A germ layer is a primary layer of

cell 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 ...

s that forms during embryonic development. 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, with c ...

s are particularly pronounced; however, all eumetazoans (

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 throug ...

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 ...

ns, produce two germ layers (the ectoderm and endoderm) making them diploblastic. Other animals such as

bilaterians 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 and ...

produce a third layer (the mesoderm) between these two layers, making them triploblastic. Germ layers eventually give rise to all of an animal’s tissues and organs through the process of

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 ...

History

Caspar Friedrich Wolff observed organization of the early embryo in leaf-like layers. In 1817,

Heinz Christian Pander Heinz Christian Pander, also Christian Heinrich Pander ( – ), was a Russian Empire ethnic Baltic German biologist and embryologist. Biography In 1817 he received his doctorate from the University of Würzburg, and spent several years (1827� ...

discovered three primordial germ layers while studying chick embryos. Between 1850 and 1855,

Robert Remak Robert Remak (26 July 1815 – 29 August 1865) was a Jewish Polish-German embryologist, physiologist, and neurologist, born in Poznań, Posen, Prussia, who discovered that the origin of cells was by the Cell division, division of pre-existing cel ...

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 " mesoderm" was introduced into English by Huxley in 1871, and " ectoderm" and " endoderm" by Lankester in 1873.

Evolution

Among

s, sponges 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 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 ...

and

Ctenophora Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), and ...

, show an increase in compartmentalization, having two germ layers, the endoderm and ectoderm. Diploblastic animals are organized into recognisable tissues. All bilaterian animals (from flatworms to humans) are triploblastic, possessing a mesoderm 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 multicell ...

. During the next stage, cleavage,

mitotic In cell biology, mitosis () is a part of the cell cycle in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis gives rise to genetically identical cells in which the total number of chromosomes is maintai ...

cell divisions transform the zygote into a hollow ball of cells, a blastula. This early embryonic form undergoes gastrulation, forming a

gastrula 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. ...

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, after about three days, the zygote forms a solid mass of cells by mitotic division, called a

morula A morula (Latin, ''morus'': mulberry) is an early-stage embryo consisting of a solid ball of cells called blastomeres, contained in mammals, and other animals within the zona pellucida shell. The blastomeres are the daughter cells of the zygot ...

. This then changes to a blastocyst, 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 p ...

, and an inner cell mass called the

embryoblast The inner cell mass (ICM) or embryoblast (known as the pluriblast in marsupials) is a structure in the early development of an embryo. It is the mass of cells inside the blastocyst that will eventually give rise to the definitive structures of th ...

. Filled with uterine fluid, the blastocyst breaks out of the

zona pellucida The zona pellucida (plural zonae pellucidae, also egg coat or pellucid zone) is a specialized extracellular matrix that surrounds the plasma membrane of mammalian oocytes. It is a vital constitutive part of the oocyte. The zona pellucida first ap ...

and undergoes implantation. The inner cell mass initially has two layers: the hypoblast and epiblast. At the end of the second week, a primitive streak appears. The epiblast in this region moves towards the primitive streak, dives down into it, and forms a new layer, called the endoderm, pushing the hypoblast out of the way (this goes on to form the amnion.) The epiblast keeps moving and forms a second layer, the mesoderm. The top layer is now called the ectoderm. Gastrulation occurs in reference to the primary body axis. Germ layer formation is linked to the primary body axis as well, however it is less reliant on it than gastrulation is. ''

Hydractinia ''Hydractinia'' is a genus of commensal athecate hydroids which belong to the family Hydractiniidae. ''Hydractinia'' species mostly live on hermit-crabbed marine gastropod shells. One species, '' Hydractinia echinata'', is commonly known as s ...

