Metencephalon

Metencephalon
	Diagram depicting the main subdivisions of the embryonic vertebrate brain. These regions will later differentiate into forebrain, midbrain and hindbrain structures.
	Pons and cerebellum.
Identifiers
MeSH	D020540
NeuroNames	543
NeuroLex ID	birnlex_965
TA	A14.1.03.004
FMA	62003
	Anatomical terms of neuroanatomy [ edit on Wikidata]

Last updated April 24, 2019

The metencephalon is the embryonic part of the hindbrain that differentiates into the pons and the cerebellum. It contains a portion of the fourth ventricle and the trigeminal nerve (CN V), abducens nerve (CN VI), facial nerve (CN VII), and a portion of the vestibulocochlear nerve (CN VIII).

An embryo is an early stage of development of a multicellular diploid eukaryotic organism. In general, in organisms that reproduce sexually, an embryo develops from a zygote, the single cell resulting from the fertilization of the female egg cell by the male sperm cell. The zygote possesses half the DNA from each of its two parents. In plants, animals, and some protists, the zygote will begin to divide by mitosis to produce a multicellular organism. The result of this process is an embryo.

The hindbrain or rhombencephalon 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.

The pons is part of the brainstem, and in humans and other bipeds lies inferior to the midbrain, superior to the medulla oblongata and anterior to the cerebellum.

Embryology

The metencephalon develops from the higher/rostral half of the embryonic rhombencephalon, and is differentiated from the myelencephalon in the embryo by approximately 5 weeks of age. By the third month, the metencephalon differentiates into its two main structures, the pons and the cerebellum.

Myelencephalon posterior region of the embryonic hindbrain

The myelencephalon or afterbrain is the most posterior region of the embryonic hindbrain, from which the medulla oblongata develops.

Functions

The pons regulates breathing through particular nuclei that regulate the breathing center of the medulla oblongata. The cerebellum works to coordinate muscle movements, maintain posture, and integrate sensory information from the inner ear and proprioceptors in the muscles and joints.

The medulla oblongata is a long stem-like structure located in the brainstem. It is anterior and partially inferior to the cerebellum. It is a cone-shaped neuronal mass responsible for autonomic (involuntary) functions ranging from vomiting to sneezing. The medulla contains the cardiac, respiratory, vomiting and vasomotor centers and therefore deals with the autonomic functions of breathing, heart rate and blood pressure.

Cerebellum region of the brain that coordinates motor functions and muscle tone

The cerebellum is a major feature of the hindbrain of all vertebrates. Although usually smaller than the cerebrum, in some animals such as the mormyrid fishes it may be as large as or even larger. In humans, the cerebellum plays an important role in motor control. It may also be involved in some cognitive functions such as attention and language as well as in regulating fear and pleasure responses, but its movement-related functions are the most solidly established. The human cerebellum does not initiate movement, but contributes to coordination, precision, and accurate timing: it receives input from sensory systems of the spinal cord and from other parts of the brain, and integrates these inputs to fine-tune motor activity. Cerebellar damage produces disorders in fine movement, equilibrium, posture, and motor learning in humans.

The inner ear is the innermost part of the vertebrate ear. In vertebrates, the inner ear is mainly responsible for sound detection and balance. In mammals, it consists of the bony labyrinth, a hollow cavity in the temporal bone of the skull with a system of passages comprising two main functional parts:

Development

At the early stages of brain development, the brain vesicles that are formed are imperative.^[1] Each brain region is characterized by its own specific architecture. These regions of the brain are determined by a combination of transcription factors and the signals that change their expression.^[1]

The isthmus is the main organizing center for the tectum and the cerebellum.^[2] The tectum is the dorsal part of the metencephalon. The tectum includes the superior and inferior colliculli, which play a part in visual and audio processing. Two of the major genes that affect the metencephalon are Fgf8 and Wnt1, which are both expressed around the isthmus. Fgf8 is also known as Fibroblast Growth Factor 8. It is a protein that is widely thought to be the most important organizing signal. Its main function is to set up and maintain the barrier between the midbrain and hindbrain, specifically between the mesencephalon and metencephalon.^[2] It also plays a large role in deciding the structure of the mid- and hindbrain. Wnt1 is a proto-oncogene protein (Wingless-type MMTV integration site family, member 1). This gene was originally thought to play a role in the development of the midbrain and hindbrain, but studies have shown that this may not be the case.^[2] Wnt1 is thought to be behind the genetic disorder called Joubert Syndrome, a disorder that affects the cerebellum.

