The cerebellum is a major feature of the hindbrain of all vertebrates. Although usually smaller than the cerebrum, in some animals such as the mormyrid fishes it may be as large as or even larger. In humans, the cerebellum plays an important role in motor control. It may also be involved in some cognitive functions such as attention and language as well as emotional control 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 coordination, precision, and accurate timing: it receives input from sensory systems of the spinal cord and from other parts of the brain, and integrates these inputs to fine-tune motor activity. Cerebellar damage produces disorders in fine movement, equilibrium, posture, and motor learning in humans.
In neuroanatomy, a nucleus is a cluster of neurons in the central nervous system, located deep within the cerebral hemispheres and brainstem. The neurons in one nucleus usually have roughly similar connections and functions. Nuclei are connected to other nuclei by tracts, the bundles (fascicles) of axons extending from the cell bodies. A nucleus is one of the two most common forms of nerve cell organization, the other being layered structures such as the cerebral cortex or cerebellar cortex. In anatomical sections, a nucleus shows up as a region of gray matter, often bordered by white matter. The vertebrate brain contains hundreds of distinguishable nuclei, varying widely in shape and size. A nucleus may itself have a complex internal structure, with multiple types of neurons arranged in clumps (subnuclei) or layers.
Eyeblink conditioning (EBC) is a form of classical conditioning that has been used extensively to study neural structures and mechanisms that underlie learning and memory. The procedure is relatively simple and usually consists of pairing an auditory or visual stimulus with an eyeblink-eliciting unconditioned stimulus (US). Naïve organisms initially produce a reflexive, unconditioned response (UR) that follows US onset. After many CS-US pairings, an association is formed such that a learned blink, or conditioned response (CR), occurs and precedes US onset. The magnitude of learning is generally gauged by the percentage of all paired CS-US trials that result in a CR. Under optimal conditions, well-trained animals produce a high percentage of CRs. The conditions necessary for, and the physiological mechanisms that govern, eyeblink CR learning have been studied across many mammalian species, including mice, rats, guinea pigs, rabbits, ferrets, cats, and humans. Historically, rabbits have been the most popular research subjects.
The inferior olivary nucleus (ION), is a structure found in the medulla oblongata underneath the superior olivary nucleus. In vertebrates, the ION is known to coordinate signals from the spinal cord to the cerebellum to regulate motor coordination and learning. These connections have been shown to be tightly associated, as degeneration of either the cerebellum or the ION results in degeneration of the other.
The cerebellar vermis is located in the medial, cortico-nuclear zone of the cerebellum, which is in the posterior fossa of the cranium. The primary fissure in the vermis curves ventrolaterally to the superior surface of the cerebellum, dividing it into anterior and posterior lobes. Functionally, the vermis is associated with bodily posture and locomotion. The vermis is included within the spinocerebellum and receives somatic sensory input from the head and proximal body parts via ascending spinal pathways.
The upper part of the posterior district of the medulla oblongata is occupied by the inferior cerebellar peduncle, a thick rope-like strand situated between the lower part of the fourth ventricle and the roots of the glossopharyngeal and vagus nerves.
The dentate nucleus is a cluster of neurons, or nerve cells, in the central nervous system that has a dentate – tooth-like or serrated – edge. It is located within the deep white matter of each cerebellar hemisphere, and it is the largest single structure linking the cerebellum to the rest of the brain. It is the largest and most lateral, or farthest from the midline, of the four pairs of deep cerebellar nuclei, the others being the globose and emboliform nuclei, which together are referred to as the interposed nucleus, and the fastigial nucleus. The dentate nucleus is responsible for the planning, initiation and control of voluntary movements. The dorsal region of the dentate nucleus contains output channels involved in motor function, which is the movement of skeletal muscle, while the ventral region contains output channels involved in nonmotor function, such as conscious thought and visuospatial function.
The globose nucleus is one of the deep cerebellar nuclei. It is located medial to the emboliform nucleus and lateral to the fastigial nucleus. This nucleus contains primarily large and small multipolar neurons.
The interposed nucleus is part of the deep cerebellar complex and is composed of the globose nucleus and the emboliform nucleus. It is located in the roof of the fourth ventricle, lateral to the fastigial nucleus. It receives its afferent supply from the anterior lobe of the cerebellum and sends output via the superior cerebellar peduncle to the red nucleus.
