Midbrain - superior colliculus.svg
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The midbrain or mesencephalon is the forward-most portion of the
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 cont ...
and is associated with vision, hearing, motor control, sleep and wakefulness, arousal ( alertness), and temperature regulation. The name comes from the Greek ''mesos'', "middle", and ''enkephalos'', "brain".


Structure

The principal regions of the midbrain are the tectum, the cerebral aqueduct, tegmentum, and the cerebral peduncles. Rostrally the midbrain adjoins the
diencephalon The diencephalon (or interbrain) is a division of the forebrain (embryonic ''prosencephalon''). It is situated between the telencephalon and the midbrain (embryonic ''mesencephalon''). The diencephalon has also been known as the 'tweenbrain in ol ...
( thalamus, hypothalamus, etc.), while caudally it adjoins the hindbrain ( pons, medulla and
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 cerebel ...
). In the rostral direction, the midbrain noticeably splays laterally. Sectioning of the midbrain is usually performed axially, at one of two levels – that of the superior colliculi, or that of the inferior colliculi. One common technique for remembering the structures of the midbrain involves visualizing these cross-sections (especially at the level of the superior colliculi) as the upside-down face of a bear, with the cerebral peduncles forming the ears, the cerebral aqueduct the mouth, and the tectum the chin; prominent features of the tegmentum form the eyes and certain sculptural shadows of the face.


Tectum

The tectum (Latin for ''roof'') is the dorsal side of the midbrain. The position of the tectum is contrasted with the tegmentum, which refers to the region in front of the ventricular system, or floor of the midbrain. It is involved in certain
reflex In biology, a reflex, or reflex action, is an involuntary, unplanned sequence or action and nearly instantaneous response to a stimulus. Reflexes are found with varying levels of complexity in organisms with a nervous system. A reflex occurs ...
es in response to visual or auditory stimuli. The reticulospinal tract, which exerts some control over alertness, takes input from the tectum, and travels both rostrally and caudally from it. The corpora quadrigemina are four mounds, called colliculi, in two pairs – a superior and an inferior pair, on the surface of the tectum. The
superior colliculi In neuroanatomy, the superior colliculus () is a structure lying on the roof of the mammalian midbrain. In non-mammalian vertebrates, the homologous structure is known as the optic tectum, or optic lobe. The adjective form ''tectal'' is commonly ...
process some visual information, aid the decussation of several fibres of the optic nerve (some fibres remain ipsilateral), and are involved with saccadic eye movements. The tectospinal tract connects the superior colliculi to the
cervical nerve A spinal nerve is a mixed nerve, which carries motor, sensory, and autonomic signals between the spinal cord and the body. In the human body there are 31 pairs of spinal nerves, one on each side of the vertebral column. These are grouped into the ...
s of the
neck The neck is the part of the body on many vertebrates that connects the head with the torso. The neck supports the weight of the head and protects the nerves that carry sensory and motor information from the brain down to the rest of the body. In ...
, and co-ordinates head and eye movements. Each superior colliculus also sends information to the corresponding lateral geniculate nucleus, with which it is directly connected. The
homologous Homology may refer to: Sciences Biology *Homology (biology), any characteristic of biological organisms that is derived from a common ancestor *Sequence homology, biological homology between DNA, RNA, or protein sequences * Homologous chrom ...
structure to the superior colliculus in non mammalian vertebrates including fish and
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s, is called the optic tectum; in those animals, the optic tectum integrates sensory information from the eyes and certain auditory reflexes. The
inferior colliculi The inferior colliculus (IC) (Latin for ''lower hill'') is the principal midbrain nucleus of the auditory pathway and receives input from several peripheral brainstem nuclei in the auditory pathway, as well as inputs from the auditory cortex. Th ...
– located just above the trochlear nerve – process certain auditory information. Each inferior colliculus sends information to the corresponding medial geniculate nucleus, with which it is directly connected.


