Anatomical terms of location
Standard anatomical terms of location deal unambiguously with the anatomy of animals, including humans. All vertebrates have the same basic body plan – they are bilaterally symmetrical in early embryonic stages and bilaterally symmetrical in adulthood; that is, they have mirror-image left and right halves if divided down the middle. For these reasons, the basic directional terms can be considered to be those used in vertebrates. By extension, the same terms are used for many other organisms as well. While these terms are standardized within specific fields of biology, there are unavoidable, sometimes dramatic, differences between some disciplines. For example, differences in terminology remain a problem that, to some extent, still separates the terminology of human anatomy from that used in the study of various other zoological categories. Standardized anatomical and zoological terms of location have been developed based on Latin and Greek words, to enable all biological and medical scientists to delineate and communicate information about animal bodies and their component organs though the meaning of some of the terms is context-sensitive.
The vertebrates and Craniata share a substantial heritage and common structure, so many of the same terms are used for location. To avoid ambiguities this terminology is based on the anatomy of each animal in a standard way. For humans, one type of vertebrate, anatomical terms may differ from other forms of vertebrates. For one reason, this is because humans have a different neuraxis and, unlike animals that rest on four limbs, humans are considered when describing anatomy as being in the standard anatomical position, thus what is on "top" of a human is the head, whereas the "top" of a dog may be its back, the "top" of a flounder could refer to either its left or its right side. For invertebrates, standard application of locational terminology becomes difficult or debatable at best when the differences in morphology are so radical that common concepts are not homologous and do not refer to common concepts. For example, many species are not bilaterally symmetrical. In these species, terminology depends on their type of symmetry.
Because animals can change orientation with respect to their environment, because appendages like limbs and tentacles can change position with respect to the main body, positional descriptive terms need to refer to the animal as in its standard anatomical position. All descriptions are with respect to the organism in its standard anatomical position when the organism in question has appendages in another position; this helps avoid confusion in terminology. In humans, this refers to the body in a standing position with arms at the side and palms facing forward. While the universal vertebrate terminology used in veterinary medicine would work in human medicine, the human terms are thought to be too well established to be worth changing. Many anatomical terms can be combined, either to indicate a position in two axes or to indicate the direction of a movement relative to the body. For example, "anterolateral" indicates a position, both anterior and lateral to the body axis. In radiology, an X-ray image may be said to be "anteroposterior", indicating that the beam of X-rays pass from their source to patient's anterior body wall through the body to exit through posterior body wall.
There is no definite limit to the contexts in which terms may be modified to qualify each other in such combinations. The modifier term is truncated and an "o" or an "i" is added in prefixing it to the qualified term. For example, a view of an animal from an aspect at once dorsal and lateral might be called a "dorsolateral" view. Again, in describing the morphology of an organ or habitus of an animal such as many of the Platyhelminthes, one might speak of it as "dorsiventrally" flattened as opposed to bilaterally flattened animals such as ocean sunfish. Where desirable three or more terms may be agglutinated or concatenated, as in "anteriodorsolateral"; such terms sometimes used to be hyphenated. There is however little basis for any strict rule to interfere with choice of convenience in such usage. Three basic reference planes are used to describe location; the sagittal plane is a plane parallel to the sagittal suture. All other sagittal planes are parallel to it, it is known as a "longitudinal plane".
The plane is perpendicular to the ground. The median plane or midsagittal plane is in the midline of the body, divides the body into left and right portions; this passes through the head, spinal cord, and, in many animals, the tail. The term "median plane" can refer to the midsagittal plane of other structures, such as a digit; the frontal plane or coronal plane divides the body into ventral portions. For post-embryonic humans a coronal plane is vertical and a transverse plane is horizontal, but for embryos and quadrupeds a coronal plane is horizontal and a transverse plane is vertical. A longitudinal plane is any plane perpendicular to the transverse plane; the coronal plane and the sagittal plane are examples of longitudinal planes. A transverse plane known as a cross-section, divides the body into cranial and caudal portions. In human anatomy: A transverse plane is an X-Z plane, parallel to the ground, which s
The vertebral column known as the backbone or spine, is part of the axial skeleton. The vertebral column is the defining characteristic of a vertebrate in which the notochord found in all chordates has been replaced by a segmented series of bone: vertebrae separated by intervertebral discs; the vertebral column houses a cavity that encloses and protects the spinal cord. There are about 50,000 species of animals; the human vertebral column is one of the most-studied examples. In a human's vertebral column there are thirty-three vertebrae; the articulating vertebrae are named according to their region of the spine. There are twelve thoracic vertebrae and five lumbar vertebrae; the number of vertebrae in a region overall the number remains the same. The number of those in the cervical region however is only changed. There are ligaments extending the length of the column at the front and the back, in between the vertebrae joining the spinous processes, the transverse processes and the vertebral laminae.
