The cerebellar vermis (from

Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power ...

''vermis,'' "worm") is located in the medial, cortico-nuclear zone of the

cerebellum The cerebellum (Latin for "little brain") 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 cere ...

, which is in the posterior fossa of the

cranium The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, ...

. The

primary fissure The monticulus of the cerebellum The cerebellum (Latin for "little brain") 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 ...

in the vermis curves

ventrolateral Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...

ly to the

superior Superior may refer to: *Superior (hierarchy), something which is higher in a hierarchical structure of any kind Places *Superior (proposed U.S. state), an unsuccessful proposal for the Upper Peninsula of Michigan to form a separate state *Lake ...

surface of the

, dividing it into

anterior Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...

and posterior

lobes Lobe may refer to: People with the name * Lobe (surname) Science and healthcare * Lobe (anatomy) * Lobe, a large-scale structure of a radio galaxy * Glacial lobe, a lobe-shaped glacier * Lobation, a characteristic of the nucleus of certain ...

. 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 A head is the part of an organism which usually includes the ears, brain, forehead, cheeks, chin, eyes, nose, and mouth, each of which aid in various sensory functions such as sight, hearing, smell, and taste. Some very simple animals may no ...

and proximal body parts via ascending spinal pathways. The

develops in a rostro-caudal manner, with rostral regions in the midline giving rise to the vermis, and

caudal Caudal may refer to: Anatomy * Caudal (anatomical term) (from Latin ''cauda''; tail), used to describe how close something is to the trailing end of an organism * Caudal artery, the portion of the dorsal aorta of a vertebrate that passes into the ...

regions developing into the cerebellar hemispheres. By 4 months of

prenatal development Prenatal development () includes the development of the embryo and of the fetus during a viviparous animal's gestation. Prenatal development starts with fertilization, in the germinal stage of embryonic development, and continues in fetal devel ...

, the vermis becomes fully foliated, while development of the hemispheres lags by 30–60 days. Postnatally,

proliferation Proliferation may refer to: Weapons *Nuclear proliferation, the spread of nuclear weapons, material, and technology *Chemical weapon proliferation, the spread of chemical weapons, material, and technology * Small arms proliferation, the spread of ...

and organization of the cellular components of the

continues, with completion of the foliation pattern by 7 months of life and final

migration Migration, migratory, or migrate may refer to: Human migration * Human migration, physical movement by humans from one region to another ** International migration, when peoples cross state boundaries and stay in the host state for some minimum l ...

, proliferation, and arborization of cerebellar neurons by 20 months. Inspection of the posterior fossa is a common feature of prenatal ultrasound and is used primarily to determine whether excess fluid or malformations of the cerebellum exist. Anomalies of the cerebellar vermis are diagnosed in this manner and include

phenotypes In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology or physical form and structure, its developmental processes, its biochemical and physiological prop ...

consistent with Dandy–Walker malformation, rhombencephalosynapsis, displaying no vermis with fusion of the cerebellar hemispheres, pontocerebellar hypoplasia, or stunted growth of the cerebellum, and neoplasms. In neonates, hypoxic injury to the cerebellum is fairly common, resulting in neuronal loss and gliosis. Symptoms of these disorders range from mild loss of fine

motor control Motor control is the regulation of movement in organisms that possess a nervous system. Motor control includes reflexes as well as directed movement. To control movement, the nervous system must integrate multimodal sensory information (both f ...

to severe

mental retardation Intellectual disability (ID), also known as general learning disability in the United Kingdom and formerly mental retardation, Rosa's Law, Pub. L. 111-256124 Stat. 2643(2010). is a generalized neurodevelopmental disorder characterized by signif ...

and

death Death is the irreversible cessation of all biological functions that sustain an organism. For organisms with a brain, death can also be defined as the irreversible cessation of functioning of the whole brain, including brainstem, and brain ...

. Karyotyping has shown that most pathologies associated with the vermis are inherited though an

autosomal recessive In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant and ...

pattern, with most known

mutations In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, mitosi ...

occurring on the

X chromosome The X chromosome is one of the two sex-determining chromosomes ( allosomes) in many organisms, including mammals (the other is the Y chromosome), and is found in both males and females. It is a part of the XY sex-determination system and XO se ...

