Mesoderm
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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 Embryology, 11th edition. 2010. The mesoderm forms mesenchyme, mesothelium, non-epithelial blood cells and
coelomocyte A coelomocyte () is a phagocytic leukocyte that appears in the bodies of animals that have a coelom. In most, it attacks and digests invading organisms such as bacteria and viruses through encapsulation and phagocytosis, though in some animals (e.g. ...
s. Mesothelium lines coeloms. Mesoderm forms the muscles in a process known as
myogenesis Myogenesis is the formation of skeletal muscular tissue, particularly during embryonic development. Muscle fibers generally form through the fusion of precursor myoblasts into multinucleated fibers called ''myotubes''. In the early development o ...
, septa (cross-wise partitions) and
mesenteries In zoology, a mesentery is a membrane inside the body cavity of an animal. The term identifies different structures in different phyla: in vertebrates it is a double fold of the peritoneum enclosing the intestines; in other organisms it forms com ...
(length-wise partitions); and forms part of the
gonad A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces sper ...
s (the rest being the
gamete A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce t ...
s). Myogenesis is specifically a function of mesenchyme. The mesoderm differentiates from the rest of the embryo through
intercellular signaling In biology, cell signaling (cell signalling in British English) or cell communication is the ability of a Cell (biology), cell to receive, process, and transmit signals with its environment and with itself. Cell signaling is a fundamental property ...
, after which the mesoderm is polarized by an organizing center. The position of the organizing center is in turn determined by the regions in which
beta-catenin Catenin beta-1, also known as beta-catenin (β-catenin), is a protein that in humans is encoded by the ''CTNNB1'' gene. Beta-catenin is a dual function protein, involved in regulation and coordination of cell–cell adhesion and gene transcrip ...
is protected from degradation by GSK-3. Beta-catenin acts as a co-factor that alters the activity of the transcription factor tcf-3 from repressing to activating, which initiates the synthesis of gene products critical for mesoderm differentiation and gastrulation. Furthermore, mesoderm has the capability to induce the growth of other structures, such as the
neural plate The neural plate is a key developmental structure that serves as the basis for the nervous system. Cranial to the primitive node of the embryonic primitive streak, ectodermal tissue thickens and flattens to become the neural plate. The region ant ...
, the precursor to the nervous system.


Definition

The mesoderm is one of the three germinal layers that appears in the third week of embryonic development. It is formed through a process called gastrulation. There are four important components, which are the axial mesoderm, the
paraxial mesoderm Paraxial mesoderm, also known as presomitic or somitic mesoderm is the area of mesoderm in the neurulating embryo that flanks and forms simultaneously with the neural tube. The cells of this region give rise to somites, blocks of tissue running ...
, the
intermediate mesoderm Intermediate mesoderm or intermediate mesenchyme is a narrow section of the mesoderm (one of the three primary germ layers) located between the paraxial mesoderm and the lateral plate of the developing embryo. The intermediate mesoderm develop ...
, and the
lateral plate mesoderm The lateral plate mesoderm is the mesoderm that is found at the periphery of the embryo. It is to the side of the paraxial mesoderm, and further to the axial mesoderm. The lateral plate mesoderm is separated from the paraxial mesoderm by a narrow ...
. The axial mesoderm gives rise to the notochord. The paraxial mesoderm forms the
somitomere In the developing vertebrate embryo, the somitomeres (or somatomeres) are collections of cell (biology), cells that are derived from the loose masses of paraxial mesoderm that are found alongside the developing neural tube. In human embryogenesis t ...
s, which give rise to mesenchyme of the head, and organize into
somite The somites (outdated term: primitive segments) are a set of bilaterally paired blocks of paraxial mesoderm that form in the embryonic stage of somitogenesis, along the head-to-tail axis in segmented animals. In vertebrates, somites subdivide ...
s in occipital and caudal segments, and give rise to
sclerotome The somites (outdated term: primitive segments) are a set of bilaterally paired blocks of paraxial mesoderm that form in the embryonic stage of somitogenesis, along the head-to-tail axis in segmented animals. In vertebrates, somites subdivide ...
s (cartilage and bone), and
dermatome Dermatome may refer to: * Dermatome (anatomy), an area of skin that is supplied by a single pair of dorsal roots * Dermatome (embryology), the portion of the embryonic paraxial mesoderm, the somite, which gives rise to dermis * Dermatome (instrument ...
s (subcutaneous tissue of the skin). Signals for somite differentiation are derived from surroundings structures, including the notochord,
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, a ...
, and epidermis. The intermediate mesoderm connects the paraxial mesoderm with the lateral plate. Eventually it differentiates into urogenital structures that consist of the kidneys, gonads, their associated ducts, and the adrenal glands. The lateral plate mesoderm gives rise to the heart, blood vessels, and blood cells of the circulatory system, as well as to the mesodermal components of the limbs. Some of the mesoderm derivatives include the muscle (smooth, cardiac, and skeletal), the muscles of the tongue (occipital somites), the pharyngeal arches muscle (muscles of mastication, muscles of facial expressions), connective tissue, the dermis and subcutaneous layer of the
skin Skin is the layer of usually soft, flexible outer tissue covering the body of a vertebrate animal, with three main functions: protection, regulation, and sensation. Other cuticle, animal coverings, such as the arthropod exoskeleton, have diffe ...
,
bone A bone is a Stiffness, rigid Organ (biology), organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red blood cell, red and white blood cells, store minerals, provid ...
and cartilage, dura mater, the
endothelium The endothelium is a single layer of squamous endothelial cells that line the interior surface of blood vessels and lymphatic vessels. The endothelium forms an interface between circulating blood or lymph in the lumen and the rest of the vesse ...
of blood vessels, red blood cells,
white blood cells White blood cells, also called leukocytes or leucocytes, are the cells of the immune system that are involved in protecting the body against both infectious disease and foreign invaders. All white blood cells are produced and derived from mult ...
, microglia, the dentin of teeth, the kidneys, and the adrenal cortex.


