In the developing

chordate A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fi ...

(including

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, wi ...

s), the neural tube is the

embryo 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 ...

nic precursor to the

central nervous system The central nervous system (CNS) is the part of the nervous system consisting primarily of the brain and spinal cord. The CNS is so named because the brain integrates the received information and coordinates and influences the activity of all pa ...

, which is made up 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 ...

and

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 spi ...

. The neural groove gradually deepens as the

neural fold The neural fold is a structure that arises during neurulation in the embryonic development of both birds and mammals among other organisms. This structure is associated with primary neurulation, meaning that it forms by the coming together of tis ...

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 28th day after conception). The

ectoderm The ectoderm is one of the three primary germ layers formed in early embryonic development. It is the outermost layer, and is superficial to the mesoderm (the middle layer) and endoderm (the innermost layer). It emerges and originates from th ...

al wall of the tube forms the

rudiment In ''rudimental drumming'', a form of percussion music, a drum rudiment is one of a number of relatively small patterns which form the foundation for more extended and complex drumming patterns. The term "drum rudiment" is most closely associ ...

of the nervous system. The centre of the tube is the ''neural canal''.It is an important structure for the development of fetus's brain and spine Neural crest

Development

The neural tube develops in two ways: primary neurulation and secondary neurulation. Primary neurulation divides the

into three cell types: * The internally located neural tube * The externally located epidermis * The

neural crest Neural crest cells are a temporary group of cells unique to vertebrates that arise from the embryonic ectoderm germ layer, and in turn give rise to a diverse cell lineage—including melanocytes, craniofacial cartilage and bone, smooth muscle, pe ...

cells, which develop in the region between the neural tube and epidermis but then migrate to new locations # Primary neurulation begins after the neural plate forms. The edges of the neural plate start to thicken and lift upward, forming the neural folds. The center of the neural plate remains grounded, allowing a U-shaped neural groove to form. This neural groove sets the boundary between the right and left sides of the embryo. The neural folds pinch in towards the midline of the embryo and fuse together to form the neural tube.Gilbert, Scott F. Developmental Biology Eighth Edition. Sunderland, Massachusetts: Sinauer Associates, Inc., 2006. # In secondary neurulation, the cells of the neural plate form a cord-like structure that migrates inside the embryo and hollows to form the tube. Each organism uses primary and secondary neurulation to varying degrees. * Neurulation in ''

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 liv ...

'' proceeds only via the secondary form. * In ''

avian Avian may refer to: *Birds or Aves, winged animals * Avian (given name) (russian: Авиа́н, link=no), a male forename Aviation * Avro Avian, a series of light aircraft made by Avro in the 1920s and 1930s * Avian Limited, a hang glider manufact ...

'' species the posterior regions of the tube develop using secondary neurulation and the anterior regions develop by primary neurulation. * In ''

mammals Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), f ...

'', secondary neurulation begins around the 35th somite. Mammalian neural tubes close in the head in the opposite order that they close in the trunk. * In the head: :#Neural crest cells migrate :#Neural tube closes :#Overlying ectoderm closes * In the trunk: :#Overlying ectoderm closes :#Neural tube closes :#Neural crest cells migrate

Structure

Four neural tube subdivisions each eventually develop into distinct regions of the central nervous system by the division of neuroepithelial cells: 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 (prosencephalon), the midbrain (mesencephalon), and hindbrain (rhombencephalon) are the three primar ...

(prosencephalon), the

midbrain 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'', " ...

(mesencephalon), 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 ...

(rhombencephalon) and the

. * The ''prosencephalon'' further goes on to develop into the telencephalon (

cerebrum 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

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 o ...

(the optic vesicles and

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 hypothalamus i ...

). * The ''mesencephalon'' stays as the midbrain. * The ''rhombencephalon'' develops into the

metencephalon 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 port ...

(the

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 V ...

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 cer ...

) and the myelencephalon (the

medulla oblongata 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 neuronal mass responsible for autonomic (involun ...

). For a short time, the neural tube is open both

cranially 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 pr ...

and caudally. These openings, called

neuropore 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 process begins when the notochord induces the for ...

s, close during the fourth week in humans. Improper closure of the neuropores can result in neural tube defects such as

anencephaly Anencephaly is the absence of a major portion of the brain, skull, and scalp that occurs during embryonic development. It is a cephalic disorder that results from a neural tube defect that occurs when the rostral (head) end of the neural tube ...

spina bifida Spina bifida (Latin for 'split spine'; SB) is a birth defect in which there is incomplete closing of the spine and the membranes around the spinal cord during early development in pregnancy. There are three main types: spina bifida occulta, ...

