A germ layer is a primary layer of cells that forms during

embryonic development 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 ...

. The three germ layers in

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, ...

s are particularly pronounced; however, all

eumetazoa Eumetazoa (), also known as diploblasts, Epitheliozoa, or Histozoa, are a proposed basal animal clade as a sister group of the Porifera (sponges). The basal eumetazoan clades are the Ctenophora and the ParaHoxozoa. Placozoa is now also seen as a ...

ns (

animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...

s that are sister taxa to the

sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through t ...

s) produce two or three primary germ layers. Some animals, like

cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that th ...

ns, produce two germ layers (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 t ...

and

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 ...

) making them

diploblastic Diploblasty is a condition of the blastula in which there are two primary germ layers: the ectoderm and endoderm. Diploblastic organisms are organisms which develop from such a blastula, and include cnidaria and ctenophora, formerly grouped togeth ...

. Other animals such as bilaterians produce a third layer (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 E ...

) between these two layers, making them

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 organogenes ...

. Germ layers eventually give rise to all of an animal’s tissues and

organ Organ may refer to: Biology * Organ (biology), a part of an organism Musical instruments * Organ (music), a family of keyboard musical instruments characterized by sustained tone ** Electronic organ, an electronic keyboard instrument ** Hammond ...

s through the process of organogenesis.

History

Caspar Friedrich Wolff Caspar Friedrich Wolff (18 January 1733 – 22 February 1794) was a German physiologist and one of the founders of embryology. Life Wolff was born in Berlin, Brandenburg. In 1759 he graduated as an M.D. from the University of Halle with his disse ...

observed organization of the early embryo in leaf-like layers. In 1817, Heinz Christian Pander discovered three primordial germ layers while studying chick embryos. Between 1850 and 1855, Robert Remak had further refined the germ cell layer (''Keimblatt'') concept, stating that the external, internal and middle layers form respectively the epidermis, the gut, and the intervening musculature and vasculature. The term "

" was introduced into English by

Huxley Huxley may refer to: People * Huxley (surname) * The British Huxley family * Thomas Henry Huxley (1825–1895), British biologist known as "Darwin's Bulldog" * Aldous Huxley (1894–1963), British writer, author of ''Brave New World'', grandson ...

in 1871, and "

" and "

" by Lankester in 1873.

Evolution

Among

sponges Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through ...

show the least amount of compartmentalization, having a single germ layer. Although they have differentiated cells (e.g. collar cells), they lack true tissue coordination.

Diploblastic Diploblasty is a condition of the blastula in which there are two primary germ layers: the ectoderm and endoderm. Diploblastic organisms are organisms which develop from such a blastula, and include cnidaria and ctenophora, formerly grouped togeth ...

animals,

Cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that th ...

and Ctenophora, show an increase in compartmentalization, having two germ layers, the

and

. Diploblastic animals are organized into recognisable tissues. All

bilaterian The Bilateria or bilaterians are animals with bilateral symmetry as an embryo, i.e. having a left and a right side that are mirror images of each other. This also means they have a head and a tail (anterior-posterior axis) as well as a belly an ...

animals (from flatworms to humans) are

, possessing a

in addition to the germ layers found in Diploblasts. Triploblastic animals develop recognizable organs.

Development

Fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Proce ...

leads to the formation of a

zygote A zygote (, ) is a eukaryotic cell formed by a fertilization event between two gametes. The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individual organism. In multicellula ...

. During the next stage, cleavage, mitotic cell divisions transform the zygote into a hollow ball of cells, a

blastula Blastulation is the stage in early animal embryonic development that produces the blastula. In mammalian development the blastula develops into the blastocyst with a differentiated inner cell mass and an outer trophectoderm. The blastula (from ...

. This early embryonic form undergoes

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. Be ...

, forming a gastrula with either two or three layers (the germ layers). In all

s, these progenitor cells differentiate into all adult tissues and organs. In the

human embryo Human embryonic development, or human embryogenesis, is the development and formation of the human embryo. It is characterised by the processes of cell division and cellular differentiation of the embryo that occurs during the early stages of de ...

, after about three days, the zygote forms a solid mass of cells by mitotic division, called a morula. This then changes to a

blastocyst The blastocyst is a structure formed in the early embryonic development of mammals. It possesses an inner cell mass (ICM) also known as the ''embryoblast'' which subsequently forms the embryo, and an outer layer of trophoblast cells called the t ...

, consisting of an outer layer called a

trophoblast The trophoblast (from Greek : to feed; and : germinator) is the outer layer of cells of the blastocyst. Trophoblasts are present four days after fertilization in humans. They provide nutrients to the embryo and develop into a large part of the pl ...

, and an inner cell mass called the embryoblast. Filled with uterine fluid, the blastocyst breaks out of the zona pellucida and undergoes implantation. The inner cell mass initially has two layers: the

hypoblast In amniote embryology, the hypoblast, is one of two distinct layers arising from the inner cell mass in the mammalian blastocyst, or from the blastodisc in reptiles and birds. The hypoblast gives rise to the yolk sac, which in turn gives rise to ...

and

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 t ...

