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Damage to this region causes disturbances of balance and gait. 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 proprioceptive input from the dorsal columns of the spinal cord including the spinocerebellar tract and from the cranial trigeminal nerve , as well as from visual and auditory systems.
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 parietal lobe via the pontine nuclei forming cortico-ponto-cerebellar pathways , and sends output mainly to the ventrolateral thalamus in turn connected to motor areas of the premotor cortex and primary motor area of the cerebral cortex and to the red nucleus.
Two types of neuron play dominant roles in the cerebellar circuit: Purkinje cells and granule cells. Three types of axons also play dominant roles: mossy fibers and climbing fibers which enter the cerebellum from outside , and parallel fibers 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.
Climbing fibers project to Purkinje cells and also send collaterals directly to the deep nuclei. The cerebellar cortex is divided into three layers.
At the bottom lies the thick granular layer, densely packed with granule cells, along with interneurons , mainly Golgi cells but also including Lugaro cells and unipolar brush cells.
In the middle lies the Purkinje layer, a narrow zone that contains the cell bodies of Purkinje cells and Bergmann glial cells.
At the top lies the molecular layer, which contains the flattened dendritic 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: stellate cells and basket cells.
The top, outermost layer of the cerebellar cortex is the molecular layer. This layer contains the flattened dendritic trees 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: stellate cells and basket cells. 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. 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 , After emitting collaterals that affect nearby parts of the cortex, their axons travel into the deep cerebellar nuclei , where they make on the order of 1, contacts each with several types of nuclear cells, all within a small domain.
Purkinje cells use GABA as their neurotransmitter, and therefore exert inhibitory effects on their targets. 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 extracellular recording techniques.
Purkinje cells normally emit action potentials at a high rate even in the absence of the synaptic input. The spike trains show a mixture of what are called simple and complex spikes.
Complex spikes are often followed by a pause of several hundred milliseconds during which simple spike activity is suppressed. A specific, recognizable feature of Purkinje neurons is the expression of calbindin.
Cerebellar granule cells , in contrast to Purkinje cells, are among the smallest neurons in the brain. A granule cell emits only four to five dendrites, each of which ends in an enlargement called a dendritic claw.
The thin, unmyelinated 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 parallel fiber ; the splitting of the vertical branch into two horizontal branches gives rise to a distinctive "T" shape.
Granule cells receive all of their input from mossy fibers, but outnumber them by 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 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 by David Marr , 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.
Mossy fibers enter the granular layer from their points of origin, many arising from the pontine nuclei , others from the spinal cord, vestibular nuclei etc.
In the human cerebellum, the total number of mossy fibers has been estimated at about million. Within the granular layer, a mossy fiber generates a series of enlargements called rosettes.
The contacts between mossy fibers and granule cell dendrites take place within structures called glomeruli. Each glomerulus has a mossy fiber rosette at its center, and up to 20 granule cell dendritic claws contacting it.
Terminals from Golgi cells infiltrate the structure and make inhibitory synapses onto the granule cell dendrites. The entire assemblage is surrounded by a sheath of glial cells.
Purkinje cells also receive input from the inferior olivary nucleus on the contralateral side of the brainstem via climbing fibers.
Although the inferior olive lies in the medulla oblongata 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 a single Purkinje cell.
The climbing fiber synapses cover the cell body and proximal dendrites; this zone is devoid of parallel fiber inputs.
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 , of one type versus exactly one of the 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. The deep nuclei 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 nuclei 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 dentate , globose , emboliform , and fastigial 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 interposed nucleus. 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.
These cells project to a variety of targets outside the cerebellum. Intermixed with them are a lesser number of small cells, which use GABA as a neurotransmitter and project exclusively to the inferior olivary nucleus , the source of climbing fibers.
Thus, the nucleo-olivary projection provides an inhibitory feedback 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.
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 Purkinje cells again in cats. 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 compartmentalized. There are large compartments that are generally known as zones ; these can be divided into smaller compartments known as microzones.
The first indications of compartmental structure came from studies of the receptive fields of cells in various parts of the cerebellar cortex.
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 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.
Oscarsson in the late s proposed that these cortical zones can be partitioned into smaller units called microzones. Microzones were found to contain on the order of Purkinje cells each, arranged in a long, narrow strip, oriented perpendicular to the cortical folds.
It is not only receptive fields that define the microzone structure: The climbing fiber input from the inferior olivary nucleus 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 gap junctions , which synchronize their activity, causing Purkinje cells within a microzone to show correlated complex spike activity on a millisecond time scale.
In , 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, plus a group of coupled olivary neurons that project to all of the included microzones as well as to the deep nuclear area.
The cerebellum is provided with blood from three paired major arteries: the superior cerebellar artery SCA , the anterior inferior cerebellar artery AICA , and the posterior inferior cerebellar artery 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 anastomose 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 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.
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 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.
Prior to the s the function of the cerebellum was almost universally believed to be purely motor-related, but newer findings have brought that view into question.
Functional imaging 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 to be confined to the cerebellum.
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 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 and behavior.
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, The Cerebellum as a Neuronal Machine by John C.
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 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 subsequent cerebellar-learning models, however, have followed Albus in assuming that climbing fiber activity would be an error signal, and would cause synchronously activated parallel fiber inputs to be weakened.
Some of these later models, such as the Adaptive Filter model of Fujita  made attempts to understand cerebellar function in terms of optimal control theory.
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.
Studies of the vestibulo—ocular reflex 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.
One of the most extensively studied cerebellar learning tasks is the eyeblink conditioning 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.