. The vermis is intimately associated with all regions of the cerebellar cortex, which can be divided into three functional parts, each having distinct connections with the

brain The brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It consists of nervous tissue and is typically located in the head ( cephalization), usually near organs for special ...

and

spinal cord The spinal cord is a long, thin, tubular structure made up of nervous tissue, which extends from the medulla oblongata in the brainstem to the lumbar region of the vertebral column (backbone). The backbone encloses the central canal of the spin ...

. These regions are the vestibulocerebellum, which is responsible primarily for the control of eye movements; the spinocerebellum, involved in fine tune body and limb movement; and the

cerebrocerebellum 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 ...

, which is associated with planning, initiation and timing of movements.

Structure

The vermis is the unpaired, median portion of the cerebellum that connects the two hemispheres. Both the vermis and the hemispheres are composed of lobules formed by groups of

folia ''La Folía'' (Spanish), or ''Follies'' (English), also known as ''folies d'Espagne'' (French), ''La Follia'' (Italian), and ''Folia'' (Portuguese), is one of the oldest remembered European musical themes, or primary material, generally melodic ...

. There are nine lobules of the vermis: lingula, central lobule, culmen, clivus, folium of the vermis, tuber, pyramid, uvula and nodule. These lobules are often difficult to observe during human anatomy classes and may vary in size, shape and number of folia. It has been shown that folia of the cerebellum exhibit frequent variations in form, number and arrangement between individuals.

Lobe anatomy

The lingula is the first lobule of the upper portion of the vermis on the superoinferior axis and pertains to the paleocerebellum together with the central lobule, culmen, pyramid and uvula. It is separated from the central lobule by the pre-central fissure. The central lobule is the second lobule of the upper portion of the vermis on the superoinferior axis. The culmen is the third and largest lobule of the upper portion of the vermis on the superoinferior axis. It is separated from the declive by the primary fissure and is related with the

quadrangular lobule of the hemisphere. The pyramid is the seventh lobule of the vermis on the superoinferior axis. It is separated from the tuber and uvula by the pre-pyramidal and secondary fissures, respectively. This lobule is related with the biventral lobule of the hemisphere. The uvula is the second largest lobule, following the culmen. It pertains to the paleocerebellum and is separated from the nodule by the posterolateral fissure.

Spinocerebellum

The spinocerebellum receives

proprioception Proprioception ( ), also referred to as kinaesthesia (or kinesthesia), is the sense of self-movement, force, and body position. It is sometimes described as the "sixth sense". Proprioception is mediated by proprioceptors, mechanosensory neurons ...

input from the dorsal columns of the

(including the

spinocerebellar tract The spinocerebellar tract is a nerve tract originating in the spinal cord and terminating in the same side ( ipsilateral) of the cerebellum. Origins of proprioceptive information Proprioceptive information is obtained by Golgi tendon organs and ...

) and from the

trigeminal nerve In neuroanatomy, the trigeminal nerve (literal translation, lit. ''triplet'' nerve), also known as the fifth cranial nerve, cranial nerve V, or simply CN V, is a cranial nerve responsible for Sense, sensation in the face and motor functions ...

, as well as from visual and auditory systems. It sends fibers to deep cerebellar nuclei that, in turn, project to both the

cerebral cortex The cerebral cortex, also known as the cerebral mantle, is the outer layer of neural tissue of the cerebrum of the brain in humans and other mammals. The cerebral cortex mostly consists of the six-layered neocortex, with just 10% consisting o ...

and the

brain stem The brainstem (or brain stem) is the posterior stalk-like part of the brain that connects the cerebrum with the spinal cord. In the human brain the brainstem is composed of the midbrain, the pons, and the medulla oblongata. The midbrain is cont ...

, thus providing modulation of descending motor systems. This region comprises the vermis and intermediate parts of the cerebellar hemispheres. Sensory information from the periphery and from the primary motor and

somatosensory cortex In physiology, the somatosensory system is the network of neural structures in the brain and body that produce the perception of touch ( haptic perception), as well as temperature ( thermoception), body position ( proprioception), and pain. It ...

terminate in this region. Purkinje cells of the vermis project to the fastigial nucleus, controlling the

axial Axial may refer to: * one of the anatomical directions describing relationships in an animal body * In geometry: :* a geometric term of location :* an axis of rotation * In chemistry, referring to an axial bond * a type of modal frame, in music * ...

and

proximal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position ...

musculature involved in the execution of limb movements. Purkinje cells in the intermediate zone of the spinocerebellum project to the interposed nuclei, which control the distal musculature components of the descending motor pathways needed for limb movement. Both of these nuclei include projections to the

motor cortex The motor cortex is the region of the cerebral cortex believed to be involved in the planning, control, and execution of voluntary movements. The motor cortex is an area of the frontal lobe located in the posterior precentral gyrus immediately ...

in the

cerebrum The cerebrum, telencephalon or endbrain is the largest part of the brain containing the cerebral cortex (of the two cerebral hemispheres), as well as several subcortical structures, including the hippocampus, basal ganglia, and olfactory bulb. ...

