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 Alertness is the state of active attention by high sensory awareness such as being watchful and prompt to meet danger or emergency, or being quick to perceive and act. It is related for psychology . A lack of alertness is a symptom of a ...

), and temperature regulation. The name comes from the Greek ''mesos'', "middle", and ''enkephalos'', "brain".

Structure

Human brainstem-thalamus posterior view description

The principal regions of the midbrain are the

tectum The midbrain or mesencephalon is the forward-most portion of the brainstem and is associated with vision, hearing, motor control, sleep and wakefulness, arousal (alertness), and temperature regulation. The name comes from the Greek ''mesos'', "m ...

, the

cerebral aqueduct The cerebral aqueduct (aqueductus mesencephali, mesencephalic duct, sylvian aqueduct or aqueduct of Sylvius) is a conduit for cerebrospinal fluid (CSF) that connects the third ventricle to the fourth ventricle of the ventricular system of the brai ...

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 the

cerebral peduncle The cerebral peduncles are the two stalks that attach the cerebrum to the brainstem. They are structures at the front of the midbrain which arise from the ventral pons and contain the large ascending (sensory) and descending (motor) nerve tra ...

s. 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 The thalamus (from Greek θάλαμος, "chamber") is a large mass of gray matter located in the dorsal part of the diencephalon (a division of the forebrain). Nerve fibers project out of the thalamus to the cerebral cortex in all directions, ...

hypothalamus The hypothalamus () is a part of the brain that contains a number of small nuclei with a variety of functions. One of the most important functions is to link the nervous system to the endocrine system via the pituitary gland. The hypothalamu ...

, etc.), while

caudally 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 ...

it adjoins the

hindbrain The hindbrain or rhombencephalon or lower brain 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. Metencephalo ...

(

pons The pons (from Latin , "bridge") is part of the brainstem that in humans and other bipeds 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 Va ...

medulla Medulla or Medullary may refer to: Science * Medulla oblongata, a part of the brain stem * Renal medulla, a part of the kidney * Adrenal medulla, a part of the adrenal gland * Medulla of ovary, a stroma in the center of the ovary * Medulla of th ...

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

, which refers to the region in front of the

ventricular system 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 ventricular system is con ...

, 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 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 ...

, which exerts some control over alertness, takes input from the tectum, and travels both rostrally and caudally from it. The

corpora quadrigemina Corpus is Latin for "body". It may refer to: Linguistics * Text corpus, in linguistics, a large and structured set of texts * Speech corpus, in linguistics, a large set of speech audio files * Corpus linguistics, a branch of linguistics Music * ...

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 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 ...

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 In humans, the tectospinal tract (or colliculospinal tract) is a nerve tract that coordinates head and eye movements. This tract is part of the extrapyramidal system and connects the midbrain tectum, and cervical regions of the spinal cord. It ...

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 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, ventral projec ...

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

fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...

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 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 ...

; 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 The trochlear nerve (), ( lit. ''pulley-like'' nerve) also known as the fourth cranial nerve, cranial nerve IV, or CN IV, is a cranial nerve that innervates just one muscle: the superior oblique muscle of the eye, which operates through the pul ...

– 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 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 left lateral v ...

(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 ve ...

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

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 ...

. The cerebral aqueduct is a narrow channel located between the tectum and the tegmentum, and is surrounded by the

periaqueductal grey The periaqueductal gray (PAG, also known as the central gray) is a brain region that plays a critical role in autonomic function, motivated behavior and behavioural responses to threatening stimuli. PAG is also the primary control center for d ...

,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 located on the midline of the brainstem and is one of the raphe nuclei. It has rostral and caudal subdivisions. * The rostral aspect of the ''dorsal'' raphe is further divided into interfascicular, ventral, ventrolatera ...

(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

by the

superior cerebellar peduncles In the human brain, the superior cerebellar peduncle (brachium conjunctivum) 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 cer ...

, 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 The median raphe nucleus (MRN or MnR), also known as the nucleus raphes medianus (NRM)Federative Committee on Anatomical Terminology (FCAT) (1998). ''Terminologia Anatomica''. Stuttgart: Thieme or superior central nucleus, is a brain region compos ...

, 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 In neuroanatomy, 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 (cross) in the brainstem, specifically in the m ...

– 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 part of the anterolateral system or the ventrolateral system, a sensory pathway to the thalamus. From the ventral posterolateral nucleus in the thalamus, sensory information is relayed upward to the somatosensory cor ...

– 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 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 the ...

– 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 The mammillary bodies are a pair of small round bodies, located on the undersurface of the brain that, as part of the diencephalon, form part of the limbic system. They are located at the ends of the anterior arches of the fornix. They consist o ...

(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

(of the

Cerebral peduncles

The

s 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

. 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

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

corticobulbar In neuroanatomy, the corticobulbar (or corticonuclear) tract is a two-neuron white matter motor pathway connecting the motor cortex in the cerebral cortex to the medullary pyramids, which are part of the brainstem's medulla oblongata (also called ...

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 neur ...

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 white matter structure situated in the inferomedial part of each cerebral hemisphere of the brain. It carries information past the basal ganglia, separating the caudate nucleus and the thalamus from the putamen and the g ...

. 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 Addiction is a neuropsychological disorder characterized by a persistent and intense urge to engage in certain behaviors, one of which is the usage of a drug, despite substantial harm and other negative consequences. Repetitive drug use o ...

, and other functions. There are two regions within the substantia nigra – one where neurons are densely packed (the

pars compacta The pars compacta (SNpc) is a portion of the ''substantia nigra'', located in the midbrain. It is formed by dopaminergic neurons and located medial to the pars reticulata. Parkinson's disease is characterized by the death of dopaminergic neurons ...

) and one where they aren't (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 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 Parkinson's disease (PD), or simply Parkinson's, is a long-term degenerative disorder of the central nervous system that mainly affects the motor system. The symptoms usually emerge slowly, and as the disease worsens, non-motor symptoms becom ...

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 basilar artery () is one of the arteries that supplies the brain with oxygen-rich blood. The two vertebral arteries and the basilar artery are known as the vertebral basilar system, which supplies blood to the posterior part of the circle of W ...

. * 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, part of the back of the human brain. The two arteries originate from the distal end of the basilar artery, where it bifur ...

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 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 fold become elevated, a ...

, 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 Anatomical terms of location#Directional terms, rostral (forward-most) portion of the brain. The forebrain (prosencephalon), the midbrain (mesencephalon), and hin ...

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, 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. In ...

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 an accumulation of cerebrospinal fluid (CSF) occurs within the brain. This typically causes increased intracranial pressure, pressure inside the skull. Older people may have headaches, double vision, poor ...

. 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

. Its

is closely associated with

motor system The motor system is the set of central and peripheral structures in the nervous system that support motor functions, i.e. movement. Peripheral structures may include skeletal muscles and neural connections with muscle tissues. Central structur ...

pathways of the

basal ganglia The basal ganglia (BG), or basal nuclei, are a group of subcortical nuclei, of varied origin, in the brains of vertebrates. In humans, and some primates, there are some differences, mainly in the division of the globus pallidus into an extern ...

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

vertebrate 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, ...

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

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

house mice The house mouse (''Mus musculus'') is a small mammal of the order Rodentia, characteristically having a pointed snout, large rounded ears, and a long and almost hairless tail. It is one of the most abundant species of the genus '' Mus''. Althoug ...

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. ()

References

{{Authority control Medical mnemonics Brain anatomy