. At the end of the second week, a

primitive streak The primitive streak is a structure that forms in the early embryo in amniotes. In amphibians the equivalent structure is the blastopore. During early embryonic development, the embryonic disc becomes oval shaped, and then pear-shaped with the ...

appears. The epiblast in this region moves towards the primitive streak, dives down into it, and forms a new layer, called the

, pushing the hypoblast out of the way (this goes on to form the

amnion The amnion is a membrane that closely covers the human and various other embryos when first formed. It fills with amniotic fluid, which causes the amnion to expand and become the amniotic sac that provides a protective environment for the develo ...

.) The epiblast keeps moving and forms a second layer, the

. The top layer is now called the

. Gastrulation occurs in reference to the primary

body axis 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 prov ...

. Germ layer formation is linked to the primary body axis as well, however it is less reliant on it than gastrulation is. '' Hydractinia'' shows that germ layer formation that transpires as a mixed delamination. In mice, germ layer differentiation is controlled by two

transcription factor In molecular biology, a transcription factor (TF) (or sequence-specific DNA-binding factor) is a protein that controls the rate of transcription of genetic information from DNA to messenger RNA, by binding to a specific DNA sequence. The fu ...

s: Sox2 and Oct4 proteins. These transcription factors cause the pluripotent mouse embryonic stem cells to select a germ layer fate. Sox2 promotes ectodermal differentiation, while Oct4 promotes mesendodermal differentiation. Each gene inhibits what the other promotes. Amounts of each protein are different throughout the genome, causing the embryonic stem cells to select their fate.

The germ layers

Endoderm

The endoderm is one of the germ layers formed during animal

. Cells migrating inward along the archenteron form the inner layer of the gastrula, which develops into the

. The endoderm consists at first of flattened cells, which subsequently become columnar. It forms the epithelial lining of the whole of the digestive tract except part of the mouth and pharynx and the terminal part of the rectum (which are lined by involutions of the ectoderm). It also forms the lining cells of all the glands which open into the digestive tract, including those of the liver and pancreas; the epithelium of the auditory tube and tympanic cavity; the trachea, bronchi, and alveoli of the lungs; the bladder and part of the urethra; and the follicle lining of the thyroid gland and thymus. The endoderm forms: the

pharynx The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its struc ...

, the

esophagus The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to the ...

, the stomach, the

small intestine The small intestine or small bowel is an organ in the gastrointestinal tract where most of the absorption of nutrients from food takes place. It lies between the stomach and large intestine, and receives bile and pancreatic juice through the p ...

, the colon, the

liver The liver is a major Organ (anatomy), organ only found in vertebrates which performs many essential biological functions such as detoxification of the organism, and the Protein biosynthesis, synthesis of proteins and biochemicals necessary for ...

, the

pancreas The pancreas is an organ of the digestive system and endocrine system of vertebrates. In humans, it is located in the abdomen behind the stomach and functions as a gland. The pancreas is a mixed or heterocrine gland, i.e. it has both an end ...

, the

bladder The urinary bladder, or simply bladder, is a hollow organ in humans and other vertebrates that stores urine from the kidneys before disposal by urination. In humans the bladder is a distensible organ that sits on the pelvic floor. Urine enters ...

, the

epithelial Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellula ...

parts of the

trachea The trachea, also known as the windpipe, is a Cartilage, cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air-breathing animals with lungs. The trachea extends ...

and bronchi, the

lungs The lungs are the primary organs of the respiratory system in humans and most other animals, including some snails and a small number of fish. In mammals and most other vertebrates, two lungs are located near the backbone on either side of th ...

, the

thyroid The thyroid, or thyroid gland, is an endocrine gland in vertebrates. In humans it is in the neck and consists of two connected lobes. The lower two thirds of the lobes are connected by a thin band of tissue called the thyroid isthmus. The thy ...

, and the

parathyroid Parathyroid glands are small endocrine glands in the neck of humans and other tetrapods. Humans usually have four parathyroid glands, located on the back of the thyroid gland in variable locations. The parathyroid gland produces and secretes par ...

Mesoderm

The mesoderm germ layer forms in 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 ...

s of

s. During

, some of the cells migrating inward contribute to the mesoderm, an additional layer between the endoderm and the

. The formation of a mesoderm leads to the development of a

coelom The coelom (or celom) is the main body cavity in most animals and is positioned inside the body to surround and contain the digestive tract and other organs. In some animals, it is lined with mesothelium. In other animals, such as molluscs, it r ...

. Organs formed inside a coelom can freely move, grow, and develop independently of the body wall while fluid cushions protects them from shocks. The mesoderm has several components which develop into tissues:

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 develops i ...

, paraxial mesoderm,

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 ...