'' 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 f ...

s: Sox2 and

Oct4 Oct-4 (octamer-binding transcription factor 4), also known as POU5F1 (POU domain, class 5, transcription factor 1), is a protein that in humans is encoded by the ''POU5F1'' gene. Oct-4 is a homeodomain transcription factor of the POU family. I ...

proteins. These transcription factors cause the

pluripotent Pluripotency: These are the cells that can generate into any of the three Germ layers which imply Endodermal, Mesodermal, and Ectodermal cells except tissues like the placenta. According to Latin terms, Pluripotentia means the ability for many thin ...

mouse

embryonic stem cell Embryonic stem cells (ESCs) are pluripotent stem cells derived from the inner cell mass of a blastocyst, an early-stage pre- implantation embryo. Human embryos reach the blastocyst stage 4–5 days post fertilization, at which time they consist ...

s 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 embryonic development. Cells migrating inward along the

archenteron The primary gut that forms during gastrulation in a developing embryo is known as the archenteron, the gastrocoel or the primitive digestive tube. It develops into the endoderm and mesoderm of an animal. Formation in sea urchins As primary mesen ...

form the inner layer of the

, which develops into the endoderm. The endoderm consists at first of flattened cells, which subsequently become columnar. It forms the epithelial lining of the whole of the

digestive tract The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans and ...

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 st ...

, 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

stomach The stomach is a muscular, hollow organ in the gastrointestinal tract of humans and many other animals, including several invertebrates. The stomach has a dilated structure and functions as a vital organ in the digestive system. The stomach i ...

, the small intestine, the colon, the

liver The liver is a major organ only found in vertebrates which performs many essential biological functions such as detoxification of the organism, and the synthesis of proteins and biochemicals necessary for digestion and growth. In humans, it ...

, 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 en ...

, the bladder, 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 intercell ...

parts of the

trachea The trachea, also known as the windpipe, is a 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 from the ...

and

bronchi A bronchus is a passage or airway in the lower respiratory tract that conducts air into the lungs. The first or primary bronchi pronounced (BRAN-KAI) to branch from the trachea at the carina are the right main bronchus and the left main bronchus. ...

, 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 ...

, 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.

Mesoderm

The mesoderm germ layer forms in the embryos of triploblastic

s. During gastrulation, some of the cells migrating inward contribute to the mesoderm, an additional layer between the endoderm and the ectoderm. The formation of a mesoderm leads to the development of a coelom. 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,

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 ...

, lateral plate mesoderm, 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 Smooth may refer to: Mathematics * Smooth function, a function that is infinitely differentiable; used in calculus and topology * Smooth manifold, a differentiable manifold for which all the transition maps are smooth functions * Smooth algebrai ...

and striated),

bone A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, ...

, cartilage, connective tissue,

adipose tissue Adipose tissue, body fat, or simply fat is a loose connective tissue composed mostly of adipocytes. In addition to adipocytes, adipose tissue contains the stromal vascular fraction (SVF) of cells including preadipocytes, fibroblasts, vascular ...

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, dermis,

Dentine Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by e ...

of teeth, genitourinary system,

serous membrane The serous membrane (or serosa) is a smooth tissue membrane of mesothelium lining the contents and inner walls of body cavities, which secrete serous fluid to allow lubricated sliding movements between opposing surfaces. The serous membrane ...

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.

Ectoderm

The ectoderm generates the outer layer of the embryo, and it forms from the embryo's epiblast. The ectoderm 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 Neural crest cells are a temporary group of cells unique to vertebrates that arise from the embryonic ectoderm germ layer, and in turn give rise to a diverse cell lineage—including melanocytes, craniofacial cartilage and bone, smooth muscle, per ...

, and the

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, ...

. The surface ectoderm develops into: epidermis, hair, 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 ca ...

, sebaceous glands,

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 ...

tooth enamel Tooth enamel is one of the four major 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. The other major tissues are dentin, ...

, the epithelium of the mouth 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 ...

. 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, melanocytes, 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 ve ...

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 sp ...

posterior pituitary The posterior pituitary (or neurohypophysis) is the posterior lobe of the pituitary gland which is part of the endocrine system. The posterior pituitary is not glandular as is the anterior pituitary. Instead, it is largely a collection of axonal ...

motor neurons 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 indirectly ...

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 In embryogenesis, Rathke's pouch is an evagination at the roof of the developing mouth in front of the buccopharyngeal membrane. It gives rise to the anterior pituitary (adenohypophysis), a part of the endocrine system. Development Rathke's pou ...

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