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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
neuron A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa. ...
al mass responsible for autonomic (involuntary) functions, ranging from
vomiting Vomiting (also known as emesis and throwing up) is the involuntary, forceful expulsion of the contents of one's stomach through the mouth and sometimes the nose. Vomiting can be the result of ailments like food poisoning, gastroenteri ...
to
sneezing A sneeze (also known as sternutation) is a semi-autonomous, convulsive expulsion of air from the lungs through the nose and mouth, usually caused by foreign particles irritating the nasal mucosa. A sneeze expels air forcibly from the mouth and ...
. The medulla contains the
cardiac The heart is a muscular organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as carbon dioxide to t ...
,
respiratory The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies gre ...
,
vomiting Vomiting (also known as emesis and throwing up) is the involuntary, forceful expulsion of the contents of one's stomach through the mouth and sometimes the nose. Vomiting can be the result of ailments like food poisoning, gastroenteri ...
and
vasomotor Vasomotor refers to actions upon a blood vessel which alter its diameter. More specifically, it can refer to vasodilator action and vasoconstrictor action. Control Sympathetic innervation Sympathetic nerve fibers travel around the tunica media of ...
centers, and therefore deals with the autonomic functions of breathing,
heart rate Heart rate (or pulse rate) is the frequency of the heartbeat measured by the number of contractions (beats) of the heart per minute (bpm). The heart rate can vary according to the body's physical needs, including the need to absorb oxygen and excr ...
and blood pressure as well as the
sleep–wake cycle A circadian rhythm (), or circadian cycle, is a natural, internal process that regulates the sleep–wake cycle and repeats roughly every 24 hours. It can refer to any process that originates within an organism (i.e., endogenous) and responds to ...
. During embryonic development, the medulla oblongata develops from the myelencephalon. The myelencephalon is a secondary vesicle which forms during the maturation of the
rhombencephalon 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. Metencephal ...
, also referred to as the hindbrain. The bulb is an archaic term for the medulla oblongata. In modern clinical usage, the word bulbar (as in
bulbar palsy Bulbar palsy refers to a range of different signs and symptoms linked to impairment of function of the glossopharyngeal nerve (CN IX), the vagus nerve (CN X), the accessory nerve (CN XI), and the hypoglossal nerve (CN XII). It is caused by a low ...
) is retained for terms that relate to the medulla oblongata, particularly in reference to
medical conditions A disease is a particular abnormal condition that negatively affects the structure or function of all or part of an organism, and that is not immediately due to any external injury. Diseases are often known to be medical conditions that a ...
. The word bulbar can refer to the nerves and tracts connected to the medulla, and also by association to those muscles
innervated 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 ...
, such as those of the
tongue The tongue is a muscular organ in the mouth of a typical tetrapod. It manipulates food for mastication and swallowing as part of the digestive process, and is the primary organ of taste. The tongue's upper surface (dorsum) is covered by taste ...
,
pharynx The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its st ...
and larynx.


Anatomy

The medulla can be thought of as being in two parts: * an upper open part or superior part where the dorsal surface of the medulla is formed by 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 ...
. * a lower closed part or inferior part where the fourth ventricle has narrowed at the
obex OBEX (abbreviation of OBject EXchange, also termed IrOBEX) is a communications protocol that facilitates the exchange of binary objects between devices. It is maintained by the Infrared Data Association but has also been adopted by the Bluetooth S ...
in the caudal medulla, and surrounds part of the
central canal The central canal (also known as spinal foramen or ependymal canal) is the cerebrospinal fluid-filled space that runs through the spinal cord. The central canal lies below and is connected to the ventricular system of the brain, from which it r ...
.