Nuclei

The interposed nucleus is smaller than the dentate nucleus but larger than the fastigial nucleus and functions to modulate muscle stretch reflexes of distal musculature. It is located

dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage * Dorsal co ...

to the fourth ventricle and lateral to the fastigial nucleus; it receives

afferent Afferent may refer to: Anatomical structures Meaning "conveying towards a center": * Afferent arterioles, blood vessels that supply the nephrons * Afferent lymphatic vessels, lymph vessels that carry lymph to a lymph node * Afferent nerve fiber ...

neuronal supply from the anterior lobe of the cerebellum and sends output via the superior cerebellar peduncle and the red nucleus. The fastigial nucleus is the most medial efferent cerebellar nucleus, targeting the pontine and medullary

reticular formation The reticular formation is a set of interconnected nuclei that are located throughout the brainstem. It is not anatomically well defined, because it includes neurons located in different parts of the brain. The neurons of the reticular formation ...

as well as the vestibular nuclei. This region deals with antigravity muscle groups and other synergies involved with standing and walking. It is thought that fastigial nuclei

axons An axon (from Greek ἄξων ''áxōn'', axis), or nerve fiber (or nerve fibre: see spelling differences), is a long, slender projection of a nerve cell, or neuron, in vertebrates, that typically conducts electrical impulses known as action po ...

are

excitatory In neuroscience, an excitatory postsynaptic potential (EPSP) is a postsynaptic potential that makes the postsynaptic neuron more likely to fire an action potential. This temporary depolarization of postsynaptic membrane potential, caused by the ...

and project beyond the

, likely using

glutamate Glutamic acid (symbol Glu or E; the ionic form is known as glutamate) is an α-amino acid that is used by almost all living beings in the biosynthesis of proteins. It is a non-essential nutrient for humans, meaning that the human body can syn ...

and

aspartate Aspartic acid (symbol Asp or D; the ionic form is known as aspartate), is an α-amino acid that is used in the biosynthesis of proteins. Like all other amino acids, it contains an amino group and a carboxylic acid. Its α-amino group is in the pro ...

neurotransmitters A neurotransmitter is a signaling molecule secreted by a neuron to affect another cell across a synapse. The cell receiving the signal, any main body part or target cell, may be another neuron, but could also be a gland or muscle cell. Neuro ...

Pathology

Malformations of the posterior fossa have been recognized more frequently during the past few

decade A decade () is a period of ten years. Decades may describe any ten-year period, such as those of a person's life, or refer to specific groupings of calendar years. Usage Any period of ten years is a "decade". For example, the statement that "d ...

s as the result of recent advances in technology. Malformations of the cerebellar vermis were first identified using pneumoencephalography, where air is injected into the

cerebrospinal fluid Cerebrospinal fluid (CSF) is a clear, colorless body fluid found within the tissue that surrounds the brain and spinal cord of all vertebrates. CSF is produced by specialised ependymal cells in the choroid plexus of the ventricles of the bra ...

spaces of the

; displaced, occluded or

dysplastic Dysplasia is any of various types of abnormal growth or development of cells (microscopic scale) or organs (macroscopic scale), and the abnormal histology or anatomical structure(s) resulting from such growth. Dysplasias on a mainly microscopic ...

structures could be identified. Upon the advent of

computerized tomography A computed tomography scan (CT scan; formerly called computed axial tomography scan or CAT scan) is a medical imaging technique used to obtain detailed internal images of the body. The personnel that perform CT scans are called radiographers ...

(CT) and magnetic resonance imaging (MRI), the resolution of cranial structures including the mid-hindbrain regions improved dramatically.