The fastigial nucleus is located in the cerebellum. It is one of the four deep cerebellar nuclei, and is grey matter embedded in the white matter of the cerebellum.
The flocculus is a small lobe of the cerebellum at the posterior border of the middle cerebellar peduncle anterior to the biventer lobule. Like other parts of the cerebellum, the flocculus is involved in motor control. It is an essential part of the vestibulo-ocular reflex, and aids in the learning of basic motor skills in the brain.
Cerebellar peduncles connect the cerebellum to the brain stem. There are six cerebellar peduncles in total, three on each side:
In the human brain, the superior cerebellar peduncle is a paired structure of white matter that connects the cerebellum to the midbrain. It consists mainly of efferent fibers, the cerebellothalamic tract that runs from a cerebellar hemisphere to the contralateral thalamus, and the cerebellorubral tract that runs from a cerebellar hemisphere to the red nucleus. It also contains afferent tracts, most prominent of which is the ventral spinocerebellar tract. Other afferent tracts are the trigeminothalamic fibers, tectocerebellar fibers, and noradrenergic fibers from the locus coeruleus. The superior peduncle emerges from the upper and medial parts of the white matter of each hemisphere and is placed under cover of the upper part of the cerebellum.
Mossy fibers are one of the major inputs to cerebellum. There are many sources of this pathway, the largest of which is the cerebral cortex, which sends input to the cerebellum via the pontocerebellar pathway. Other contributors include the vestibular nerve and nuclei, the spinal cord, the reticular formation, and feedback from deep cerebellar nuclei. Axons enter the cerebellum via the middle and inferior cerebellar peduncles, where some branch to make contact with deep cerebellar nuclei. They ascend into the white matter of the cerebellum, where each axon branches to innervate granule cells in several cerebellar folia.
The emboliform nucleus is a deep cerebellar nucleus that lies immediately to the medial side of the nucleus dentatus, and partly covering its hilum. It is one among the four pairs of deep cerebellar nuclei, which are from lateral to medial: the dentate, interposed, and fastigial nuclei. These nuclei can be seen using Weigert's elastic stain.
The anatomy of the cerebellum can be viewed at three levels. At the level of gross anatomy, the cerebellum consists of a tightly folded and crumpled layer of cortex, with white matter underneath, several deep nuclei embedded in the white matter, and a fluid-filled ventricle in the middle. At the intermediate level, the cerebellum and its auxiliary structures can be broken down into several hundred or thousand independently functioning modules or compartments known as microzones. At the microscopic level, each module consists of the same small set of neuronal elements, laid out with a highly stereotyped geometry.
The name granule cell has been used for a number of different types of neurons whose only common feature is that they all have very small cell bodies. Granule cells are found within the granular layer of the cerebellum, the dentate gyrus of the hippocampus, the superficial layer of the dorsal cochlear nucleus, the olfactory bulb, and the cerebral cortex.
Cerebellar cognitive affective syndrome (CCAS), also called Schmahmann's syndrome is a condition that follows from lesions (damage) to the cerebellum of the brain. It refers to a constellation of deficits in the cognitive domains of executive function, spatial cognition, language, and affect resulting from damage to the cerebellum. Impairments of executive function include problems with planning, set-shifting, abstract reasoning, verbal fluency, and working memory, and there is often perseveration, distractibility and inattention. Language problems include dysprosodia, agrammatism and mild anomia. Deficits in spatial cognition produce visual–spatial disorganization and impaired visual–spatial memory. Personality changes manifest as blunting of affect or disinhibited and inappropriate behavior. These cognitive impairments result in an overall lowering of intellectual function. CCAS challenges the traditional view of the cerebellum being responsible solely for regulation of motor functions. It is now thought that the cerebellum is responsible for monitoring both motor and nonmotor functions. The nonmotor deficits described in CCAS are believed to be caused by dysfunction in cerebellar connections to the cerebral cortex and limbic system.
Unipolar brush cells (UBCs) are a class of excitatory glutamatergic interneuron found in the granular layer of the cerebellar cortex and also in the granule cell domain of the cochlear nucleus.