Cerebral aqueduct

The cerebral aqueduct is the part of the ventricular system which links the third ventricle (rostrally) with the fourth ventricle (caudally); as such it is responsible for continuing the circulation of cerebrospinal fluid. The cerebral aqueduct is a narrow channel located between the tectum and the tegmentum, and is surrounded by the periaqueductal grey,Martin. Neuroanatomy Text and Atlas, Second edition. 1996, pp. 522-525. which has a role in analgesia, quiescence, and bonding. The dorsal raphe nucleus (which releases
serotonin Serotonin () or 5-hydroxytryptamine (5-HT) is a monoamine neurotransmitter. Its biological function is complex and multifaceted, modulating mood, cognition, reward, learning, memory, and numerous physiological processes such as vomiting and vas ...
in response to certain neural activity) is located at the ventral side of the periaqueductal grey, at the level of the inferior colliculus. The nuclei of two pairs of
cranial nerve Cranial nerves are the nerves that emerge directly from the brain (including the brainstem), of which there are conventionally considered twelve pairs. Cranial nerves relay information between the brain and parts of the body, primarily to and ...
s are similarly located at the ventral side of the periaqueductal grey – the pair of
oculomotor nuclei The fibers of the oculomotor nerve arise from a nucleus in the midbrain, which lies in the gray substance of the floor of the cerebral aqueduct and extends in front of the aqueduct for a short distance into the floor of the third ventricle. From t ...
(which control the eyelid, and most eye movements) is located at the level of the superior colliculus, while the pair of trochlear nuclei (which helps focus vision on more proximal objects) is located caudally to that, at the level of the inferior colliculus, immediately lateral to the dorsal raphe nucleus. The oculomotor nerve emerges from the nucleus by traversing the ventral width of the tegmentum, while the trochlear nerve emerges via the tectum, just below the inferior colliculus itself; the trochlear is the only cranial nerve to exit the brainstem dorsally. The Edinger-Westphal nucleus (which controls the shape of the lens and size of the pupil) is located between the oculomotor nucleus and the cerebral aqueduct.


Tegmentum

The
midbrain tegmentum The midbrain is anatomically delineated into the tectum (roof) and the tegmentum (floor). The midbrain tegmentum extends from the substantia nigra to the cerebral aqueduct in a horizontal section of the midbrain. It forms the floor of the midbrain ...
is the portion of the midbrain ventral to the cerebral aqueduct, and is much larger in size than the tectum. It communicates with 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 cerebel ...
by the superior cerebellar peduncles, which enter at the caudal end, medially, on the ventral side; the cerebellar peduncles are distinctive at the level of the inferior colliculus, where they
decussate Decussation is used in biological contexts to describe a crossing (due to the shape of the Roman numeral for ten, an uppercase 'X' (), ). In Latin anatomical terms, the form is used, e.g. . Similarly, the anatomical term chiasma is named aft ...
, but they dissipate more rostrally. Between these peduncles, on the ventral side, is the median raphe nucleus, which is involved in memory consolidation. The main bulk of the tegmentum contains a complex synaptic network of neurons, primarily involved in homeostasis and reflex actions. It includes portions of the
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 ...
. A number of distinct
nerve tract A nerve tract is a bundle of nerve fibers (axons) connecting nuclei of the central nervous system. In the peripheral nervous system this is known as a nerve, and has associated connective tissue. The main nerve tracts in the central nervous syste ...
s between other parts of the brain pass through it. The medial lemniscus – a narrow ribbon of fibres – passes through in a relatively constant axial position; at the level of the inferior colliculus it is near the lateral edge, on the ventral side, and retains a similar position rostrally (due to widening of the tegmentum towards the rostral end, the position can appears more medial). The spinothalamic tract – another ribbon-like region of fibres – are located at the lateral edge of the tegmentum; at the level of the inferior colliculus it is immediately dorsal to the medial lemiscus, but due to the rostral widening of the tegmentum, is lateral of the medial lemiscus at the level of the superior colliculus. A prominent pair of round, reddish, regions – the red nuclei (which have a role in motor co-ordination) – are located in the rostral portion of the midbrain, somewhat medially, at the level of the superior colliculus. The
rubrospinal tract The rubrospinal tract is a part of the nervous system. It is a part of the lateral indirect extra-pyramidal tract. Structure In the midbrain, it originates in the magnocellular red nucleus, crosses to the other side of the midbrain, and desce ...
emerges from the red nucleus and descends caudally, primarily heading to the cervical portion of the spine, to implement the red nuclei's decisions. The area between the red nuclei, on the ventral side – known as the ventral tegmental area – is the largest
dopamine Dopamine (DA, a contraction of 3,4-dihydroxyphenethylamine) is a neuromodulatory molecule that plays several important roles in cells. It is an organic compound, organic chemical of the catecholamine and phenethylamine families. Dopamine const ...
-producing area in the brain, and is heavily involved in the neural reward system. The ventral tegmental area is in contact with parts of the forebrain – the mammillary bodies (from the
Diencephalon The diencephalon (or interbrain) is a division of the forebrain (embryonic ''prosencephalon''). It is situated between the telencephalon and the midbrain (embryonic ''mesencephalon''). The diencephalon has also been known as the 'tweenbrain in ol ...
) and hypothalamus (of the
diencephalon The diencephalon (or interbrain) is a division of the forebrain (embryonic ''prosencephalon''). It is situated between the telencephalon and the midbrain (embryonic ''mesencephalon''). The diencephalon has also been known as the 'tweenbrain in ol ...
).


