412:978
4369:doi
4326:doi
4322:109
4289:doi
4242:doi
4238:128
4199:doi
4165:doi
4102:doi
4008:doi
3971:PMC
3963:doi
3924:doi
3838:PMC
3830:doi
3826:106
3789:doi
3785:123
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