The midbrain or mesencephalon is the uppermost 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 conti ...

connecting the

diencephalon In the human brain, 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 t ...

and

cerebrum The cerebrum (: cerebra), 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 olfac ...

with the

pons The pons (from Latin , "bridge") is part of the brainstem that in humans and other mammals, lies inferior to the midbrain, superior to the medulla oblongata and anterior to the cerebellum. The pons is also called the pons Varolii ("bridge of ...

. It consists of the cerebral peduncles,

tegmentum The tegmentum (from Latin for "covering") is a general area within the brainstem. The tegmentum is the ventral part of the midbrain and the tectum is the dorsal part of the midbrain. It is located between the ventricular system and distinctive b ...

, and

tectum The midbrain or mesencephalon is the uppermost portion of the brainstem connecting the diencephalon and cerebrum with the pons. It consists of the cerebral peduncles, tegmentum, and tectum. It is functionally associated with vision, hearing, mo ...

. It is functionally associated with vision, hearing, motor control, sleep and wakefulness, arousal ( alertness), and temperature regulation.Breedlove, Watson, & Rosenzweig. Biological Psychology, 6th Edition, 2010, pp. 45-46 The name ''mesencephalon'' comes from the Greek ''mesos'', "middle", and ''enkephalos'', "brain".

Structure

Human brainstem-thalamus posterior view description

The midbrain is the shortest segment of the brainstem, measuring less than 2cm in length. It is situated mostly in the

posterior cranial fossa The posterior cranial fossa is the part of the cranial cavity located between the foramen magnum, and tentorium cerebelli. It is formed by the sphenoid bones, temporal bones, and occipital bone. It lodges the cerebellum, and parts of the brai ...

, with its superior part extending above the tentorial notch. The principal regions of the midbrain are the

, the

cerebral aqueduct The cerebral aqueduct (aqueduct of the midbrain, aqueduct of Sylvius, Sylvian aqueduct, mesencephalic duct) is a small, narrow tube connecting the third and fourth ventricles of the brain. The cerebral aqueduct is a midline structure that passe ...

, and the cerebral peduncles. Rostrally the midbrain adjoins the

(

thalamus The thalamus (: thalami; from Greek language, Greek Wikt:θάλαμος, θάλαμος, "chamber") is a large mass of gray matter on the lateral wall of the third ventricle forming the wikt:dorsal, dorsal part of the diencephalon (a division of ...

hypothalamus The hypothalamus (: hypothalami; ) is a small part of the vertebrate brain that contains a number of nucleus (neuroanatomy), nuclei with a variety of functions. One of the most important functions is to link the nervous system to the endocrin ...

, etc.), while

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

it adjoins the

hindbrain The hindbrain, rhombencephalon (shaped like a rhombus) is a developmental categorization of portions of the central nervous system in vertebrates. It includes the medulla, pons, and cerebellum. Together they support vital bodily processes. Met ...

(

, medulla and

cerebellum The cerebellum (: cerebella or cerebellums; 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 it or eve ...

). 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 part of the midbrain dorsal to the

.The position of the tectum is contrasted with the

, which refers to the region in front of the

ventricular system In neuroanatomy, the ventricular system is a set of four interconnected cavities known as cerebral ventricles in the brain. Within each ventricle is a region of choroid plexus which produces the circulating cerebrospinal fluid (CSF). The ventric ...

, 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 process some visual information, aid the

decussation 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 Chiasm (anatomy), chi ...

of several fibres of the

optic nerve In neuroanatomy, the optic nerve, also known as the second cranial nerve, cranial nerve II, or simply CN II, is a paired cranial nerve that transmits visual system, visual information from the retina to the brain. In humans, the optic nerve i ...

(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 t ...

s of the

neck The neck is the part of the body in many vertebrates that connects the head to the torso. It supports the weight of the head and protects the nerves that transmit sensory and motor information between the brain and the rest of the body. Addition ...

, and co-ordinates head and eye movements. Each superior colliculus also sends information to the corresponding

lateral geniculate nucleus In neuroanatomy, the lateral geniculate nucleus (LGN; also called the lateral geniculate body or lateral geniculate complex) is a structure in the thalamus and a key component of the mammalian visual pathway. It is a small, ovoid, Anatomical ter ...