, and chorda-mesoderm. The chorda-mesoderm develops into the notochord. The intermediate mesoderm develops into kidneys and gonads. The paraxial mesoderm develops into cartilage, skeletal muscle, and dermis. The lateral plate mesoderm develops into the circulatory system (including the heart and spleen), the wall of the gut, and wall of the human body. Through cell signaling cascades and interactions with the ectodermal and endodermal cells, the mesodermal cells begin the process of differentiation. The mesoderm forms: muscle ( smooth and

striated Striations means a series of ridges, furrows or linear marks, and is used in several ways: * Glacial striation * Striation (fatigue), in material * Striation (geology), a ''striation'' as a result of a geological fault * Striation Valley, in ...

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 ...

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 an ...

connective tissue Connective tissue is one of the four primary types of animal tissue, along with epithelial tissue, muscle tissue, and nervous tissue. It develops from the mesenchyme derived from the mesoderm the middle embryonic germ layer. Connective tiss ...

, adipose tissue,

circulatory system The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, tha ...

lymphatic system The lymphatic system, or lymphoid system, is an organ system in vertebrates that is part of the immune system, and complementary to the circulatory system. It consists of a large network of lymphatic vessels, lymph nodes, lymphatic or lymphoid o ...

dermis The dermis or corium is a layer of skin between the epidermis (with which it makes up the cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from stress and strain. It is divided i ...

, Dentine of teeth,

genitourinary system The genitourinary system, or urogenital system, are the organs of the reproductive system and the urinary system. These are grouped together because of their proximity to each other, their common embryological origin and the use of common pathwa ...

, serous membranes,

spleen The spleen is an organ found in almost all vertebrates. Similar in structure to a large lymph node, it acts primarily as a blood filter. The word spleen comes .

and

notochord In anatomy, the notochord is a flexible rod which is similar in structure to the stiffer cartilage. If a species has a notochord at any stage of its life cycle (along with 4 other features), it is, by definition, a chordate. The notochord consis ...

Ectoderm

The ectoderm generates the outer layer of the embryo, and it forms from the embryo's

. The

develops into the

surface ectoderm The surface ectoderm (or external ectoderm) forms the following structures: *Skin (only epidermis; dermis is derived from mesoderm) (along with glands, hair, and nails) *Epithelium of the mouth and nasal cavity and glands of the mouth and nasal cav ...

, neural crest, and the neural tube. The surface ectoderm develops into:

epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water rele ...

hair Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and f ...

, nails,

lens of the eye The lens, or crystalline lens, is a transparent biconvex structure in the eye that, along with the cornea, helps to refract light to be focused on the retina. By changing shape, it functions to change the focal length of the eye so that it can ...

sebaceous glands A sebaceous gland is a microscopic exocrine gland in the skin that opens into a hair follicle to secrete an oily or waxy matter, called sebum, which lubricates the hair and skin of mammals. In humans, sebaceous glands occur in the greatest number ...

cornea The cornea is the transparent front part of the eye that covers the iris, pupil, and anterior chamber. Along with the anterior chamber and lens, the cornea refracts light, accounting for approximately two-thirds of the eye's total optical power ...

tooth enamel Tooth enamel is one of the four major Tissue (biology), tissues that make up the tooth in humans and many other animals, including some species of fish. It makes up the normally visible part of the tooth, covering the Crown (tooth), crown. The ...

, the epithelium of the

mouth In animal anatomy, the mouth, also known as the oral cavity, or in Latin cavum oris, is the opening through which many animals take in food and issue vocal sounds. It is also the cavity lying at the upper end of the alimentary canal, bounded on ...

and

nose A nose is a protuberance in vertebrates that houses the nostrils, or nares, which receive and expel air for respiration alongside the mouth. Behind the nose are the olfactory mucosa and the sinuses. Behind the nasal cavity, air next passes th ...

. The neural crest of the ectoderm develops into:

peripheral nervous system The peripheral nervous system (PNS) is one of two components that make up the nervous system of bilateral animals, with the other part being the central nervous system (CNS). The PNS consists of nerves and ganglia, which lie outside the brain ...

adrenal medulla The adrenal medulla ( la, medulla glandulae suprarenalis) is part of the adrenal gland. It is located at the center of the gland, being surrounded by the adrenal cortex. It is the innermost part of the adrenal gland, consisting of chromaffin cel ...

melanocytes Melanocytes are melanin-producing neural crest-derived cells located in the bottom layer (the stratum basale) of the skin's epidermis, the middle layer of the eye (the uvea), the inner ear, vaginal epithelium, meninges, bones, and heart. ...

, facial cartilage. The neural tube of the ectoderm develops into:

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 a v ...

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 ...

, posterior pituitary, motor neurons,

retina The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which then ...

. Note: The anterior pituitary develops from the ectodermal tissue of Rathke's pouch.

Neural crest

Because of its great importance, the neural crest is sometimes considered a fourth germ layer. It is, however, derived from the ectoderm.

References

{{DEFAULTSORT:Germ Layer Embryology Developmental biology Gastrulation

History

Evolution

Development

The germ layers

Endoderm

Mesoderm

Ectoderm

Neural crest

See also

References