Otx1 and Otx2 are genes that play important parts in the development of the brain, and studies have shown that their roles change throughout the brain’s development.^[3] It is thought that at the stage of brain development wherein the rostral brain is regionalized into its different parts (telencephalon, diencephalon, metencephalon, and mesencephalon) that Otx2 and Otx1 protect the caudalization of the diencephalon and mesencephalon into metencephalon.^[3]

Related Research Articles

The central nervous system (CNS) is the part of the nervous system consisting of the brain and spinal cord. The CNS is so named because it integrates the received information and coordinates and influences the activity of all parts of the bodies of bilaterally symmetric animals—that is, all multicellular animals except sponges and radially symmetric animals such as jellyfish—and it contains the majority of the nervous system. Many consider the retina and the optic nerve, as well as the olfactory nerves and olfactory epithelium as parts of the CNS, synapsing directly on brain tissue without intermediate ganglia. As such, the olfactory epithelium is the only central nervous tissue in direct contact with the environment, which opens up for therapeutic treatments. The CNS is contained within the dorsal body cavity, with the brain housed in the cranial cavity and the spinal cord in the spinal canal. In vertebrates, the brain is protected by the skull, while the spinal cord is protected by the vertebrae. The brain and spinal cord are both enclosed in the meninges. Within the CNS, the interneuronal space is filled with a large amount of supporting non-nervous cells called neuroglial cells.

The thalamus is a large mass of gray matter in the dorsal part of the diencephalon of the brain with several functions such as relaying of sensory signals, including motor signals to the cerebral cortex, and the regulation of consciousness, sleep, and alertness.

In the developing chordate, 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, and ultimately the folds meet and coalesce in the middle line and convert the groove into the closed neural tube. In humans, neural tube closure usually occurs by the fourth week of pregnancy. The ectodermal wall of the tube forms the rudiment of the nervous system. The centre of the tube is the neural canal.

The brainstem is the posterior part of the brain, continuous with the spinal cord. In the human brain the brainstem includes the midbrain, and the pons and medulla oblongata of the hindbrain. Sometimes the diencephalon, the caudal part of the forebrain, is included.

Forebrain rostral-most (forward-most) portion of the brain

In the anatomy of the brain of vertebrates, the forebrain or prosencephalon is the rostral (forward-most) portion of the brain. The forebrain (prosencephalon), the midbrain (mesencephalon), and hindbrain (rhombencephalon) are the three primary brain vesicles during the early development of the nervous system. The forebrain controls body temperature, reproductive functions, eating, sleeping, and the display of emotions.

The midbrain or mesencephalon is a portion of the central nervous system associated with vision, hearing, motor control, sleep/wake, arousal (alertness), and temperature regulation.

The diencephalon is a division of the forebrain, and is situated between the telencephalon and the midbrain. It consists of structures that are on either side of the third ventricle, including the thalamus, the hypothalamus, the epithalamus and the subthalamus.

Neurulation refers to the folding process in vertebrate embryos, which includes the transformation of the neural plate into the neural tube. The embryo at this stage is termed the neurula.

The tegmentum 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 basal or ventral structures at each level. It forms the floor of the midbrain (mesencephalon) whereas the tectum forms the ceiling. It is a multisynaptic network of neurons that is involved in many subconscious homeostatic and reflexive pathways. It is a motor center that relays inhibitory signals to the thalamus and basal nuclei preventing unwanted body movement. The tegmentum area includes various different structures, such as the "rostral (=frontal/cranial/oral) end of the reticular formation, several nuclei controlling eye movements, the periaqueductal gray matter, the red nucleus, the substantia nigra, and the ventral tegmental area".

The zona limitans intrathalamica (ZLI) is a lineage-restriction compartment and primary developmental boundary in the vertebrate forebrain that serves as a signaling center and a restrictive border between the thalamus and the prethalamus.

