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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 56 57 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 14 this complex of motor symptoms is called ataxia 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 5 if cerebellar dysfunction is indicated a magnetic resonance imaging scan can be used to obtain a detailed picture of any structural alterations that may exist 58 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 59 60 some forms of migraine headache may also produce temporary dysfunction of the cerebellum of variable severity 61 infection can result in cerebellar damage in such conditions as the prion diseases 62 and miller fisher syndrome a variant of guillain barré syndrome aging edit 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 according to an epigenetic biomarker of tissue age known as an epigenetic clock it is about 15 years younger than expected in a centenarian 63 further gene expression patterns in the human cerebellum show less age related alteration than that in the cerebral cortex 64 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 65 66 developmental and degenerative disorders edit ultrasound image of the fetal head at 19 weeks of pregnancy in a modified axial section showing the normal fetal cerebellum and cisterna magna 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 67 visualization of the fetal cerebellum by ultrasound scan at 18 to 20 weeks of pregnancy can be used to screen for fetal neural tube defects with a sensitivity rate of up to 99 68 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 69 as cgnps terminally differentiate into cerebellar granule cells 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 69 mutations that abnormally activate sonic hedgehog signaling predispose to cancer of the cerebellum medulloblastoma in humans with gorlin syndrome and in genetically engineered mouse models 70 71 congenital malformation or underdevelopment hypoplasia of the cerebellar vermis is a characteristic of both dandy walker syndrome and joubert syndrome 72 73 in very rare cases the entire cerebellum may be absent 74 the inherited neurological disorders machado joseph disease ataxia telangiectasia and friedreich s ataxia cause progressive neurodegeneration linked to cerebellar loss 59 67 congenital brain malformations outside the cerebellum can in turn cause herniation of cerebellar tissue as seen in some forms of arnold chiari malformation 75 other conditions that are closely linked to cerebellar degeneration include the idiopathic progressive neurological disorders multiple system atrophy and ramsay hunt syndrome type i 76 77 and paraneoplastic cerebellar degeneration an autoimmune disorder in which tumors elsewhere in the body elicit an autoimmune response that causes neuronal loss in the cerebellum 78 cerebellar atrophy can result from chronic thiamine vitamin b1 deficiency as seen in dry beriberi and in wernicke korsakoff syndrome 79 or from vitamin e deficiency 67 cerebellar atrophy has been observed in many other neurological disorders including huntington s disease multiple sclerosis 62 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 67 pain edit there is a general consensus that the cerebellum is involved in pain processing 80 81 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 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 26 the magnitude of this remodeling and the induction of neuron progenitor markers suggest the contribution of adult neurogenesis to these changes comparative anatomy and evolution edit cross section of the brain of a porbeagle shark with the cerebellum highlighted in blue the circuits in the cerebellum are similar across all classes of vertebrates including fish reptiles birds and mammals 82 there is also an analogous brain structure in cephalopods with well developed brains such as octopuses 83 this has been taken as evidence that the cerebellum performs functions important to all animal species with a brain 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 84 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 84 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 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 85 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 86 the size of the human cerebellum compared to the rest of the brain has been increasing in size while the cerebrum decreased in size 87 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 88 the lateral hemispheres of the cerebellum are now 2 7 times greater in both humans and apes than they are in monkeys 87 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 hominidae evolution due to the cerebellum s role in cognitive functions the increase in its size may have played a role in cognitive expansion 87 cerebellum like structures edit 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 82 the only cerebellum like structure found in mammals is the dorsal cochlear nucleus dcn one of the two primary sensory nuclei that receive input directly from the auditory nerve 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 89 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 82 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 82 in fact james m 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 90 despite bower s viewpoint there is also strong evidence that the cerebellum directly influences motor output in mammals 91 92 history edit base of the human brain as drawn by andreas vesalius in 1543 descriptions edit even the earliest anatomists were able to recognize the cerebellum by its distinctive appearance aristotle and herophilus quoted in galen called it the παρεγκεφαλίς parenkephalis as opposed to the ἐγκέφαλος enkephalos or brain proper galen s extensive description is the earliest that survives he speculated that the cerebellum was the source of motor nerves 93 further significant developments did not come until the renaissance vesalius discussed the cerebellum briefly and the anatomy was described more thoroughly by thomas willis 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 luigi rolando in 1809 established the key finding that damage to the cerebellum results in motor disturbances jean pierre flourens 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 the lesion is very extensive 94 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 5 etymology edit the name cerebellum is a diminutive of cerebrum brain 95 it can be translated literally as little brain the latin name is a direct translation of the ancient greek παρεγκεφαλίς parenkephalis which was used in the works of aristotle the first known writer to describe the structure 96 no other name is used in the english language literature but historically a variety of greek or latin derived names have been used including cerebrum parvum 97 encephalion 98 encranion 97 cerebrum posterius 99 and parencephalis 97 references edit this article was submitted to wikijournal of medicine for external academic peer review in 2016 reviewer reports the updated content was reintegrated into the wikipedia page under a cc by sa 3 0 license 2016 the version of record as reviewed is marion wright skaggs william nielsen finn et al 2016 the cerebellum pdf wikijournal of medicine 3 1 doi 10 15347 wjm 2016 001 wikidata q44001486 hodos w 2009 evolution of ce...