. The

dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage * Dorsal ...

part of the neural tube contains the alar plate, which is associated primarily with sensation. The

ventral 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 pr ...

part of the neural tube contains the basal plate, which is primarily associated with motor (i.e.,

muscle Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of m ...

) control.

Dorsal-ventral patterning

The neural tube patterns along the dorsal-ventral axis to establish defined compartments of neural progenitor cells that lead to distinct classes of neurons. According to the French flag model of

morphogenesis Morphogenesis (from the Greek ''morphê'' shape and ''genesis'' creation, literally "the generation of form") is the biological process that causes a cell, tissue or organism to develop its shape. It is one of three fundamental aspects of dev ...

, this patterning occurs early in development and results from the activity of several secreted signaling molecules.

Sonic hedgehog Sonic hedgehog protein (SHH) is encoded for by the ''SHH'' gene. The protein is named after the character ''Sonic the Hedgehog''. This signaling molecule is key in regulating embryonic morphogenesis in all animals. SHH controls organogenesis an ...

(Shh) is a key player in patterning the ventral axis, while bone morphogenic proteins (BMPs) and Wnt family members play an important role in patterning the dorsal axis. Other factors shown to provide positional information to the neural progenitor cells include fibroblast growth factors (FGFs) and

retinoic acid Retinoic acid (used simplified here for all-''trans''-retinoic acid) is a metabolite of vitamin A1 (all-''trans''- retinol) that mediates the functions of vitamin A1 required for growth and development. All-''trans''-retinoic acid is required in ...

. Retinoic acid is required ventrally along with Shh to induce Pax6 and Olig2 during differentiation of motor neurons. Three main ventral cell types are established during early neural tube development: the ''

floor plate The floor plate is a structure integral to the developing nervous system of vertebrate organisms. Located on the ventral midline of the embryonic neural tube, the floor plate is a specialized glial structure that spans the anteroposterior axis ...

cells'', which form at the ventral midline during the neural fold stage; as well as the more dorsally located

motor neuron A motor neuron (or motoneuron or efferent neuron) is a neuron whose cell body is located in the motor cortex, brainstem or the spinal cord, and whose axon (fiber) projects to the spinal cord or outside of the spinal cord to directly or indirectly ...

s and

interneuron Interneurons (also called internuncial neurons, relay neurons, association neurons, connector neurons, intermediate neurons or local circuit neurons) are neurons that connect two brain regions, i.e. not direct motor neurons or sensory neurons. ...

s. These cell types are specified by the secretion of the Shh from the notochord (located ventrally to the neural tube), and later from the floor plate cells. Shh acts as a morphogen, meaning that it acts in a concentration-dependent manner to specify cell types as it moves further from its source. The following is a proposed mechanism for how Shh patterns the ventral neural tube: A gradient of Shh that controls the expression of a group of homeodomain (HD) and basic Helix-Loop-Helix (bHLH) transcription factors is created. These transcription factors are grouped into two protein classes based on how Shh affects them. Class I is inhibited by Shh, whereas Class II is activated by Shh. These two classes of proteins then cross-regulate each other to create more defined boundaries of expression. The different combinations of expression of these transcription factors along the dorsal-ventral axis of the neural tube are responsible for creating the identity of the neuronal progenitor cells. Five molecularly distinct groups of ventral neurons form from these neuronal progenitor cells in vitro. Also, the position at which these neuronal groups are generated ''in vivo'' can be predicted by the concentration of Shh required for their induction in vitro. Studies have shown that neural progenitors can evoke different responses based on the length of exposure to Shh, with a longer exposure time resulting in more ventral cell types. At the dorsal end of the neural tube, BMPs are responsible for neuronal patterning. BMP is initially secreted from the overlying ectoderm. A secondary signaling center is then established in the roof plate, the dorsal most structure of the neural tube. BMP from the dorsal end of the neural tube seems to act in the same concentration-dependent manner as Shh in the ventral end. This was shown using zebrafish mutants that had varying amounts of BMP signaling activity. Researchers observed changes in dorsal-ventral patterning, for example, zebrafish deficient in certain BMPs showed a loss of dorsal sensory neurons and an expansion of interneurons. Shh gradient in the neural tube

References

External links

* *
Diagram at embryology.med.unsw.edu.au

{{DEFAULTSORT:Neural Tube Embryology of nervous system Neurulation Ectoderm

Development

Structure

Dorsal-ventral patterning

See also

References

External links