External surfaces

The anterior median fissure contains a fold of
pia mater Pia mater ( or ),Entry "pia mater"
in
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 ...
in a small triangular area, termed the foramen cecum. On either side of this fissure are raised areas termed the medullary pyramids. The pyramids house the
pyramidal tracts The pyramidal tracts include both the corticobulbar tract and the corticospinal tract. These are aggregations of efferent nerve fibers from the upper motor neurons that travel from the cerebral cortex and terminate either in the brainstem (''cort ...
–the corticospinal and the
corticobulbar tract 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 ...
s of the nervous system. At the caudal part of the medulla these tracts cross over in the
decussation of the pyramids In neuroanatomy, the medullary pyramids are paired white matter structures of the brainstem's medulla oblongata that contain motor fibers of the corticospinal and corticobulbar tracts – known together as the pyramidal tracts. The lower limit ...
obscuring the fissure at this point. Some other fibers that originate from the anterior median fissure above the decussation of the pyramids and run laterally across the surface of the pons are known as the
anterior external arcuate fibers 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 ...
. The region between the anterolateral and posterolateral sulcus in the upper part of the medulla is marked by a pair of swellings known as olivary bodies (also called ''olives''). They are caused by the largest nuclei of the olivary bodies, the inferior olivary nuclei. The posterior part of the medulla between the posterior median sulcus and the posterolateral sulcus contains tracts that enter it from the posterior funiculus of the spinal cord. These are the gracile fasciculus, lying medially next to the midline, and the cuneate fasciculus, lying laterally. These fasciculi end in rounded elevations known as the gracile and the cuneate tubercles. They are caused by masses of
gray matter Grey matter is a major component of the central nervous system, consisting of neuronal cell bodies, neuropil (dendrites and unmyelinated axons), glial cells (astrocytes and oligodendrocytes), synapses, and capillaries. Grey matter is distingui ...
known as the gracile nucleus and the
cuneate nucleus In neuroanatomy, the dorsal column nuclei are a pair of nuclei in the dorsal columns in the brainstem. The name refers collectively to the cuneate nucleus and gracile nucleus, which are present at the bottom of the medulla oblongata. Both nucl ...
. The soma (cell bodies) in these nuclei are the second-order neurons of the posterior column-medial lemniscus pathway, and their axons, called the
internal arcuate fibers In neuroanatomy, the internal arcuate fibers or internal arcuate tract are the axons of second-order sensory neurons that compose the gracile and cuneate nuclei of the medulla oblongata. These second-order neurons begin in the gracile and cunea ...
or fasciculi, decussate from one side of the medulla to the other to form 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 ...
. Just above the tubercles, the posterior aspect of the medulla is occupied by a triangular fossa, which forms the lower part of the floor of 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 ...
. The fossa is bounded on either side by the
inferior cerebellar peduncle The upper part of the posterior district of the medulla oblongata is occupied by the inferior cerebellar peduncle, a thick rope-like strand situated between the lower part of the fourth ventricle and the roots of the glossopharyngeal and vagus nerv ...
, which connects the medulla to the cerebellum. The lower part of the medulla, immediately lateral to the cuneate fasciculus, is marked by another longitudinal elevation known as the tuberculum cinereum. It is caused by an underlying collection of gray matter known as the spinal trigeminal nucleus. The gray matter of this nucleus is covered by a layer of nerve fibers that form the spinal tract of the
trigeminal nerve In neuroanatomy, the trigeminal nerve ( lit. ''triplet'' nerve), also known as the fifth cranial nerve, cranial nerve V, or simply CN V, is a cranial nerve responsible for sensation in the face and motor functions such as biting and chew ...
. The base of the medulla is defined by the commissural fibers, crossing over from the ipsilateral side in the spinal cord to the contralateral side in the brain stem; below this is the spinal cord.


Blood supply

Blood to the medulla is supplied by a number of
arteries An artery (plural arteries) () is a blood vessel in humans and most animals that takes blood away from the heart to one or more parts of the body (tissues, lungs, brain etc.). Most arteries carry oxygenated blood; the two exceptions are the pu ...
. * Anterior spinal artery: This supplies the whole medial part of the medulla oblongata. * Posterior inferior cerebellar artery: This is a major branch of the vertebral artery, and supplies the posterolateral part of the medulla, where the main sensory tracts run and synapse. It also supplies part of the cerebellum. * Direct branches of the vertebral artery: The vertebral artery supplies an area between the anterior spinal and posterior inferior cerebellar arteries, including the solitary nucleus and other sensory nuclei and fibers. *
Posterior spinal artery The posterior spinal artery (dorsal spinal arteries) arises from the vertebral artery in 25% of humans or the posterior inferior cerebellar artery in 75% of humans, adjacent to the medulla oblongata. It supplies the grey and white posterior column ...
: This supplies the dorsal column of the closed medulla containing fasciculus gracilis, gracile nucleus, fasciculus cuneatus, and
cuneate nucleus In neuroanatomy, the dorsal column nuclei are a pair of nuclei in the dorsal columns in the brainstem. The name refers collectively to the cuneate nucleus and gracile nucleus, which are present at the bottom of the medulla oblongata. Both nucl ...
.