Paired box gene 2, also known as PAX2 is a protein which in humans is encoded by the PAX2 gene.

Fibroblast growth factor 8 is a protein that in humans is encoded by the FGF8 gene.

The rhombic lip is a posterior section of the developing metencephalon which can be recognized transiently within the vertebrate embryo. It extends posteriorly from the roof of the fourth ventricle to dorsal neuroepithelial cells. The rhombic lip can be divided into eight structural units based on rhombomeres 1-8 (r1-r8), which can be recognized at early stages of hindbrain development. Producing granule cells and five brainstem nuclei, the rhombic lip plays an important role in developing a complex cerebellar neural system.

Brain vesicles are the bulge-like features of the early development of the neural tube in vertebrates. Vesicle formation begins shortly after anterior neural tube closure at about embryonic day 9.0 in the mouse and the fourth and fifth gestational week in human development. In zebrafish and chicken embryos, brain vesicles form by about 24 hours and 48 hours post-conception, respectively. Initially there are three primary brain vesicles: prosencephalon, mesencephalon, and rhombencephalon. These develop into five secondary brain vesicles – the prosencephalon is subdivided into the telencephalon and diencephalon, and the rhombencephalon into the metencephalon and myelencephalon. During these early vesicle stages, the walls of the neural tube contain neural stem cells in a region called the neuroepithelium or ventricular zone. These neural stem cells divide rapidly, driving growth of the early brain, but later, these stem cells begin to generate neurons through the process of neurogenesis.

The isthmic organizer, or isthmus organizer, also known as the midbrain−hindbrain boundary (MHB), is a secondary organizer region that develops at the junction of the midbrain and metencephalon. The MHB expresses signaling molecules that regulate the differentiation and patterning of the adjacent neuroepithelium. This allows for the development of the midbrain and hindbrain as well as the specification of neuronal subtypes in these regions. The fact that the MHB is sufficient for the development of the mid and hindbrain was shown in an experiment where quail MHB cells transplanted into the forebrain of a chick were able to induce an ectopic midbrain and cerebellum.

Three flexures form in the part of the embryonic neural tube that develops into the brain. At four weeks gestational age in the human embryo the neural tube has developed at the cranial end into three swellings – the primary brain vesicles. The space into which the cranial part of the neural tube is developing is limited. This limitation causes the neural tube to bend, or flex, at two ventral flexures – the rostral cephalic flexure, and the caudal cervical flexure. It also bends dorsally into the pontine flexure. These flexures have formed by the time that the primary brain vesicles have developed into five secondary brain vesicles in the fifth week.

References

1 2 Nakamura, H., and Watanabe, Y. Isthmus organizer and regionalization of the mesencephalon and metencephalon. Int. J. Dev. Biol. 49: 231-235 (2005). doi: 10.1387/ijdb.041964hn.
1 2 3 Matsunaga, E., Katahira, T., and Nakamura, H. "Role of Lmx1b and Wnt1 in mesencephalon and metencephalon development""Development" (2002)
1 2 Sakuri, Y., Kurokawa, D., Kiyonari, H., Kajikawa, E., Suda, Y., and Aizawa, S. "Otx2 and Otx1 protect diencephalon and mesencephalon from caudalization into metencephalon during early brain regionalization" ^{[ permanent dead link ]} "Developmental Biology" (2010).

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[Nakamura-1] 1 2 Nakamura, H., and Watanabe, Y. Isthmus organizer and regionalization of the mesencephalon and metencephalon. Int. J. Dev. Biol. 49: 231-235 (2005). doi: 10.1387/ijdb.041964hn.

[Matsunaga-2] 1 2 3 Matsunaga, E., Katahira, T., and Nakamura, H. "Role of Lmx1b and Wnt1 in mesencephalon and metencephalon development""Development" (2002)

[Sakuri-3] 1 2 Sakuri, Y., Kurokawa, D., Kiyonari, H., Kajikawa, E., Suda, Y., and Aizawa, S. "Otx2 and Otx1 protect diencephalon and mesencephalon from caudalization into metencephalon during early brain regionalization" ^{[ permanent dead link ]} "Developmental Biology" (2010).