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.
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 "learning theories" and "performance theories"—that is, theories that make use of synaptic plasticity 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 mathematical models and simulated using computers. Perhaps the earliest "performance" theory was the "delay line" hypothesis of Valentino Braitenberg.
The original theory put forth by Braitenberg and Roger Atwood in proposed that slow propagation of signals along parallel fibers imposes predictable delays that allow the cerebellum to detect time relationships within a certain window.
Theories in the "learning" category almost all derive from publications by Marr and Albus. Marr's 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 also formulated his version as a software algorithm he called a CMAC Cerebellar Model Articulation Controller , which has been tested in a number of applications.
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 flocculonodular lobe 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.
Other manifestations include hypotonia decreased muscle tone , dysarthria problems with speech articulation , dysmetria problems judging distances or ranges of movement , dysdiadochokinesia inability to perform rapid alternating movements such as walking , impaired check reflex or rebound phenomenon, and intention tremor involuntary movement caused by alternating contractions of opposing muscle groups.
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.
To identify cerebellar problems, neurological examination includes assessment of gait a broad-based gait being indicative of ataxia , finger-pointing tests and assessment of posture.
The list of medical problems that can produce cerebellar damage is long, including stroke , hemorrhage , swelling of the brain cerebral edema , tumors , alcoholism , physical trauma such as gunshot wounds or explosives, and chronic degenerative conditions such as olivopontocerebellar atrophy.
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 epigenetic clock : it is about 15 years younger than expected in a centenarian.
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.
In normal development, endogenous sonic hedgehog 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 in the mouse.
Congenital malformation or underdevelopment hypoplasia of the cerebellar vermis is a characteristic of both Dandy—Walker syndrome and Joubert syndrome.
Other conditions that are closely linked to cerebellar degeneration include the idiopathic progressive neurological disorders multiple system atrophy and Ramsay Hunt syndrome type I ,   and the autoimmune disorder paraneoplastic cerebellar degeneration , in which tumors elsewhere in the body elicit an autoimmune response that causes neuronal loss in the cerebellum.
Cerebellar atrophy has been observed in many other neurological disorders including Huntington's disease , multiple sclerosis ,  essential tremor , progressive myoclonus epilepsy , and Niemann—Pick disease.
Cerebellar atrophy can also occur as a result of exposure to toxins including heavy metals or pharmaceutical or recreational drugs. There is a general consensus that the cerebellum is involved in pain processing.
Some of this information is transferred to the motor system inducing a conscious motor avoidance of pain, graded according to pain intensity. 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 macroscopic anatomical remodeling of the hindbrain, including the cerebellum. The circuits in the cerebellum are similar across all classes of vertebrates , including fish, reptiles, birds, and mammals.
There is considerable variation in the size and shape of the cerebellum in different vertebrate species. In amphibians , it is little developed, and in lampreys , and hagfish , 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 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.
The cerebellum of cartilaginous and bony fishes 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 deep cerebellar nuclei.
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 mormyrid fish a family of weakly electrosensitive freshwater fish , the cerebellum is considerably larger than the rest of the brain put together.
The largest part of it is a special structure called the valvula , which has an unusually regular architecture and receives much of its input from the electrosensory system.
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, and in Homo sapiens until the middle Pleistocene 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 expanded.
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 Hominidae evolution.
Due to the cerebellum's role in cognitive functions, the increase in its size may have played a role in cognitive expansion.
Most vertebrate species have a cerebellum and one or more cerebellum-like structures, brain areas that resemble the cerebellum in terms of cytoarchitecture and neurochemistry.
The DCN is a layered structure, with the bottom layer containing granule cells similar to those of the cerebellum, giving rise to parallel fibers that rise to the superficial layer and travel across it horizontally.
The superficial layer contains a set of GABAergic neurons called cartwheel cells that resemble Purkinje cells anatomically and chemically—they receive parallel fiber input, but do not have any inputs that resemble climbing fibers.
The output neurons of the DCN are pyramidal cells. 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.
Most species of fish and amphibians possess a lateral line 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 goes to the dorsal octavolateral nucleus, which also has a cerebellum-like structure.
In ray-finned fishes by far the largest group , the optic tectum has a layer—the marginal layer—that is cerebellum-like.
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 modifiable synapses , 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.
Bower 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.
Even the earliest anatomists were able to recognize the cerebellum by its distinctive appearance. It is intriguing to think that there might be analogs of '"fractured somatotopy"' in the cognitive parts of the cerebellum that could help support highly complex, sophisticated cognitive functions, such as language or abstract reasoning, Sereno said.
And that's just how the cerebellum is set up. Until now, the cerebellum was thought to be involved mainly in basic functions like movement, but its expansion over time and its new inputs from cortical areas involved in cognition suggest that it can also process advanced concepts like mathematical equations.
For instance, there is some recent evidence that people who suffer cerebellum damage have difficulty processing emotion.
Sereno began working on imaging while at the University College London, and continued to work on processing the imaging data and analysis at SDSU.
Materials provided by San Diego State University. Original written by Padma Nagappan. Note: Content may be edited for style and length. Science News.
Cerebellum highlighted in illustration of brain stock image. Journal Reference : Martin I. ScienceDaily, 31 July San Diego State University. Retrieved September 2, from www.
Multifunctional Small Brains Nov. This discovery can help understand the consequences The cerebellum, a brain structure traditionally considered to be involved in Researchers found a direct neural connection The analysis of thousands Below are relevant articles that may interest you.
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