Cerebellum - Knowledge

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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. 1515:

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.

974: 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: 1524: 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: 1249: 59: 1050: 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.

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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

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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

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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.

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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

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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

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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

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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

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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

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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: 1235:

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. 1293:

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

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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. 825:

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

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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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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.

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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.

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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

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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

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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

6890: 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. 549:

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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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 (

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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 5210: 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

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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 5340: 1705:

headache may also produce temporary dysfunction of the cerebellum, of variable severity. Infection can result in cerebellar damage in such conditions as

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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).

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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

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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

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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?".

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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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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

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that rise to the superficial layer and travel across it horizontally. The superficial layer contains a set of GABAergic neurons called

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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".

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or Schmahmann's syndrome has been described in adults and children. Estimates based on functional mapping of the cerebellum using

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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: 1418: 1354: 4769:

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).

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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: 4747: 1324: 1179:

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: 5733:

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: 3998:

Doya K (October 1999). "What are the computations of the cerebellum, the basal ganglia and the cerebral cortex?".

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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".

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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: 5021:

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

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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: 2460:

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

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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: 6471: 3418: 2211: 1845: 1771: 1504: 1471: 1100: 846: 638: 423: 1830: 6838: 6831: 6390: 5318: 3899: 2097: 1818: 1663: 1320: 598: 139: 17: 985: 6825: 6446: 6340: 1806: 886: 455: 163: 159: 6194: 4159: 204: 6745: 6430: 6382: 3534:

Manni E, Petrosini L (March 2004). "A century of cerebellar somatotopy: a debated representation".

2550: 1891: 1778: 1659: 1608:

The lower trace shows an attempt by a patient with cerebellar disease to reproduce the upper trace.

1428: 1290: 1224: 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: 3949:

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.

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In addition to its direct role in motor control, the cerebellum is necessary for several types of

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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: 1841: 1783: 1432: 1407: 1220: 1128: 1091: 1057: 997:. After emitting collaterals that affect nearby parts of the cortex, their axons travel into the 962: 835: 759: 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)".

2245: 973: 6842: 6627: 6493: 6400: 6349: 6249: 4547: 4154: 3870:. Michael Adamaszek, Mario Manto, Dennis J. L. G. Schutter. Cham, Switzerland: Springer. 2022. 2545: 1990: 1915:

motor system inducing a conscious motor avoidance of pain, graded according to pain intensity.

1826: 1694: 1444: 1374: 1299: 1189: 1184: 1170: 1064:. These enlargements are sites of excitatory input from mossy fibers and inhibitory input from 998: 869: 583: 477: 415: 400: 236: 224: 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. 4514: 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

6910: 6858: 6811: 6784: 6576: 6540: 6199: 4172: 2237: 2216: 1822: 1556: 1548: 1511: 1382: 705: 578: 534: 439: 5862:"Precise control of movement kinematics by optogenetic inhibition of Purkinje cell activity" 4539: 1666:

scan can be used to obtain a detailed picture of any structural alterations that may exist.

6488: 6137: 5994: 5742: 5647: 5484: 4415: 4105: 3966: 3237:"The excitatory synaptic action of climbing fibres on the Purkinje cells of the cerebellum" 3072: 2815: 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" 2241: 730:(forming cortico-ponto-cerebellar pathways), and sends output mainly to the ventrolateral 8: 6905: 6821: 6794: 6735: 6585: 6562: 6535: 6423: 5162: 3818:"The organization of the human cerebellum estimated by intrinsic functional connectivity" 2606:, Abingdon, Oxon ; New York, NY : Routledge, 2018.: Routledge, pp. 53–62, 2081: 2073: 2068:

goes to the dorsal octavolateral nucleus, which also has a cerebellum-like structure. In

1899: 1814: 1622: 1536: 1366: 814: 684:, although it also receives visual and other sensory input. Damage to this region causes 673: 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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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

6647: 6641: 6018: 6006: 5937: 5910: 5886: 5861: 5801: 5611: 5586: 5539: 5496: 5441: 5416: 5392: 5367: 5135: 5116: 5100: 5081: 4937: 4898: 4850: 4823: 4795: 4717: 4692: 4634: 4538:

Schmitz TJ (2007). "Examination of Coordination". In O'Sullivan SB, Schmitz TJ (eds.).