Cerebral peduncles

The cerebral peduncles each form a lobe ventrally of the tegmentum, on either side of the midline. Beyond the midbrain, between the lobes, is the
interpeduncular fossa The interpeduncular fossa is a deep depression of the ventral surface of the midbrain between the two crura cerebri. It has been found in humans and macaques, but not in rats or mice, showing that this is a relatively new evolutionary region. A ...
, which is a cistern filled with cerebrospinal fluid . The majority of each lobe constitutes the
cerebral crus The cerebral crus (crus cerebri) is the Anatomical terms of location, anterior portion of the cerebral peduncle which contains the motor tracts, travelling from the cerebral cortex to the pons and spine. The plural of which is cerebral crura. In ...
. The cerebral crus are the main tracts descending from the thalamus to caudal parts of the central nervous system; the central and medial ventral portions contain the corticobulbar and corticospinal tracts, while the remainder of each crus primarily contains tracts connecting the cortex to the pons. Older texts refer to the crus cerebri as the ''cerebral peduncle''; however, the latter term actually covers all fibres communicating with the cerebrum (usually via the diencephalon), and therefore would include much of the tegmentum as well. The remainder of the crus pedunculi – small regions around the main cortical tracts – contain tracts from the internal capsule. The portion of the lobes in connection with the tegmentum, except the most lateral portion, is dominated by a blackened band – the
substantia nigra The substantia nigra (SN) is a basal ganglia structure located in the midbrain that plays an important role in reward and movement. ''Substantia nigra'' is Latin for "black substance", reflecting the fact that parts of the substantia nigra app ...
(literally ''black substance'') – which is the only part of the basal ganglia system outside the forebrain. It is ventrally wider at the rostral end. By means of the basal ganglia, the substantia nigra is involved in motor-planning,
learning Learning is the process of acquiring new understanding, knowledge, behaviors, skills, value (personal and cultural), values, attitudes, and preferences. The ability to learn is possessed by humans, animals, and some machine learning, machines ...
, addiction, and other functions. There are two regions within the substantia nigra – one where neurons are densely packed (the pars compacta) and one where they aren't (the pars reticulata), which serve a different role from one another within the basal ganglia system. The substantia nigra has extremely high production of melanin (hence the colour), dopamine, and
noradrenalin Norepinephrine (NE), also called noradrenaline (NA) or noradrenalin, is an organic chemical in the catecholamine family that functions in the brain and body as both a hormone and neurotransmitter. The name "noradrenaline" (from Latin '' ad'', ...
; the loss of dopamine-producing neurons in this region contributes to the progression of Parkinson's disease.


