Climbing fiber | |
---|---|
Details | |
Location | Inferior olive and Cerebellum [ citation needed ] |
Shape | Unique projection neuron (see text) |
Function | Unique excitatory function (see text) |
Neurotransmitter | Glutamate |
Presynaptic connections | Inferior olive |
Postsynaptic connections | Purkinje cells |
Anatomical terms of neuroanatomy |
Climbing fibers are the name given to a series of neuronal projections from the inferior olivary nucleus located in the medulla oblongata. [1] [2]
These axons pass through the pons and enter the cerebellum via the inferior cerebellar peduncle where they form synapses with the deep cerebellar nuclei and Purkinje cells. Each climbing fiber will form synapses with 1-10 Purkinje cells.
Early in development, Purkinje cells are innervated by multiple climbing fibers, but as the cerebellum matures, these inputs gradually become eliminated resulting in a single climbing fiber input per Purkinje cell.
These fibers provide very powerful, excitatory input to the cerebellum which results in the generation of complex spike excitatory postsynaptic potential (EPSP) in Purkinje cells. [1] In this way climbing fibers (CFs) perform a central role in motor behaviors. [3]
The climbing fibers carry information from various sources such as the spinal cord, vestibular system, red nucleus, superior colliculus, reticular formation and sensory and motor cortices. Climbing fiber activation is thought to serve as a motor error signal sent to the cerebellum, and is an important signal for motor timing. In addition to the control and coordination of movements, [4] the climbing fiber afferent system contributes to sensory processing and cognitive tasks likely by encoding the timing of sensory input independently of attention or awareness. [5] [6] [7]
In the central nervous system, these fibers are able to undergo remarkable regenerative modifications in response to injuries, being able to generate new branches by sprouting to innervate surrounding Purkinje cells if these lose their CF innervation. [8] This kind of injury-induced sprouting has been shown to need the growth associated protein GAP-43. [9] [10] [11]
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 it 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 neurophysiology, long-term depression (LTD) is an activity-dependent reduction in the efficacy of neuronal synapses lasting hours or longer following a long patterned stimulus. LTD occurs in many areas of the CNS with varying mechanisms depending upon brain region and developmental progress.
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.
In neuroanatomy, a neural pathway is the connection formed by axons that project from neurons to make synapses onto neurons in another location, to enable neurotransmission. Neurons are connected by a single axon, or by a bundle of axons known as a nerve tract, or fasciculus. Shorter neural pathways are found within grey matter in the brain, whereas longer projections, made up of myelinated axons, constitute white matter.
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.
Basket cells are inhibitory GABAergic interneurons of the brain, found throughout different regions of the cortex and cerebellum.
Purkinje cells or Purkinje neurons, named for Czech physiologist Jan Evangelista Purkyně who identified them in 1837, are a unique type of prominent large neurons located in the cerebellar cortex of the brain. With their flask-shaped cell bodies, many branching dendrites, and a single long axon, these cells are essential for controlling motor activity. Purkinje cells mainly release GABA neurotransmitter, which inhibits some neurons to reduce nerve impulse transmission. Purkinje cells efficiently control and coordinate the body's motor motions through these inhibitory actions.
The spinocerebellar tract is a nerve tract originating in the spinal cord and terminating in the same side (ipsilateral) of the cerebellum.
There are four deep cerebellar nuclei embedded in the white matter of the medullary centre. The nuclei are the fastigial, globose, emboliform, and dentate nuclei.
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.
In neuroscience, Golgi cells are the most abundant inhibitory interneurons found within the granular layer of the cerebellum. Golgi cells can be found in the granular layer at various layers. The Golgi cell is essential for controlling the activity of the granular layer. They were first identified as inhibitory in 1964. It was also the first example of an inhibitory feedback network in which the inhibitory interneuron was identified anatomically. Golgi cells produce a wide lateral inhibition that reaches beyond the afferent synaptic field and inhibit granule cells via feedforward and feedback inhibitory loops. These cells synapse onto the dendrite of granule cells and unipolar brush cells. They receive excitatory input from mossy fibres, also synapsing on granule cells, and parallel fibers, which are long granule cell axons. Thereby this circuitry allows for feed-forward and feed-back inhibition of granule cells.
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.
The cochlear nuclear (CN) complex comprises two cranial nerve nuclei in the human brainstem, the ventral cochlear nucleus (VCN) and the dorsal cochlear nucleus (DCN). The ventral cochlear nucleus is unlayered whereas the dorsal cochlear nucleus is layered. Auditory nerve fibers, fibers that travel through the auditory nerve carry information from the inner ear, the cochlea, on the same side of the head, to the nerve root in the ventral cochlear nucleus. At the nerve root the fibers branch to innervate the ventral cochlear nucleus and the deep layer of the dorsal cochlear nucleus. All acoustic information thus enters the brain through the cochlear nuclei, where the processing of acoustic information begins. The outputs from the cochlear nuclei are received in higher regions of the auditory brainstem.
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.
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Cerebellar granule cells form the thick granular layer of the cerebellar cortex and are among the smallest neurons in the brain. Cerebellar granule cells are also the most numerous neurons in the brain: in humans, estimates of their total number average around 50 billion, which means that they constitute about 3/4 of the brain's neurons.
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.
Tom Otis is an American researcher, academic and author. He is the Chief Scientific Officer at the Sainsbury Wellcome Centre for Neural Circuits and Behaviour and holds a Professorship in Neuroscience at University College London.
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