Joubert syndrome

Joubert syndrome (JS) is one of the most commonly diagnosed syndromes associated with the molar tooth sign (MTS), or

hypotonia Hypotonia is a state of low muscle tone (the amount of tension or resistance to stretch in a muscle), often involving reduced muscle strength. Hypotonia is not a specific medical disorder, but a potential manifestation of many different diseases ...

infancy An infant or baby is the very young offspring of human beings. ''Infant'' (from the Latin word ''infans'', meaning 'unable to speak' or 'speechless') is a formal or specialised synonym for the common term ''baby''. The terms may also be used to ...

with later development of

ataxia Ataxia is a neurological sign consisting of lack of voluntary coordination of muscle movements that can include gait abnormality, speech changes, and abnormalities in eye movements. Ataxia is a clinical manifestation indicating dysfunction of t ...

, developmental delays,

, abnormal breathing patterns, abnormal eye movements specific to oculomotor

apraxia Apraxia is a motor disorder caused by damage to the brain (specifically the posterior parietal cortex or corpus callosum), which causes difficulty with motor planning to perform tasks or movements. The nature of the damage determines the disor ...

, or the presence of the MTS on the cranial MRI. JS is an

condition with an estimated

prevalence In epidemiology, prevalence is the proportion of a particular population found to be affected by a medical condition (typically a disease or a risk factor such as smoking or seatbelt use) at a specific time. It is derived by comparing the number o ...

of 1: 100,000.

Dandy Walker malformation

Dandy Walker malformation A dandy is a man who places particular importance upon physical appearance, refined language, and leisurely hobbies, pursued with the appearance of nonchalance. A dandy could be a self-made man who strove to imitate an aristocratic lifestyle desp ...

is a relatively common

congenital A birth defect, also known as a congenital disorder, is an abnormal condition that is present at birth regardless of its cause. Birth defects may result in disabilities that may be physical, intellectual, or developmental. The disabilities c ...

brain malformation with a prevalence of 1:30,000 live births. Dandy Walker malformation is characterized by enlarged posterior fossa and in which the cerebellar vermis is completely absent, or present in a rudimentary form, sometimes rotated accompanied by an elevation of the fourth ventricle. It is also commonly associated with

dysplasia Dysplasia is any of various types of abnormal growth or development of cells ( microscopic scale) or organs (macroscopic scale), and the abnormal histology or anatomical structure(s) resulting from such growth. Dysplasias on a mainly microscopi ...

s of

brainstem The brainstem (or brain stem) is the posterior stalk-like part of the brain that connects the cerebrum with the spinal cord. In the human brain the brainstem is composed of the midbrain, the pons, and the medulla oblongata. The midbrain is ...

nuclei. DWM has been reported to be in association with a wide array of

chromosomal A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...

anomalies, including

trisomy 18 A trisomy is a type of polysomy in which there are three instances of a particular chromosome, instead of the normal two. A trisomy is a type of aneuploidy (an abnormal number of chromosomes). Description and causes Most organisms that reprod ...

, trisomy 9, and trisomy 13. Surveys suggest that prenatal exposure to teratogens such as

rubella Rubella, also known as German measles or three-day measles, is an infection caused by the rubella virus. This disease is often mild, with half of people not realizing that they are infected. A rash may start around two weeks after exposure and ...

alcohol Alcohol most commonly refers to: * Alcohol (chemistry), an organic compound in which a hydroxyl group is bound to a carbon atom * Alcohol (drug), an intoxicant found in alcoholic drinks Alcohol may also refer to: Chemicals * Ethanol, one of sev ...

are correlated with development of Dandy Walker malformation.

Rhombencephalosynapsis

Rhombencephalosynapsis is an anomaly characterized by the absence or severe dysgenesis of the cerebellar vermis with fusion of the cerebellar hemispheres, peduncles, and dentate nuclei. Diagnostic features include fusion of the midbrain colliculi,

hydrocephalus Hydrocephalus is a condition in which an accumulation of cerebrospinal fluid (CSF) occurs within the brain. This typically causes increased pressure inside the skull. Older people may have headaches, double vision, poor balance, urinary in ...

, absence of the

corpus callosum The corpus callosum (Latin for "tough body"), also callosal commissure, is a wide, thick nerve tract, consisting of a flat bundle of commissural fibers, beneath the cerebral cortex in the brain. The corpus callosum is only found in placental ...

other midline structural brain malformations.

Damage

Lesions to the vermis commonly give rise to

clinical depression Major depressive disorder (MDD), also known as clinical depression, is a mental disorder characterized by at least two weeks of pervasive low mood, low self-esteem, and loss of interest or pleasure in normally enjoyable activities. Introdu ...