, with which it is directly connected. The homologous structure to the superior colliculus in non mammalian vertebrates including

fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...

and

amphibian Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...

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 – located just above the trochlear nerve – process certain auditory information. Each inferior colliculus sends information to the corresponding

medial geniculate nucleus The medial geniculate nucleus (MGN) or medial geniculate body (MGB) is part of the auditory thalamus and represents the thalamic relay between the inferior colliculus (IC) and the auditory cortex (AC). It is made up of a number of sub-nuclei that ...

, with which it is directly connected.

Cerebral aqueduct

The

is the part of the

which links the

third ventricle The third ventricle is one of the four connected cerebral ventricles of the ventricular system within the mammalian brain. It is a slit-like cavity formed in the diencephalon between the two thalami, in the midline between the right and lef ...

(rostrally) with the

fourth ventricle The fourth ventricle is one of the four connected fluid-filled cavities within the human brain. These cavities, known collectively as the ventricular system, consist of the left and right lateral ventricles, the third ventricle, and the fourth ...

(caudally); as such it is responsible for continuing the circulation of

cerebrospinal fluid Cerebrospinal fluid (CSF) is a clear, colorless Extracellular fluid#Transcellular fluid, transcellular body fluid found within the meninges, meningeal tissue that surrounds the vertebrate brain and spinal cord, and in the ventricular system, ven ...

. 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 The dorsal raphe nucleus is one of the raphe nuclei. It is situated in the brainstem at the midline. It has rostral and caudal subdivisions: * The rostral aspect of the ''dorsal'' raphe is further divided into interfascicular, ventral, ventrolater ...

(which releases

serotonin Serotonin (), also known as 5-hydroxytryptamine (5-HT), is a monoamine neurotransmitter with a wide range of functions in both the central nervous system (CNS) and also peripheral tissues. It is involved in mood, cognition, reward, learning, ...

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 f ...

s are similarly located at the ventral side of the periaqueductal grey – the pair of oculomotor nuclei (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 midbra ...

is the portion of the midbrain ventral to the cerebral aqueduct, and is much larger in size than the tectum. It communicates with the

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 in the brainstem that spans from the lower end of the medulla oblongata to the upper end of the midbrain. The neurons of the reticular formation make up a complex set of neural networks ...

. A number of distinct

nerve tract A nerve tract is a bundle of nerve fibers (axons) connecting Nucleus (neuroanatomy), nuclei of the central nervous system. In the peripheral nervous system, this is known as a nerve fascicle, and has associated nervous tissue, connective tissue. T ...

s between other parts of the brain pass through it. The

medial lemniscus The medial lemniscus, also known as Reil's band or Reil's ribbon (for German anatomist Johann Christian Reil), is a large ascending bundle of heavily myelinated axons that decussate in the brainstem The brainstem (or brain stem) is the pos ...

– 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 The spinothalamic tract is a nerve tract in the anterolateral system in the spinal cord. This tract is an ascending sensory pathway to the thalamus. From the ventral posterolateral nucleus in the thalamus, sensory information is relayed upwar ...

– 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 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 The ventral tegmental area (VTA) (tegmentum is Latin for ''covering''), also known as the ventral tegmental area of Tsai, or simply ventral tegmentum, is a group of neurons located close to the midline on the floor of the midbrain. The VTA is th ...

– 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 chemical of the catecholamine and phenethylamine families. It is an amine synthesized ...

-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 In the human brain, 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 t ...

) and

(of the

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 cerebral crus, cerebal crura. It has been found in humans and macaques, but not in rats or mice, showing that this is a relatively new evolutiona ...

, which is a cistern filled with

. The majority of each lobe constitutes the cerebral crus. The cerebral crus are the main tracts descending from the

to caudal parts of the central nervous system; the central and medial ventral portions contain the corticobulbar and

corticospinal tract The corticospinal tract is a white matter motor pathway starting at the cerebral cortex that terminates on lower motor neurons and interneurons in the spinal cord, controlling movements of the limbs and trunk. There are more than one million neu ...

s, while the remainder of each crus primarily contains tracts connecting the cortex to the

. 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 internal capsule is a paired white matter structure, as a two-way nerve tract, tract, carrying afferent nerve fiber, ascending and efferent nerve fiber, descending axon, fibers, to and from the cerebral cortex. The internal capsule is situate ...

. 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 a ...