Development

The medulla oblongata forms in
fetal 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 ...
from the myelencephalon. The final differentiation of the medulla is seen at week 20 gestation. Neuroblasts from the alar plate 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, ...
at this level will produce the sensory nuclei of the medulla. The basal plate neuroblasts will give rise to the motor nuclei. * Alar plate neuroblasts give rise to: ** The solitary nucleus, which contains the
general visceral afferent fibers A general officer is an officer of high rank in the armies, and in some nations' air forces, space forces, and marines or naval infantry. In some usages the term "general officer" refers to a rank above colonel."general, adj. and n.". OED O ...
for
taste The gustatory system or sense of taste is the sensory system that is partially responsible for the perception of taste (flavor). Taste is the perception produced or stimulated when a substance in the mouth reacts chemically with taste receptor ...
, as well as the special visceral afferent column. ** The spinal trigeminal nerve nuclei which contains the general somatic afferent column. ** The cochlear and
vestibular nuclei The vestibular nuclei (VN) are the cranial nuclei for the vestibular nerve located in the brainstem. In Terminologia Anatomica they are grouped in both the pons and the medulla in the brainstem. Structure Path The fibers of the vestibular n ...
, which contain the
special somatic afferent Special somatic afferent fibers (SSA) are the afferent nerve fibers that carry information from the special senses of vision, hearing and balance. The cranial nerves containing SSA fibers are the optic nerve (II) and the vestibulocochlear nerve (VI ...
column. ** The
inferior olivary nucleus The inferior olivary nucleus (ION), is a structure found in the medulla oblongata underneath the superior olivary nucleus.Gado, Thomas A. Woolsey; Joseph Hanaway; Mokhtar H. (2003). The brain atlas a visual guide to the human central nervous syst ...
, which relays to the cerebellum. ** The
dorsal column nuclei In neuroanatomy, the dorsal column nuclei are a pair of nuclei in the dorsal columns in the brainstem. The name refers collectively to the cuneate nucleus and gracile nucleus, which are present at the bottom of the medulla oblongata. Both nucl ...
, which contain the gracile and cuneate nuclei. * Basal plate neuroblasts give rise to: ** The
hypoglossal nucleus The hypoglossal nucleus is a cranial nerve nucleus, found within the medulla. Being a motor nucleus, it is close to the midline. In the open medulla, it is visible as what is known as the '' hypoglossal trigone'', a raised area (medial to the v ...
, which contains general somatic efferent fibers. ** The
nucleus ambiguus The nucleus ambiguus ("ambiguous nucleus" in English) is a group of large motor neurons, situated deep in the medullary reticular formation named by Jacob Clarke. The nucleus ambiguus contains the cell bodies of neurons that innervate the muscle ...
, which form the
special visceral efferent Special visceral efferent fibers (SVE) are the efferent nerve fibers that provide motor innervation to the muscles of the pharyngeal arches in humans, and the branchial arches in fish. Some sources prefer the term "branchiomotor" or "branchial eff ...
. ** The
dorsal nucleus of vagus nerve The dorsal nucleus of vagus nerve (or posterior nucleus of vagus nerve or dorsal vagal nucleus or nucleus dorsalis nervi vagi or nucleus posterior nervi vagi) is a cranial nerve nucleus for the vagus nerve in the medulla that lies ventral to the f ...
and the inferior salivatory nucleus, both of which form the
general visceral efferent fibers General visceral efferent fibers (GVE) or visceral efferents or autonomic efferents, are the efferent nerve fibers of the autonomic nervous system (also known as the ''visceral efferent nervous system'' that provide motor innervation to smooth mu ...
.


Function

The medulla oblongata connects the higher levels of the
brain A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a ve ...
to the
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 sp ...
, and is responsible for several functions of the
autonomous nervous system The autonomic nervous system (ANS), formerly referred to as the vegetative nervous system, is a division of the peripheral nervous system that supplies internal organs, smooth muscle and glands. The autonomic nervous system is a control system t ...
which include: * The
control of ventilation The control of ventilation refers to the physiological mechanisms involved in the control of breathing, which is the movement of air into and out of the lungs. Ventilation facilitates respiration. Respiration refers to the utilization of oxygen and ...
via signals from the carotid and aortic bodies.
Respiration Respiration may refer to: Biology * Cellular respiration, the process in which nutrients are converted into useful energy in a cell ** Anaerobic respiration, cellular respiration without oxygen ** Maintenance respiration, the amount of cellul ...
is regulated by groups of
chemoreceptor A chemoreceptor, also known as chemosensor, is a specialized sensory receptor which transduces a chemical substance (endogenous or induced) to generate a biological signal. This signal may be in the form of an action potential, if the chemorecept ...
s. These sensors detect changes in the acidity of the blood; if, for example, the blood becomes too acidic, the medulla oblongata sends electrical signals to intercostal and phrenical muscle tissue to increase their contraction rate and increase oxygenation of the
blood Blood is a body fluid in the circulatory system of humans and other vertebrates that delivers necessary substances such as nutrients and oxygen to the cells, and transports metabolic waste products away from those same cells. Blood in the cir ...
. The
ventral respiratory group The respiratory center is located in the medulla oblongata and pons, in the brainstem. The respiratory center is made up of three major respiratory groups of neurons, two in the medulla and one in the pons. In the medulla they are the dorsal res ...
and the
dorsal respiratory group The respiratory center is located in the medulla oblongata and pons, in the brainstem. The respiratory center is made up of three major respiratory groups of neurons, two in the medulla and one in the pons. In the medulla they are the dorsal res ...
are neurons involved in this regulation. The pre-Bötzinger complex is a cluster of interneurons involved in the respiratory function of the medulla. * Cardiovascular center – sympathetic, parasympathetic nervous system *
Vasomotor center The vasomotor center (VMC) is a portion of the medulla oblongata. Together with the cardiovascular center and respiratory center, it regulates blood pressure. It also has a more minor role in other homeostatic processes. Upon increase in carbon ...
baroreceptors * Reflex centers of vomiting, coughing, sneezing and swallowing. These reflexes which include the
pharyngeal reflex The pharyngeal reflex or gag reflex is a reflex muscular contraction of the back of the throat, evoked by touching the roof of the mouth, the back of the tongue, the area around the tonsils, the uvula, and the back of the throat. It, along with othe ...
, the swallowing reflex (also known as the palatal reflex), and the masseter reflex can be termed ''bulbar'' reflexes.
/ref>