4484: 4441: 4428: 4403: 4384: 4341: 4257: 4214: 4117: 4060: 3975: 3950: 3912: 3893: 3842: 3817: 3711: 3684: 3665: 3594: 3559: 3516: 3447: 3422: 3390: 3365: 3343: 3261: 3236: 3212: 3187: 3168: 3084: 3041: 2992: 2924: 2779: 2752: 2485: 2261: 1982: 1879: 1849: 1767: 1759: 1642: 1541: 1349: 1209: 1201: 1030: 739: 610: 582:(tree of life) because of its branched, tree-like appearance in cross-section—are four 333: 6053:

Discoveries in the human brain. Neuroscience prehistory, brain structure, and function

5828: 5754: 5670: 5635: 5383: 4011: 3653: 3156: 6706: 6659: 6602: 6521: 6277: 6010: 5965: 5942: 5891: 5842: 5832: 5793: 5758: 5715: 5711: 5675: 5616: 5565: 5531: 5488: 5446: 5397: 5140: 5120: 5073: 5038: 5000: 4942: 4890: 4855: 4819: 4800: 4722: 4708: 4613: 4576: 4551: 4540: 4476: 4472: 4433: 4376: 4333: 4298: 4249: 4245: 4206: 4109: 4038: 4015: 3980: 3931: 3881: 3871: 3847: 3798: 3757: 3716: 3657: 3617: 3590: 3551: 3508: 3452: 3395: 3335: 3266: 3217: 3160: 3125: 3076: 3033: 2984: 2947: 2928: 2849: 2831: 2784: 2731: 2651: 2626: 2607: 2578: 2559: 2511: 2489: 2477: 2404: 2363: 2326: 2286: 2257: 1903: 1895: 1690: 1626: 1616: 1594:

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

1560: 1269:"fractured somatotopy". A clearer indication of compartmentalization is obtained by 1083:

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.

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Connecting the cerebellum to different parts of the nervous system are three paired

6498: 6145: 6002: 5932: 5922: 5881: 5877: 5873: 5824: 5785: 5750: 5707: 5665: 5655: 5606: 5598: 5523: 5514:

Woodhams PL (July 1977). "The ultrastructure of a cerebellar analogue in octopus".

5480: 5436: 5428: 5387: 5379: 5292: 5130: 5112: 5065: 5030: 4932: 4902: 4882: 4845: 4835: 4790: 4782: 4712: 4704: 4605: 4468: 4423: 4368: 4345: 4325: 4288: 4241: 4218: 4198: 4164: 4121: 4101: 4007: 3970: 3962: 3923: 3837: 3829: 3788: 3747: 3706: 3696: 3649: 3586: 3543: 3500: 3442: 3438: 3434: 3385: 3377: 3331: 3327: 3300: 3256: 3252: 3248: 3207: 3199: 3172: 3152: 3115: 3088: 3068: 3023: 2976: 2914: 2839: 2823: 2774: 2764: 2555: 2469: 2435: 2394: 2355: 2322: 2318: 2294: 2276: 2145:

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

1646: 1013: 712: 685: 677: 502: 473: 465: 380: 372: 353: 349: 79: 6204: 4232:

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 6652: 6635: 6597: 6557: 6188: 6163: 4840: 3120: 3103: 2439: 1944: 1936: 1791: 1787: 1735: 1727: 1523: 1496: 1378: 1197: 1193: 990: 782:

fiber inputs each carry fiber-specific information; the cerebellum also receives

735: 716: 654: 606: 530: 376: 217: 192: 6711: 4917: 4662:"Veterans of Iraq, Afghanistan Show Brain Changes Related to Explosion Exposure" 4609: 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 6755: 6750: 6530: 6367: 6241: 5640:

Proceedings of the National Academy of Sciences of the United States of America

4770: 4635:"NINDS Ataxias and Cerebellar or Spinocerebellar Degeneration Information Page" 3793: 3776: 2827: 2423: 2085: 2069: 2057: 2053: 2049: 2045: 2041: 2033: 1751: 1678: 1483: 1479: 1475: 1436: 1286: 1279: 1228: 1149: 1115: 928: 891: 879: 791: 775: 771: 727: 709: 697: 634: 614: 435: 431: 427: 411: 361: 337: 6169: 6150: 6123: 5789: 5432: 4372: 4168: 3885: 3752: 3735: 3701: 3304: 3028: 3011: 2980: 2573:

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

526: 461: 392: 313: 241: 6415: 5660: 3188:"PCP4: a regulator of aldosterone synthesis in human adrenocortical tissues" 2919: 2902: 2281: 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

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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

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enter the granular layer from their points of origin, many arising from the

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A Latin dictionary founded on Andrews' edition of Freund's Latin dictionary

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Ghez C, Fahn S (1985). "The cerebellum". In Kandel ER, Schwartz JH (eds.).

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in the adult brain, initiating the organization of new cerebellar lobules.

689: 569: 542: 529:, which also includes the pons; the metencephalon is the upper part of the 469: 434:

of the granule cells; the other is an extremely strong input from a single

404: 5846: 5527: 5042: 5004: 4786: 4617: 4480: 4329: 4210: 3833: 3761: 3456: 3366:"Peripheral nerve injury induces adult brain neurogenesis and remodelling" 3270: 2298: 1025:

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

5535: 4253: 2473: 2138: 2012: 1731: 1528: 1303: 1285:

It is not only receptive fields that define the microzone structure: The

1270: 958: 911: 831: 810: 802: 743: 701: 557: 341: 3816:

Buckner RL, Krienen FM, Castellanos A, Diaz JC, Yeo BT (November 2011).

3203: 2244:). The updated content was reintegrated into the Knowledge page under a 1931: 1726:: it is about 15 years younger than expected in a centenarian. Further, 1086: 6725: 5636:"Reciprocal evolution of the cerebellum and neocortex in fossil humans" 5069: 4277:"Neural substrates of eyeblink conditioning: acquisition and retention" 4202: 3381: 3059:

Strick PL, Dum RP, Fiez JA (2009). "Cerebellum and nonmotor function".

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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:

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The smallest region, the flocculonodular lobe, is often called the

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are in front of the cerebellum. It is separated from the overlying

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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".

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Ivry RB, Spencer RM, Zelaznik HN, Diedrichsen J (December 2002).

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Standring S, Borley NR, et al., eds. (2008). "Chapter 20".

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provide a teaching signal that induces synaptic modification in

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Raz N, Gunning-Dixon F, Head D, Williamson A, Acker JD (2001).

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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".

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with a highly regular arrangement, the most important being

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Synonymia anatomica. Synonymik der anatomischen Nomenclatur

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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"

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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

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Further significant developments did not come until the

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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

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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

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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

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to be confined to the cerebellum. In particular, the

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as a neurotransmitter and project exclusively to the

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Clarke E, O'Malley CD (1996). "Ch. 11: Cerebellum".

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Albus JS (1971). "A theory of cerebellar function".

