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In
anatomy Anatomy () is the branch of biology concerned with the study of the structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old science, having i ...
, the notochord is a flexible rod which is similar in structure to the stiffer
cartilage Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck ...
. If a
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
has a notochord at any stage of its life cycle (along with 4 other features), it is, by definition, a
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 ...
. The notochord consists of inner, vacuolated cells covered by
fibrous Fiber or fibre (from la, fibra, links=no) is a natural or artificial substance that is significantly longer than it is wide. Fibers are often used in the manufacture of other materials. The strongest engineering materials often incorporat ...
and
elastic Elastic is a word often used to describe or identify certain types of elastomer, elastic used in garments or stretchable fabrics. Elastic may also refer to: Alternative name * Rubber band, ring-shaped band of rubber used to hold objects togethe ...
sheaths, lies along the anteroposterior axis (''front to back''), is usually closer to the dorsal than the ventral surface of 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 spe ...
, and is composed of cells derived from the
mesoderm The mesoderm is the middle layer of the three germ layers that develops during gastrulation in the very early development of the embryo of most animals. The outer layer is the ectoderm, and the inner layer is the endoderm.Langman's Medical Emb ...
. The most commonly cited functions of the notochord are: as a midline tissue that provides directional signals to surrounding tissue during development, as a skeletal (structural) element, and as a vertebral precursor. In lancelets the notochord persists throughout life as the main structural support of the body. In
tunicate A tunicate is a marine invertebrate animal, a member of the subphylum Tunicata (). It is part of the Chordata, a phylum which includes all animals with dorsal nerve cords and notochords (including vertebrates). The subphylum was at one t ...
s the notochord is present only in the larval stage, being completely absent in the adult animal. In these invertebrate chordates, the notochord is not vacuolated. In all
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, with ...
s other than the hagfish, the notochord is integrated into the
vertebral column The vertebral column, also known as the backbone or spine, is part of the axial skeleton. The vertebral column is the defining characteristic of a vertebrate in which the notochord (a flexible rod of uniform composition) found in all chordate ...
, with its original structure being (nearly) retained in the
intervertebral disc An intervertebral disc (or intervertebral fibrocartilage) lies between adjacent vertebrae in the vertebral column. Each disc forms a fibrocartilaginous joint (a symphysis), to allow slight movement of the vertebrae, to act as a ligament to h ...
s as the nucleus pulposus.


Structure

The notochord is a long, rodlike structure that develops dorsal to the gut and ventral to the neural tube. The notochord is composed primarily of a core of glycoproteins, encased in a sheath of collagen fibers wound into two opposing helices. The glycoproteins are stored in vacuolated, turgid cells. The angle between these fibers determines whether increased pressure in the core will result in shortening and thickening versus lengthening and thinning. Contraction of these muscle fibers result in a side-to-side motion resembling swimming. The stiffened notochord prevents movement through telescoping motion, such as that of an
earthworm An earthworm is a terrestrial invertebrate that belongs to the phylum Annelida. They exhibit a tube-within-a-tube body plan; they are externally segmented with corresponding internal segmentation; and they usually have setae on all segments. T ...
.