Blood Supply

The midbrain is supplied by the following arteries: * The tectum is supplied by the
superior cerebellar artery The superior cerebellar artery (SCA) is an artery of the head. It arises near the end of the basilar artery. It is a branch of the basilar artery. It supplies parts of the cerebellum, the midbrain, and other nearby structures. It is the cause of ...
. * The central part of the tegmentum is supplied by the paramedian branches of the basilar artery. * The lateral part of the midbrain is supplied by the posterior cerebral artery.


Development

During
embryonic development An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sperm ...
, the midbrain (also known as the mesencephalon) arises from the second
vesicle Vesicle may refer to: ; In cellular biology or chemistry * Vesicle (biology and chemistry), a supramolecular assembly of lipid molecules, like a cell membrane * Synaptic vesicle ; In human embryology * Vesicle (embryology), bulge-like features o ...
of the neural tube, while the interior of this portion of the tube becomes the cerebral aqueduct. Unlike the other two vesicles – the forebrain and hindbrain – the midbrain does not develop further subdivision for the remainder of neural development. It does not split into other brain areas. while the forebrain, for example, divides into the telencephalon and the
diencephalon The diencephalon (or interbrain) is a division of the forebrain (embryonic ''prosencephalon''). It is situated between the telencephalon and the midbrain (embryonic ''mesencephalon''). The diencephalon has also been known as the 'tweenbrain in ol ...
. Throughout embryonic development, the cells within the midbrain continually multiply; this happens to a much greater extent ventrally than it does dorsally. The outward expansion compresses the still-forming cerebral aqueduct, which can result in partial or total obstruction, leading to congenital hydrocephalus. The tectum is derived in embryonic development from the
alar plate Daminozide—also known as aminozide, Alar, Kylar, SADH, B-995, B-nine, and DMASA,—is a plant growth regulator, a chemical sprayed on fruit to regulate growth, make harvest easier, and keep apples from falling off the trees before they ripen s ...
of the neural tube.


Function

The mesencephalon is considered part of the
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 cont ...
. Its
substantia nigra The substantia nigra (SN) is a basal ganglia structure located in the midbrain that plays an important role in reward and movement. ''Substantia nigra'' is Latin for "black substance", reflecting the fact that parts of the substantia nigra app ...
is closely associated with motor system pathways of the basal ganglia. The human mesencephalon is archipallian in origin, meaning that its general architecture is shared with the most ancient of vertebrates.
Dopamine Dopamine (DA, a contraction of 3,4-dihydroxyphenethylamine) is a neuromodulatory molecule that plays several important roles in cells. It is an organic compound, organic chemical of the catecholamine and phenethylamine families. Dopamine const ...
produced in the
substantia nigra The substantia nigra (SN) is a basal ganglia structure located in the midbrain that plays an important role in reward and movement. ''Substantia nigra'' is Latin for "black substance", reflecting the fact that parts of the substantia nigra app ...
and ventral tegmental area plays a role in movement, movement planning, excitation, motivation and habituation of species from humans to the most elementary animals such as insects. Laboratory house mice from lines that have been
selectively bred Selective breeding (also called artificial selection) is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant mal ...
for high voluntary
wheel running A hamster wheel or running wheel is an exercise device used primarily by hamsters and other rodents, but also by other cursorial animals when given the opportunity. Most of these devices consist of a runged or ridged wheel held on a stand by ...
have enlarged midbrains. The midbrain helps to relay information for vision and hearing.


Related terms

The term "tectal plate" or "quadrigeminal plate" is used to describe the junction of the gray and white matter in the embryo. ()


See also

*
List of regions in the human brain The human brain anatomical regions are ordered following standard neuroanatomy hierarchies. Functional, connective, and developmental regions are listed in parentheses where appropriate. Hindbrain (rhombencephalon) Myelencephalon * Med ...


References

{{Authority control Medical mnemonics Brain anatomy