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1971 that a cerebellar Purkinje cell functions as a

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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: 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: 703: 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: 258: 255: 252: 250: 246: 243: 240: 238: 234: 231: 228: 226: 222: 219: 216: 213: 209: 206: 203: 201: 197: 194: 191: 189: 185: 182: 179: 177: 173: 168: 165: 161: 158: 156: 152: 149: 145: 141: 138: 136: 132: 129: 126: 122: 117: 76: 74:Pronunciation 72: 67: 60: 55: 48: 43: 38: 33: 19: 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: 6087: 6081: 6072: 6052: 6046: 6037: 6031: 5990: 5986: 5980: 5961: 5955: 5918: 5914: 5904: 5869: 5865: 5855: 5820: 5814: 5781: 5777: 5771: 5738: 5734: 5728: 5703: 5699: 5643: 5639: 5629: 5594: 5590: 5580: 5561: 5519: 5515: 5509: 5476: 5472: 5424: 5420: 5410: 5378:(1): 14–27. 5375: 5371: 5361: 5349:. Retrieved 5345:the original 5335: 5323:. Retrieved 5319:the original 5309: 5297:. Retrieved 5293:the original 5283: 5271:. Retrieved 5267:the original 5257: 5245:. Retrieved 5231: 5219:. Retrieved 5215:the original 5205: 5193:. Retrieved 5189:the original 5179: 5167:. Retrieved 5163:the original 5153: 5108: 5104: 5094: 5061: 5057: 5051: 5026: 5022: 4996: 4992: 4986: 4976: 4928: 4924: 4911: 4878: 4874: 4868: 4831: 4828:PLOS Biology 4827: 4813: 4778: 4774: 4764: 4752:. Retrieved 4748:the original 4700: 4696: 4686: 4674:. Retrieved 4665: 4655: 4643:. Retrieved 4601: 4597: 4591: 4572: 4566: 4541: 4533: 4522:. Retrieved 4515:the original 4510: 4497: 4464: 4461:Neuroscience 4460: 4454: 4411: 4407: 4397: 4364: 4360: 4354: 4321: 4317: 4311: 4284: 4280: 4270: 4237: 4233: 4227: 4194: 4190: 4184: 4173:the original 4150: 4146: 4097: 4093: 4076: 4072: 4061: 4053: 4034: 4028: 4003: 3999: 3993: 3958: 3954: 3944: 3919: 3915: 3908: 3866: 3860: 3825: 3821: 3811: 3784: 3780: 3770: 3743: 3739: 3729: 3692: 3688: 3678: 3648:(6): 732–9. 3645: 3641: 3613: 3607: 3582: 3578: 3572: 3542:(3): 241–9. 3539: 3535: 3529: 3496: 3492: 3430: 3426: 3373: 3369: 3323: 3319: 3313: 3296: 3292: 3244: 3240: 3230: 3195: 3191: 3181: 3148: 3144: 3138: 3111: 3107: 3097: 3064: 3060: 3054: 3019: 3015: 3005: 2972: 2968: 2962: 2943: 2937: 2910: 2906: 2896: 2887: 2811: 2807: 2797: 2760: 2756: 2746: 2727: 2723: 2647: 2641: 2617:, retrieved 2603: 2593: 2575:Neuroscience 2574: 2568: 2541: 2537: 2508:Neuroscience 2507: 2465: 2461: 2431: 2427: 2417: 2390: 2386: 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 18:Cerebellar 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: 5678: 5671:553338 5668: 5619: 5609: 5568: 5542: 5536:864041 5534: 5499: 5491: 5449: 5439: 5400: 5390: 5143: 5133: 5123: 5084: 5076: 5041: 5003: 4945: 4935: 4901: 4893: 4858: 4848: 4803: 4793: 4725: 4715: 4616: 4579: 4554: 4550:–225. 4487: 4479: 4444: 4436: 4387: 4379: 4344: 4336: 4301: 4260: 4254:194656 4252: 4217: 4209: 4157: 4120: 4112: 4041: 4018: 3983: 3973: 3934: 3884: 3874: 3850: 3840: 3801: 3760: 3719: 3709: 3668: 3660: 3620: 3597: 3562: 3554: 3519: 3511: 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:. 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