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3/28/2019 Mesenchyme - Wikipedia

Mesenchyme
Mesenchy m e, in v ertebrate embry ology , is a ty pe of connectiv e tissue
Mesenchyme
found mostly during the dev elopment of the embry o. [1 ] It is composed
mainly of ground substance with few cells or fibers. It can also refer to a
group of mucoproteins found in certain ty pes of cy sts (etc.), resembling
mucus. It is most easily found as a component of Wharton's jelly .

The v itreous of the ey eball is a similar tissue. [2 ]

In inv ertebrate zoology , the term refers to free cells loosely arranged in a
matrix. [3 ]

Contents
Terminology
In vertebrate embryology Transmission electron micrograph
Structure of mesenchyme displaying the
Development ultrastructure of a typical cell and
Epithelial to mesenchymal transition
matrix.
Implantation
Primary mesenchyme
Neural mesenchyme

In invertebrate zoology
References
External links

Mesenchyme (pointer) stained with


Terminology H&E
"Mesenchy me" is a term introduced by Oscar Hertwig in 1881. [3 ] Details
Carnegie 6b (http://genex.hgu.mr
In order to differentiate the use of the word mesenchy me in inv ertebrate
stage c.ac.uk/Databases/Hum
zoology (an ecto- or entomesodermal middle lay er of some inv ertebrates)
anAnatomy/new/carneg
and the use in v ertebrate embry ology (that is, undifferentiated tissue found
ie6b.shtml)
in embry onic true mesoderm - entomesoderm - from which are deriv ed all
connectiv e tissues, blood v essels, blood cells, the ly mphatic sy stem, and Precursor Primarily mesoderm
the heart), some authors prefer to use the term mesoglea (in wider sense) in Identifiers
lieu of mesenchy me when referring to the middle lay ers of sponges and
TE E5.16.4.0.3.0.18 (http
diploblasts, reserv ing the term mesenchy me for the embry ological sense.
s://www.unifr.ch/ifaa/Pu
Howev er, Brusca & Brusca discourage this usage, using mesoglea in its
blic/EntryPage/ViewTE/
strict sense (noncellular mesenchy me), and preferring to maintain both the
TEe0516.html)
embry ological and zoological senses for the term mesenchy me. [4 ]
Anatomical terminology

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Finally , some similar terms used in botany generally are differentiated by the suffix "a": mesenchy ma (a tissue
between xy lem and phloem in roots), collenchy ma (primordial leaf tissues) and parenchy ma (supportiv e tissues). [4 ]

In vertebrate embryology

Structure
Mesenchy me is characterized morphologically by a prominent ground substance matrix containing a loose
aggregate of reticular fibrils and unspecialized cells. [5 ] Mesenchy mal cells can migrate easily , in contrast to
epithelial cells, which lack mobility and are organized into closely adherent sheets, and are polarized in an apical-
basal orientation.

Development
The mesenchy me originates from the mesoderm. [6 ] From the mesoderm, the mesenchy me appears as an
embry ologically primitiv e "soup". This "soup" exists as a combination of the mesenchy mal cells plus serous fluid plus
the many different tissue proteins. Serous fluid is ty pically stocked with the many serous elements, such as sodium
and chloride. The mesenchy me dev elops into the tissues of the ly mphatic and circulatory sy stems, as well as the
musculoskeletal sy stem. This latter sy stem is characterized as connectiv e tissues throughout the body , such as bone,
muscle and cartilage. A malignant cancer of mesenchy mal cells is a ty pe of sarcoma. [7 ][8 ]

Epithelial to mesenchymal transition


The first emergence of mesenchy me occurs during gastrulation from the epithelial–mesenchy mal transition (EMT)
process. This transition occurs through the loss of epithelial cadherin, tight junctions, and adherens junctions on the
cell membranes of epithelial cells. [9 ] The surface molecules undergo endocy tosis and the microtubule cy toskeleton
loses shape, enabling mesenchy me to migrate along the extracellular matrix (ECM). Epithelial–mesenchy mal
transition occurs in embry onic cells that require migration through or ov er tissue, and can be followed with a
mesenchy mal–epithelial transition to produce secondary epithelial tissues. Embry ological mesenchy mal cells
expresses fibroblast-specific protein (Fsp1), which is indicativ e of their shared properties with the migratory adult
fibroblasts, and c-Fos, an oncogene associated with the down-regulation of epithelial cadherin. [1 0 ][1 1 ] Both
formation of the primitiv e streak and mesenchy mal tissue is dependent on the Wnt/β-catenin pathway . [1 2 ] Specific
markers of mesenchy mal tissue include the additional expression of ECM factors such as fibronectin and
v itronectin. [1 3 ]