Development

During the third week, a process called gastrulation creates a mesodermal layer between the endoderm and the ectoderm. This process begins with the formation of a primitive streak on the surface of the epiblast. The cells of the layers move between the epiblast and the hypoblast, and begin to spread laterally and cranially. The cells of the epiblast move toward the primitive streak and slip beneath it, in a process called "invagination". Some of the migrating cells displace the hypoblast and create the endoderm, and other cells migrate between the endoderm and the epiblast to create the mesoderm. The remaining cells form the ectoderm. After that, the epiblast and the hypoblast establish contact with the extraembryonic mesoderm until they cover the yolk sac and amnion. They move onto either side of the
prechordal plate In the development Development or developing may refer to: Arts *Development hell, when a project is stuck in development *Filmmaking, development phase, including finance and budgeting *Development (music), the process thematic material is ...
. The prechordal cells migrate to the midline to form the notochordal plate. The chordamesoderm is the central region of trunk mesoderm. This forms the notochord, which induces the formation of the neural tube, and establishes the anterior-posterior body axis. The notochord extends beneath the neural tube from the head to the tail. The mesoderm moves to the midline until it covers the notochord. When the mesoderm cells proliferate, they form the paraxial mesoderm. In each side, the mesoderm remains thin, and is known as the lateral plate. The intermediate mesoderm lies between the paraxial mesoderm and the lateral plate. Between days 13 and 15, the proliferation of extraembryonic mesoderm, primitive streak, and embryonic mesoderm take place. The notochord process occurs between days 15 and 17. Eventually, the development of the notochord canal and the axial canal takes place between days 17 and 19, when the first three somites are formed.


Paraxial mesoderm

During the third week, the paraxial mesoderm is organized into segments. If they appear in the cephalic region and grow with cephalocaudal direction, they are called somitomeres. If they appear in the cephalic region but establish contact with the neural plate, they are known as neuromeres, which later will form the mesenchyme in the head. The somitomeres organize into somites which grow in pairs. In the fourth week the somites lose their organization and cover the notochord and spinal cord to form the backbone. In the fifth week, there are 4 occipital somites, 8 cervical, 12 thoracic, 5 lumbar, 5 sacral and 8 to 10 coccygeal that will form the axial skeleton. Somitic derivatives are determined by local signaling between adjacent embryonic tissues, in particular the neural tube, notochord, surface ectoderm and the somitic compartments themselves. The correct specification of the deriving tissues, skeletal, cartilage, endothelia and connective tissue is achieved by a sequence of morphogenic changes of the paraxial mesoderm, leading to the three transitory somitic compartments: dermomyotome, myotome and sclerotome. These structures are specified from dorsal to ventral and from medial to lateral. each somite will form its own sclerotome that will differentiate into the tendon cartilage and bone component. Its myotome will form the muscle component and the dermatome that will form the dermis of the back. The myotome and dermatome have a nerve component.


