Notochord

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In anatomy, the notochord is a flexible rod

made out of a material similar to cartilage.
If a species has a notochord at any stage
of its life cycle, it is, by definition, a
chordate. In vertebrates the notochord
becomes part of the vertebral column. The
notochord lies along the anteroposterior
("head to tail") axis, is usually closer to the
ventral than the dorsal surface of the
animal, and is composed of cells derived
from the mesoderm. The notochord has
been observed to have many functions
including developmental functions. The
most commonly cited functions are as a
site of muscle attachment, vertebral
precursor, and as a midline tissue that
provides signals to the surrounding tissue
during development.[1]
 Notochord

Transverse section of a chick embryo of forty-five

hours’ incubation.

Details

Precursor axial mesoderm

Gives rise to nucleus pulposus

Identifiers

Latin notochorda

MeSH D009672

TE E5.0.1.1.0.0.8

Anatomical terminology
Notochords are thought to be
advantageous (both in an evolutionary and
developmental context) because they
provide(d) rigid structure for muscle
attachment, but were still flexible. In some
chordates, it persists throughout life as the
main structural support of the body, while
in most vertebrates it becomes the
nucleus pulposus of the intervertebral
disc.[1] The notochord plays a key role in
signaling and coordinating development.
Embryos of vertebrates still form transient
notochord structures today during the
gastrulation phase of development. The
notochord is found ventral to the neural
tube.
Development
Notogenesis is the development of the
notochord by the epiblasts that make up
the floor of the amnion cavity.[2] The
progenitor notochord is derived from cells
migrating from the primitive node and
pit.[3] The notochord forms during
gastrulation 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 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, where it ends in a
hook-like extremity in the region of the
future dorsum sellae of the sphenoid
bone. Initially it exists between the neural
tube and the endoderm of the yolk-sac, but
soon becomes separated from them by
the mesoderm, which grows medially and
surrounds it. From the mesoderm
surrounding the neural tube and
notochord, the skull, vertebral column, and
the membranes of the brain and medulla
spinalis are developed.

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, from
which notochordal cells largely regress.[4]
In humans, by the age of 4, all notochord
residue is replaced by a population of
chondrocyte-like cells of unclear origin.[5]
Persistence of notochordal cells within the
vertebra may cause a pathologic
condition: persistent notochordal canal.[6]
They are also found to persist in the
nasopharyngeal space and, in such an
unusual instance, may give rise to a
Tornwaldt cyst.

Neurology
Research into the notochord has played a
key role in understanding the development
of the central nervous system. 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 neurons in the dorsal tube. Motor
neuron formation generally occurs in the
ventral neural tube, while the dorsal tube
generally forms sensory cells.

The notochord secretes a protein called

sonic hedgehog (SHH), a key morphogen
regulating organogenesis and having a
critical role in signaling the development
of motor neurons.[7] The secretion of SHH
by the notochord establishes the ventral
pole of the dorsal-ventral axis in the
developing embryo.

Evolution

A dissected spotted African lungfish showing the

notochord

The notochord is the defining feature of

chordates, 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.[8] Pikaia appears to have a
proto-notochord, and notochords are
present in several basal chordates such as
Haikouella, Haikouichthys, and
Myllokunmingia, all from the Cambrian.
The Ordovician oceans included many
diverse species of Agnatha and early
Gnathostomata which possessed
notochords, either with attached bony
elements or without, most notably the
conodonts,[9] placoderms[10] and
ostracoderms. Even after the evolution of
the vertebral column in chondrichthyes
and osteichthyes, 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 have been
comprehensively reviewed (Annona, G.,
Holland, N. D., and D'Aniello, S. 2015.
Evolution of the notochord. EvoDevo 6:
article 30). 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.

Structure
The notochord is a long rod like structure
that develops between dorsal nervous
system and gut. The notochord is
composed primarily of a core of
glycoproteins, encased in a sheath of
collagen fibers wound into two opposing
helices.The angle between these fibers
determines whether increased pressure in
the core will result in shortening and
thickening versus lengthening and
thinning.[11]

Organisms which retain a

post-embryonic notochord
Acipenseriformes (paddlefish and
sturgeon)[12]
Amphioxus
Tunicate larvae
 Hagfish
Lamprey
Coelacanth
African lungfish
Tadpoles
Ostracoderms (extinct)

Additional images
Surface view of embryo of Concolor
gibbon (Hylobates concolor).

Diagram of a transverse section, showing

the mode of formation of the amnion in
the chick.
Section through the head of a human
embryo, about twelve days old, in the
region of the hind-brain.

Transverse section of human embryo eight

and a half to nine weeks old.
References
1. Stemple, Derek L. (2005-06-01).
"Structure and function of the notochord: an
essential organ for chordate" .
Development. 132 (11): 2503–2512.
doi:10.1242/dev.01812 . ISSN 0950-1991 .
PMID 15890825 .
2. The trilaminar germ disk (3rd week)
3. Hood, Rousseaux, Blakley, Ronald D.,
Colin G., Patricia M. (29 May 2007).
"Embryo and Fetus" . Handbook of
Toxicologic Pathology (Second Edition).
Academic Press, Published by Elsevier Inc.
2: Pages 895–936. doi:10.1016/b978-0-12-
330215-1.50047-8 .
4. Choi, K.; Cohn, Martin J.; Harfe, Brian D.
(2009). "Identification of Nucleus Pulposus
Precursor Cells and Notochordal Remnants
in the Mouse: Implications for Disk
Degeneration and Chordoma Formation" .
Developmental Dynamics. 237 (12): 3953–
3958. doi:10.1002/dvdy.21805 .
PMC 2646501 . PMID 19035356 .
5. Urban, J. P. G. (2000). "The Nucleus of
the Intervertebral Disc from Development to
Degeneration". Integrative and Comparative
Biology. 40: 53. doi:10.1093/icb/40.1.53 .
6. Christopherson, Lr; Rabin, Bm; Hallam,
Dk; Russell, Ej (1 January 1999).
"Persistence of the notochordal canal: MR
and plain film appearance" (Free full text).
AJNR. American journal of neuroradiology.
20 (1): 33–6. ISSN 0195-6108 .
PMID 9974055 .
7. Echelard, Y; Epstein, Dj; St-Jacques, B;
Shen, L; Mohler, J; Mcmahon, Ja; Mcmahon,
Ap (December 1993). "Sonic hedgehog, a
member of a family of putative signaling
molecules, is implicated in the regulation of
CNS polarity". Cell. 75 (7): 1417–30.
doi:10.1016/0092-8674(93)90627-3 .
PMID 7916661 .
8. Kardong, Kenneth V. (1995). Vertebrates:
comparative anatomy, function, evolution.
McGraw-Hill. pp. 55, 57. ISBN 0-697-21991-
7.
9. "Archived copy" . Archived from the
original on 2006-03-13. Retrieved
2007-09-05.
10. "Archived copy" . Archived from the
original on 2010-12-20. Retrieved
2009-11-21.
11. M. A. R. Koehl. "Mechanical Design of
Fiber-Wound Hydraulic Skeletons: The
Stiffening and Straightening of Embryonic
Notochords" .
12. Joseph J. Luczkovich; Philip J. Motta;
Stephen F. Norton; Karel F. Liem (17 April
2013). Ecomorphology of fishes . Springer
Science & Business Media. p. 201.
ISBN 978-94-017-1356-6.

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