Role in signaling and development

The notochord plays a key role in signaling and coordinating development.
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 spe ...
s of modern vertebrates form transient notochord structures during
gastrulation Gastrulation is the stage in the early embryonic development of most animals, during which the blastula (a single-layered hollow sphere of cells), or in mammals the blastocyst is reorganized into a multilayered structure known as the gastrula. ...
. The notochord is found ventral to the neural tube. Notogenesis is the development of the notochord by
epiblast In amniote embryonic development, the epiblast (also known as the primitive ectoderm) is one of two distinct cell layers arising from the inner cell mass in the mammalian blastocyst, or from the blastula in reptiles and birds, the other layer is ...
s that form the floor of the amnion cavity. The progenitor notochord is derived from cells migrating from the primitive node and pit. The notochord forms during
gastrulation Gastrulation is the stage in the early embryonic development of most animals, during which the blastula (a single-layered hollow sphere of cells), or in mammals the blastocyst is reorganized into a multilayered structure known as the gastrula. ...
and soon after induces the formation of the neural plate ( neurulation), synchronizing the development of the neural tube. On the ventral aspect of the neural groove an axial thickening of the
endoderm Endoderm is the innermost of the three primary germ layers in the very early embryo. The other two layers are the ectoderm (outside layer) and mesoderm (middle layer). Cells migrating inward along the archenteron form the inner layer of the gast ...
takes place. (In bipedal chordates, e.g. humans, this surface is properly referred to as the anterior surface). This thickening appears as a furrow (the chordal furrow) the margins of which anastomose (come into contact), and so convert it into a solid rod of polygonal-shaped cells (the notochord) which is then separated from the endoderm. In vertebrates, it extends throughout the entire length of the future vertebral column, and reaches as far as the anterior end of 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'', ...
, where it ends in a hook-like extremity in the region of the future
dorsum sellae The dorsum sellae is part of the sphenoid bone in the skull. Together with the basilar part of the occipital bone it forms the clivus. In the sphenoid bone, the anterior boundary of the sella turcica is completed by two small eminences, one o ...
of the
sphenoid bone The sphenoid bone is an unpaired bone of the neurocranium. It is situated in the middle of the skull towards the front, in front of the basilar part of the occipital bone. The sphenoid bone is one of the seven bones that articulate to form the o ...
. Initially, it exists between the neural tube and the endoderm of the yolk-sac; soon, the notochord becomes separated from them by the
mesoderm The mesoderm is the middle layer of the three germ layers that develops during gastrulation in the very early development of the embryo of most animals. The outer layer is the ectoderm, and the inner layer is the endoderm.Langman's Medical Emb ...
, which grows medially and surrounds it. From the mesoderm surrounding the neural tube and notochord, the
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
, vertebral column, and the membranes of the
brain A brain is an organ (biology), 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 Visual perception, vision. I ...
and medulla spinalis are developed. Because it originates from the primitive node and is ultimately positioned with the mesodermal space, it is considered to be derived from mesoderm. A postembryonic vestige of the notochord is found in the nucleus pulposus of the intervertebral discs. Isolated notochordal remnants may escape their lineage-specific destination in the nucleus pulposus and instead attach to the outer surfaces of the
vertebral bodies The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic i ...
, from which notochordal cells largely regress.


In amphibians and fish

During development of amphibians and fish, the notochord induces development of the hypochord through secretion of vascular endothelial growth factor. The hypochord is a transient structure ventral to the notochord, and is primarily responsible for correct development of the dorsal aorta. ''Notochord flexion'', when the notochord bends to form a part of the developing caudal fin, is a hallmark of an early growth stage of some fish.


In humans

By the age of 4, all notochord residue is replaced by a population of
chondrocyte Chondrocytes (, from Greek χόνδρος, ''chondros'' = cartilage + κύτος, ''kytos'' = cell) are the only cells found in healthy cartilage. They produce and maintain the cartilaginous matrix, which consists mainly of collagen and prote ...
-like cells of unclear origin. Persistence of notochordal cells within the vertebra may cause a pathologic condition: persistent notochordal canal. If the notochord and the nasopharynx do not separate properly during embryonic development, a depression (Tornwaldt bursa) or
Tornwaldt cyst A Tornwaldt cyst also spelt as Thornwaldt or Thornwald cyst is a benign cyst located in the upper posterior nasopharynx. It can be seen on computed tomography (CT) or magnetic resonance imaging Magnetic resonance imaging (MRI) is a medical ...
may form. The cells are the likely precursors to a rare cancer called chordoma.


Neurology

Research into the notochord has played a key role in understanding the development of 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 p ...
. By transplanting and expressing a second notochord near the dorsal neural tube, 180 degrees opposite of the normal notochord location, one can induce the formation of
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 indirect ...
s in the dorsal tube. Motor neuron formation generally occurs in the ventral neural tube, while the dorsal tube generally forms sensory
cells Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery w ...
. The notochord secretes a protein called sonic hedgehog (SHH), a key
morphogen A morphogen is a substance whose non-uniform distribution governs the pattern of tissue development in the process of morphogenesis or pattern formation, one of the core processes of developmental biology, establishing positions of the vario ...
regulating organogenesis and having a critical role in signaling the development of motor neurons. The secretion of SHH by the notochord establishes the ventral pole of the dorsal-ventral axis in the developing embryo.