Molecular regulation of somite differentiation

Surrounding structures such as the notochord, neural tube, epidermis and lateral plate mesoderm send signals for somite differentiation Notochord protein accumulates in presomitic mesoderm destined to form the next somite and then decreases as that somite is established. The notochord and the neural tube activate the protein SHH which helps the somite to form its sclerotome. The cells of the sclerotome express the protein PAX1 that induces the cartilage and bone formation. The neural tube activates the protein WNT1 that expresses PAX 2 so the somite creates the myotome and dermatome. Finally, the neural tube also secretes neurotrophin 3 (NT-3), so that the somite creates the dermis. Boundaries for each somite are regulated by
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 ...
(RA) and a combination of FGF8 and WNT3a. So
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 ...
is an endogenous signal that maintains the bilateral synchrony of mesoderm segmentation and controls bilateral symmetry in vertebrates. The bilaterally symmetric body plan of vertebrate embryos is obvious in somites and their derivates such as the vertebral column. Therefore, asymmetric somite formation correlates with a left-right desynchronization of the segmentation oscillations. Many studies with ''Xenopus'' and zebrafish have analyzed the factors of this development and how they interact in signaling and transcription. However, there are still some doubts in how the prospective mesodermal cells integrate the various signals they receive and how they regulate their morphogenic behaviours and cell-fate decisions. Human embryonic stem cells for example have the potential to produce all of the cells in the body and they are able to self-renew indefinitely so they can be used for a large-scale production of therapeutic cell lines. They are also able to remodel and contract collagen and were induced to express muscle actin. This shows that these cells are multipotent cells.


Intermediate mesoderm

The intermediate mesoderm connects the paraxial mesoderm with the lateral plate mesoderm, and differentiates into urogenital structures. In upper thoracic and cervical regions, this forms the nephrotomes. In caudal regions, it forms the nephrogenic cord. It also helps to develop the excretory units of the urinary system and the gonads.


Lateral plate mesoderm

The
lateral plate mesoderm The lateral plate mesoderm is the mesoderm that is found at the periphery of the embryo. It is to the side of the paraxial mesoderm, and further to the axial mesoderm. The lateral plate mesoderm is separated from the paraxial mesoderm by a narrow ...
splits into the parietal (somatic) and visceral (splanchnic) layers. The formation of these layers starts with the appearance of intercellular cavities. The somatic layer depends upon a continuous layer with mesoderm that covers the amnion. The splanchnic layer depends upon a continuous layer that covers the yolk sac. The two layers cover the intraembryonic cavity. The parietal layer, together with overlying ectoderm, forms the lateral body wall folds. The visceral layer forms the walls of the gut tube. Mesoderm cells of the parietal layer form the mesothelial membranes or serous membranes, which line the peritoneal, pleural, and pericardial cavities.


See also

*
Histogenesis Histogenesis is the formation of different tissues from undifferentiated cells. These cells are constituents of three primary germ layers, the endoderm, mesoderm, and ectoderm. The science of the microscopic structures of the tissues formed wi ...
*
Organogenesis Organogenesis is the phase of embryonic development that starts at the end of gastrulation and continues until birth. During organogenesis, the three germ layers formed from gastrulation (the ectoderm, endoderm, and mesoderm) form the internal org ...
*
Triploblastic Triploblasty is a condition of the gastrula in which there are three primary germ layers: the ectoderm, mesoderm, and endoderm. Germ cells are set aside in the embryo at the blastula stage, which are incorporated into the gonads during organo ...
* List of human cell types derived from the germ layers


References


Further reading

* * * * * *


External links

* * {{Embryology Germ layers Developmental biology Embryology Gastrulation