Implantation
The first cells of the embry o to undergo EMT and form mesenchy me are the extra-embry onic cells of the
trophectoderm. These migrate from the body of the blastocy st into the endometrial lay er of the uterus in order to
contribute to the formation of the anchored placenta. [1 4 ]

Primary mesenchyme
Primary mesenchy me is the first embry onic mesenchy mal tissue to emerge, and it is produced from EMT in epiblast
cells. In the epiblast, it is induced by the primitiv e streak through Wnt signaling, and produces endoderm and
mesoderm from a transitory tissue called mesendoderm during the process of gastrulation. [1 5 ]

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The formation of primary mesenchy me depends on the expression of WNT3. Other deficiencies in the Wnt signaling
pathway , such as in Nodal (a TGF-beta protein), will lead to defectiv e mesoderm formation. [9 ]

The tissue lay ers formed from the primitiv e streak inv aginate together into the embry o and the induced
mesenchy mal stem cells will ingress and form the mesoderm. Mesodermal tissue will continue to differentiate
and/or migrate throughout the embry o to ultimately form most connectiv e tissue lay ers of the body . [1 6 ]

Neural mesenchyme
Embry ological mesenchy me is particularly transitory and soon differentiates after migration. Neural mesenchy me
forms soon after primary mesenchy me formation. [1 7 ]

The interaction with ectoderm and somite-forming morphogenic factors cause some primary mesenchy me to form
neural mesenchy me, or paraxial mesoderm, and contribute to somite formation. Neural mesenchy me soon
undergoes a mesenchy mal–epithelial transition under the influence of WNT6 produced by ectoderm to form
somites. [1 8 ] These structures will undergo a secondary EMT as the somite tissue migrates later in dev elopment to
form structural connectiv e tissue such as cartilage and skeletal muscle. [1 9 ]

Neural crest cells (NCCs) form from neuroectoderm, instead of the primary mesenchy me, from morphogenic signals
of the neural crest. The EMT occurs as a result of Wnt signaling, the influence of Sox genes and the loss of E-cadherin
from the cell surface. NCCs additionally require the repression of N-cadherin, and neural cell adhesion molecule.
NCCs ingress into the embry o from the epithelial neuroectodermal lay er and migrate throughout the body in order
form multiple peripheral nerv ous sy stem (PNS) cells and melanocy tes. Migration of NCCs is primarily induced by
BMP signaling and its inhibitor, Noggin. [2 0 ][2 1 ]

In invertebrate zoology
In some inv ertebrates, e.g., Porifera, Cnidaria, Ctenophora and some triploblasts (the acoelomates), the term
"mesenchy me" refers to a more-or-less solid but looselly organized tissue consisting of a gel matrix (the mesoglea)
with v arious cellular and fibrous inclusions, located between epidermis and gastrodermis. In some cases, the
mesoglea is noncellular. [2 2 ]

In sponges, the mesenchyme is called mesohyl.[4]


In diploblasts (Cnidaria and Ctenophora), the mesenchyme is fully ectodermally derived. This kind of mesenchyme is
called ectomesodermal, and is not considered true mesoderm.
In triploblastic acoelomates (e.g., flatworms), the term parenchyme is sometimes used for the middle (mesenchymal)
layer, in which the dense layer includes tissues derived from both ecto- and entomesoderm (true mesoderm, derived
from entoderm).
When cellular material is sparse or densely packed, the mesenchy me may be sometimes called collenchy me (e.g.,
cnidarians) or parenchy me (e.g., flatworms), respectiv ely . [4 ] When no cellular material is present (e.g., in
Hy drozoa), the lay er is properly called mesoglea. [4 ]

In some colonial cnidarians, the mesenchy me is perforated by gastrov ascular channels continuous among colony
members. This entire matrix of common basal material is called coenenchy me. [4 ]

References
1. "Blue Histology - Connective Tissues" (http://www.lab.anhb.uwa.edu.au/mb140/CorePages/Connective/Connect.htm#me
senchymal). Retrieved 2008-12-05.