Evolution in chordates

The notochord is the defining feature ( synapomorphy) of
chordates 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 five ...
, and was present throughout life in many of the earliest chordates. Although the stomochord of hemichordates was once thought to be homologous, it is now viewed as a convergence. Pikaia appears to have a proto-notochord, and notochords are present in several basal chordates such as Haikouella,
Haikouichthys ''Haikouichthys'' is an extinct genus of craniate (animals with notochords and distinct heads) that lived 518 million years ago, during the Cambrian explosion of multicellular life. ''Haikouichthys'' had a defined skull and other characteristi ...
, and
Myllokunmingia ''Myllokunmingia'' is a genus of basal chordate from the Lower Cambrian Maotianshan shales of China 518 to 490 mya and is thought to be a vertebrate, although this is not conclusively proven. The species M. fengjiaoa is 28 mm long and 6&nbs ...
, all from the
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ...
. The Ordovician oceans included many diverse species of Agnatha and early
Gnathostomata Gnathostomata (; from Greek: (') "jaw" + (') "mouth") are the jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all living vertebrates, including humans. In addition to opposing jaws, livi ...
which possessed notochords, either with attached bony elements or without, most notably the
conodont Conodonts ( Greek ''kōnos'', " cone", + ''odont'', "tooth") are an extinct group of agnathan (jawless) vertebrates resembling eels, classified in the class Conodonta. For many years, they were known only from their tooth-like oral elements, whi ...
s, placoderms, and
ostracoderm Ostracoderms () are the armored jawless fish of the Paleozoic Era. The term does not often appear in classifications today because it is paraphyletic (excluding jawed fishes) (may also be polyphyletic if anaspids are closer to cyclostomes) and ...
s. Even after the evolution of the vertebral column in chondrichthyes and
osteichthyes Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartil ...
, these taxa remained common and are well represented in the fossils record. Several species (see list below) have reverted to the primitive state, retaining the notochord into adulthood, though the reasons for this are not well understood. Scenarios for the evolutionary origin of the notochord were comprehensively reviewed by Annona, Holland, and D'Aniello (2015). They point out that, although many of these ideas have not been well supported by advances in molecular phylogenetics and developmental genetics, two of them have actually been revived under the stimulus of modern molecular approaches (the first proposes that the notochord evolved ''de novo'' in chordates, and the second derives it from a homologous structure, the axochord, that was present in annelid-like ancestors of the chordates). Deciding between these two scenarios (or possibly another yet to be proposed) should be facilitated by much more thorough studies of gene regulatory networks in a wide spectrum of animals.


Post-embryonic retention

In most vertebrates, the notochord develops into secondary structures. In other
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 ...
s, the notochord is retained as an essential anatomical structure. The evolution of the notochord within the phylum Chordata is considered in detail by Holland and Somorjai (2020). The following organisms retain a post-embryonic notochord: *''
Acipenseriformes Acipenseriformes is an order of basal ray-finned fishes that includes living and fossil sturgeons and paddlefishes (Acipenseroidei), as well as the extinct families Chondrosteidae and Peipiaosteidae. They are the second earliest diver ...
'' (
paddlefish Paddlefish (family Polyodontidae) are a family of ray-finned fish belonging to order Acipenseriformes, and one of two living groups of the order alongside sturgeons (Acipenseridae). They are distinguished from other fish by their titular elong ...
and
sturgeon Sturgeon is the common name for the 27 species of fish belonging to the family Acipenseridae. The earliest sturgeon fossils date to the Late Cretaceous, and are descended from other, earlier acipenseriform fish, which date back to the Early ...
) *Lancelet ( Amphioxus) *
Tunicate A tunicate is a marine invertebrate animal, a member of the subphylum Tunicata (). It is part of the Chordata, a phylum which includes all animals with dorsal nerve cords and notochords (including vertebrates). The subphylum was at one t ...
(larval stage only) * Hagfish * Lamprey * Coelacanth * African lungfish *
Tadpole A tadpole is the larval stage in the biological life cycle of an amphibian. Most tadpoles are fully aquatic, though some species of amphibians have tadpoles that are terrestrial. Tadpoles have some fish-like features that may not be found ...
s *
Ostracoderms Ostracoderms () are the armored jawless fish of the Paleozoic Era. The term does not often appear in classifications today because it is paraphyletic (excluding jawed fishes) (may also be polyphyletic if anaspids are closer to cyclostomes) and t ...
(extinct)


Within Amphioxus

The notochord of the lancelet protrudes beyond the anterior end of the neural tube. This projection serves a second purpose in allowing the animal to burrow within the sediment of shallow waters. There, amphioxus is a filter feeder and spends most of its life partially submerged within the sediment.


Additional images

File:Gray14.png, Surface view of embryo of Concolor gibbon ('' Hylobates concolor''). File:Gray29.png, Diagram of a transverse section, showing the mode of formation of the amnion in the chick. File:Gray898.png, Section through the head of a human embryo, about twelve days old, in the region of the hind-brain. File:Gray1111.png, Transverse section of human embryo eight and a half to nine weeks old.


References

{{Authority control Embryology of nervous system *