, inappropriate emotional displays (e.g. unwarranted giggling) in addition to movement disorders.

Comparative anatomy

Early neurophysiologists suggest that

retina The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which then ...

l and

inertia Inertia is the idea that an object will continue its current motion until some force causes its speed or direction to change. The term is properly understood as shorthand for "the principle of inertia" as described by Newton in his first law o ...

l signals were selected for about 450 million years ago by primitive

brainstem The brainstem (or brain stem) is the posterior stalk-like part of the brain that connects the cerebrum with the spinal cord. In the human brain the brainstem is composed of the midbrain, the pons, and the medulla oblongata. The midbrain is ...

-cerebellar circuitry because of their relationship with the environment. Microscopically, it is evident that

Purkinje cell Purkinje cells, or Purkinje neurons, are a class of GABAergic inhibitory neurons located in the cerebellum. They are named after their discoverer, Czech anatomist Jan Evangelista Purkyně, who characterized the cells in 1839. Structure The ...

precursors arose from granule cells, first forming in irregular patterns, then progressively becoming organized in a layered fashion. Evolutionarily, the Purkinje cells then developed extensive dendritic trees that increasingly became confined to a single plane, through which the

of granule cells threaded, eventually forming a neuronal grid of right angles. The origin of the

is in close association with that of the nuclei of the vestibular cranial nerve and lateral line

nerves A nerve is an enclosed, cable-like bundle of nerve fibers (called axons) in the peripheral nervous system. A nerve transmits electrical impulses. It is the basic unit of the peripheral nervous system. A nerve provides a common pathway for the e ...

, perhaps suggesting that this part of the cerebellum originated as a means of carrying out transformations of the coordinate system from input data of the

vestibular organ The vestibular system, in vertebrates, is a sensory system that creates the sense of balance and spatial orientation for the purpose of coordinating movement with balance. Together with the cochlea, a part of the auditory system, it constitutes ...

and the lateral line organs. This suggests that the function of the cerebellum evolved as a mode of computing and representing an image relating to the position of the body in space. The cerebellar vermis evolved in conjunction with the hemispheres; this is seen in

lampreys Lampreys (sometimes inaccurately called lamprey eels) are an ancient extant lineage of jawless fish of the order Petromyzontiformes , placed in the superclass Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like ...

and higher

vertebrates Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, wi ...

In fish

, the cerebellar vermis develops between two bilaterally symmetrical formations located

to the upper end of the

medulla oblongata The medulla oblongata or simply medulla is a long stem-like structure which makes up the lower part of the brainstem. It is anterior and partially inferior to the cerebellum. It is a cone-shaped neuronal mass responsible for autonomic (involun ...

, or rhombencephalon. This is the region of termination for the fibers of the vestibular nerve and lateral line nerves; thus, these are the oldest

paths to the cerebellum and cerebellar vermis. In bony fish, or teleosts, it has been proposed that the cerebellar auricles, which receive a large amount of input from the vestibulolateral line system, constitute the vestibulocerebellum and are homologues of the flocculonodular lobe of higher vertebrates along with the corpus cerebelli, which receives spinocerebellar and tectocerebellar fibers. The

labyrinth In Greek mythology, the Labyrinth (, ) was an elaborate, confusing structure designed and built by the legendary artificer Daedalus for King Minos of Crete at Knossos. Its function was to hold the Minotaur, the monster eventually killed by the ...

and the lateral line organs of

have structural and functional similarity. An important difference between the two structures is that the arrangement of the lateral line organs are such that they are sensitive to relative motion of the fluid surrounding the animal, whereas the labyrinths, having very similar sensing mechanisms, are sensitive to

endolymph Endolymph is the fluid contained in the membranous labyrinth of the inner ear. The major cation in endolymph is potassium, with the values of sodium and potassium concentration in the endolymph being 0.91 mM and 154 mM, respectively. ...

, providing information concerning the animal's own equilibrium of the body and orientation in space.

Additional images

File:Vermis and paravermis by Sanjoy Sanyal.webm, Dissection video. Explains the position of vermis. File:Cerebellar vermis --- animation.gif, Animation. Vermis is highlighted in red. File:Slide2BRA.JPG, Brainstem. Posterior view. File:Human brain midsagittal view description.JPG, Midsagittal view File:CerebellumDiv.png, Schematic representation of the major anatomical subdivisions of the cerebellum.

References

External links

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at University of Texas Southwestern Medical Center at Dallas
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