(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, Attitude (psychology), attitudes, and preferences. The ability to learn is possessed by humans, non-human animals, and ...

addiction Addiction is a neuropsychological disorder characterized by a persistent and intense urge to use a drug or engage in a behavior that produces natural reward, despite substantial harm and other negative consequences. Repetitive drug use can ...

, and other functions. There are two regions within the substantia nigra – one where neurons are densely packed (the pars compacta) and one where they are not (the

pars reticulata The pars reticulata (SNpr) is a portion of the substantia nigra and is located lateral to the pars compacta. Most of the neurons that project out of the pars reticulata are inhibitory GABAergic neurons (i.e., these neurons release GABA, which is ...

), 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 a hormone, neurotransmitter and neuromodulator. The name "noradrenaline" (from L ...

; the loss of dopamine-producing neurons in this region contributes to the progression of

Parkinson's disease Parkinson's disease (PD), or simply Parkinson's, is a neurodegenerative disease primarily of the central nervous system, affecting both motor system, motor and non-motor systems. Symptoms typically develop gradually and non-motor issues become ...

Blood supply

The midbrain is supplied by the following

arteries An artery () is a blood vessel in humans and most other animals that takes oxygenated blood away from the heart in the systemic circulation to one or more parts of the body. Exceptions that carry deoxygenated blood are the pulmonary arteries in ...

: * 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 t ...

. * 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 The posterior cerebral artery (PCA) is one of a pair of cerebral arteries that supply oxygenated blood to the occipital lobe, as well as the medial and inferior aspects of the temporal lobe of the human brain. The two arteries originate from the d ...

Venous blood Venous blood is deoxygenated blood which travels from the peripheral blood vessels, through the venous system into the right atrium of the heart. Deoxygenated blood is then pumped by the right ventricle to the lungs via the pulmonary artery whi ...

from the midbrain is mostly drained into the basal vein as it passes around the peduncle. Some venous blood from the colliculi drains to the great cerebral vein.

Development

During

embryonic development In developmental biology, animal embryonic development, also known as animal embryogenesis, is the developmental stage of an animal embryo. Embryonic development starts with the fertilization of an egg cell (ovum) by a sperm, sperm cell (spermat ...

, the midbrain (also known as the mesencephalon) arises from the second vesicle of the

neural tube In the developing chordate (including vertebrates), the neural tube is the embryonic precursor to the central nervous system, which is made up of the brain and spinal cord. The neural groove gradually deepens as the neural folds become elevated, ...

, while the interior of this portion of the tube becomes the cerebral aqueduct. Unlike the other two vesicles – the

forebrain In the anatomy of the brain of vertebrates, the forebrain or prosencephalon is the rostral (forward-most) portion of the brain. The forebrain controls body temperature, reproductive functions, eating, sleeping, and the display of emotions. Ve ...

and

– 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 The cerebrum (: cerebra), 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 olf ...

and the

. 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 Hydrocephalus is a condition in which cerebrospinal fluid (CSF) builds up within the brain, which can cause pressure to increase in the skull. Symptoms may vary according to age. Headaches and double vision are common. Elderly adults with n ...

. The tectum is derived in embryonic development from the alar plate of the neural tube.

Function

The midbrain is the uppermost part of the

. Its

is closely associated with

motor system The motor system is the set of central nervous system, central and peripheral nervous system, peripheral structures in the nervous system that support motor functions, i.e. movement. Peripheral structures may include skeletal muscles and Efferen ...

pathways of the

basal ganglia The basal ganglia (BG) or basal nuclei are a group of subcortical Nucleus (neuroanatomy), nuclei found in the brains of vertebrates. In humans and other primates, differences exist, primarily in the division of the globus pallidus into externa ...

. The human midbrain is archipallian in origin, meaning that its general architecture is shared with the most ancient of

vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...

Dopamine Dopamine (DA, a contraction of 3,4-dihydroxyphenethylamine) is a neuromodulatory molecule that plays several important roles in cells. It is an organic chemical of the catecholamine and phenethylamine families. It is an amine synthesized ...

produced in the substantia nigra and

plays a role in movement, movement planning, excitation, motivation and habituation of species from humans to the most elementary animals such as insects. Laboratory mice from lines that have been selectively bred for high voluntary wheel running 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. ()

References

{{Authority control Medical mnemonics Brain anatomy