Clinical significance

A Vascular occlusion, blood vessel blockage (such as in a stroke) will injure the
pyramidal tract The pyramidal tracts include both the corticobulbar tract and the corticospinal tract. These are aggregations of efferent nerve fibers from the upper motor neurons that travel from the cerebral cortex and terminate either in the brainstem (''co ...
,
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 ...
, and the
hypoglossal nucleus The hypoglossal nucleus is a cranial nerve nucleus, found within the medulla. Being a motor nucleus, it is close to the midline. In the open medulla, it is visible as what is known as the '' hypoglossal trigone'', a raised area (medial to the v ...
. This causes a syndrome called
medial medullary syndrome Medial medullary syndrome, also known as inferior alternating syndrome, hypoglossal alternating hemiplegia, lower alternating hemiplegia, or Dejerine syndrome, is a type of alternating hemiplegia characterized by a set of clinical features result ...
.
Lateral medullary syndrome Lateral medullary syndrome is a neurological disorder causing a range of symptoms due to ischemia in the lateral part of the medulla oblongata in the brainstem. The ischemia is a result of a blockage most commonly in the vertebral artery or the p ...
can be caused by the blockage of either the posterior inferior cerebellar artery or of the vertebral arteries. Progressive bulbar palsy (PBP) is a disease that attacks the nerves supplying the bulbar muscles. Infantile progressive bulbar palsy is progressive bulbar palsy in children.


Other animals

Both lampreys and hagfish possess a fully developed medulla oblongata. Since these are both very similar to early
agnathans Agnatha (, Ancient Greek 'without jaws') is an infraphylum of jawless fish in the phylum Chordata, subphylum Vertebrata, consisting of both present (cyclostomes) and extinct (conodonts and ostracoderms) species. Among recent animals, cyclosto ...
, it has been suggested that the medulla evolved in these early fish, approximately 505 million years ago. Haycock, ''Being and Perceiving'' The status of the medulla as part of the primordial reptilian brain is confirmed by its disproportionate size in modern reptiles such as the crocodile,
alligator An alligator is a large reptile in the Crocodilia order in the genus ''Alligator'' of the family Alligatoridae. The two extant species are the American alligator (''A. mississippiensis'') and the Chinese alligator (''A. sinensis''). Additiona ...
, and monitor lizard.


Additional images

File:Cerebrum lobes.svg, Lobes File:Medulla Oblongata and Cerebellum.svg, Cross section of the medulla (in red) and surrounding tissues. File:Gray679.png, Anteroinferior view of the medulla oblongata and pons. File:Gray724.png, Base of brain. File:Gray768.png, Diagram showing the positions of the three principal subarachnoid cisternæ. File:Slide11ee.JPG, Medulla oblongata File:Medulla oblongata - posterior - very low mag.jpg,
Micrograph A micrograph or photomicrograph is a photograph or digital image taken through a microscope or similar device to show a magnified image of an object. This is opposed to a macrograph or photomacrograph, an image which is also taken on a m ...
of the posterior portion of the ''open part'' of the medulla oblongata, showing 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 ...
(top of image) and the nuclei of CN XII (medial) and CN X (lateral). H&E-LFB stain.


References

*


External links

* {{DEFAULTSORT:Medulla Oblongata