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2. Singh, Inderbir (April 15, 2007). Textbook of Human Histology With Colour Atlas. Jaypee Brothers Medical Publishers..
ISBN 978-81-7179-967-1.
3. Collins, P.; Billett, F. S. (1995). "The terminology of early development: History, concepts, and current usage". Clinical
Anatomy. 8 (6): 418–425. doi:10.1002/ca.980080610 (https://doi.org/10.1002%2Fca.980080610). PMID 8713164 (https://
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6. Kierszenbaum, Abraham L.; Tres, Laura (2015). Histology and Cell Biology: An Introduction to Pathology E-Book (http
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Williams & Wilkins. p. 83. ISBN 978-0-7817-8577-8.
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gov/pmc/articles/PMC2689101). Journal of Clinical Investigation. 119 (6): 1420–8. doi:10.1172/JCI39104 (https://doi.org/
10.1172%2FJCI39104). PMC 2689101 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2689101). PMID 19487818 (http
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estrogen receptor activation involves nuclear translocation of beta-catenin and upregulation of beta-catenin/lymphoid
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of Cell Biology. 148 (1): 173–88. doi:10.1083/jcb.148.1.173 (https://doi.org/10.1083%2Fjcb.148.1.173). PMC 3207144 (ht
tps://www.ncbi.nlm.nih.gov/pmc/articles/PMC3207144). PMID 10629227 (https://www.ncbi.nlm.nih.gov/pubmed/1062922
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formation in the mouse embryo". Developmental Dynamics. 231 (2): 416–24. doi:10.1002/dvdy.20135 (https://doi.org/10.
1002%2Fdvdy.20135). PMID 15366019 (https://www.ncbi.nlm.nih.gov/pubmed/15366019).
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14. Yamakoshi, S; Bai, R; Chaen, T; Ideta, A; Aoyagi, Y; Sakurai, T; Konno, T; Imakawa, K (2012). "Expression of
mesenchymal-related genes by the bovine trophectoderm following conceptus attachment to the endometrial
epithelium". Reproduction (Cambridge, England). 143 (3): 377–87. doi:10.1530/REP-11-0364 (https://doi.org/10.1530%2
FREP-11-0364). PMID 22157247 (https://www.ncbi.nlm.nih.gov/pubmed/22157247).
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rg/10.1007%2Fbf00318331). PMID 3518538 (https://www.ncbi.nlm.nih.gov/pubmed/3518538).
16. Hay, E. D. (2005). "The mesenchymal cell, its role in the embryo, and the remarkable signaling mechanisms that create
it". Developmental Dynamics. 233 (3): 706–20. doi:10.1002/dvdy.20345 (https://doi.org/10.1002%2Fdvdy.20345).
PMID 15937929 (https://www.ncbi.nlm.nih.gov/pubmed/15937929).
17. Mareschi, K; Novara, M; Rustichelli, D; Ferrero, I; Guido, D; Carbone, E; Medico, E; Madon, E; Vercelli, A; Fagioli, F
(2006). "Neural differentiation of human mesenchymal stem cells: Evidence for expression of neural markers and eag K+
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6%2Fj.exphem.2006.06.020). PMID 17046576 (https://www.ncbi.nlm.nih.gov/pubmed/17046576).
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process of the segmental plate mesoderm leading to somite formation". Developmental Biology. 271 (1): 198–209.
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m.nih.gov/pubmed/15196961).

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19. Stockdale, F. E.; Nikovits Jr, W; Christ, B (2000). "Molecular and cellular biology of avian somite development".
Developmental Dynamics. 219 (3): 304–21. doi:10.1002/1097-0177(2000)9999:9999<::AID-DVDY1057>3.0.CO;2-5 (http
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External links
Histology at ucsd.edu (http://meded.ucsd.edu/hist-img-bank/chapter_1/Slide_7_umbilical/index.htm)

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