The vertebrae in the human vertebral column are divided into different regions, which correspond to the curves of the spinal column. The articulating vertebrae are named according to their region of the spine. Vertebrae in these regions are alike, with minor variation; these regions are called the cervical spine, thoracic spine, lumbar spine and coccyx. There are twelve thoracic vertebrae and five lumbar vertebrae; the number of vertebrae in a region overall the number remains the same. The number of those in the cervical region however is only changed; the vertebrae of the cervical and lumbar spines are independent bones, quite similar. The vertebrae of the sacrum and coccyx are fused and unable to move independently. Two special vertebrae are the axis, on which the head rests. A typical vertebra consists of two parts: the vertebral arch; the vertebral arch is posterior. Together, these enclose the vertebral foramen; because the spinal cord ends in the lumbar spine, the sacrum and coccyx are fused, they do not contain a central foramen.
The vertebral arch is formed by a pair of pedicles and a pair of laminae, supports seven processes, four articular, two transverse, one spinous, the latter being known as the neural spine. Two transverse processes and one spinous process are posterior to the vertebral body; the spinous process comes out the back, one transverse process comes out the left, one on the right. The spinous processes of the cervical and lumbar regions can be felt through the skin. Above and below each vertebra are joints called facet joints; these restrict the range of movement possible, are joined by a thin portion of the neural arch called the pars interarticularis. In between each pair of vertebrae are two small holes called intervertebral foramina; the spinal nerves leave the spinal cord through these holes. Individual vertebrae are named according to their position. From top to bottom, the vertebrae are: Cervical spine: 7 vertebrae Thoracic spine: 12 vertebrae Lumbar spine: 5 vertebrae Sacrum: 5 vertebrae Coccyx: 4 vertebrae The upper cervical spine has a curve, convex forward, that begins at the axis at the apex of the odontoid process or dens, ends at the middle of the second thoracic vertebra.
This inward curve is known as a lordotic curve. The thoracic curve, concave forward, begins at the middle of the second and ends at the middle of the twelfth thoracic vertebra, its most prominent point behind corresponds to the spinous process of the seventh thoracic vertebra. This curve is known as a kyphotic curve; the lumbar curve is more marked in the female than in the male. It is convex anteriorly, the convexity of the lower three vertebrae being much greater than that of the upper two; this curve is described as a lordotic curve. The sacral curve begins at the sacrovertebral articulation, ends at the point of the coccyx; the thoracic and sacral kyphotic curves are termed primary curves, because they are present in the fetus. The cervical and lumbar curves are compensatory or secondary, are developed after birth; the cervical curve forms when the infant is able to sit upright. The lumbar curve forms from twelve to eighteen months, when the child begins to walk. Anterior surfaceWhen viewed from in front, the width of the bodies of the vertebrae is seen to increase from the second cervical to the first thoracic.
From this point there is a rapid diminution, to the apex of the coccyx. Posterior surfaceFrom behind, the vertebral column presents in the median line the spinous processes. In the cervical region these are short and bifid. In the upper part of the thoracic region they are directed obliquely downward.
A chordate is an animal constituting the phylum Chordata. During some period of their life cycle, chordates possess a notochord, a dorsal nerve cord, pharyngeal slits, an endostyle, a post-anal tail: these five anatomical features define this phylum. Chordates are bilaterally symmetric; the Chordata and Ambulacraria together form the superphylum Deuterostomia. Chordates are divided into three subphyla: Vertebrata. There are extinct taxa such as the Vetulicolia. Hemichordata has been presented as a fourth chordate subphylum, but now is treated as a separate phylum: hemichordates and Echinodermata form the Ambulacraria, the sister phylum of the Chordates. Of the more than 65,000 living species of chordates, about half are bony fish that are members of the superclass Osteichthyes. Chordate fossils have been found from as early as the Cambrian explosion, 541 million years ago. Cladistically, vertebrates - chordates with the notochord replaced by a vertebral column during development - are considered to be a subgroup of the clade Craniata, which consists of chordates with a skull.
The Craniata and Tunicata compose the clade Olfactores. Chordates form a phylum of animals that are defined by having at some stage in their lives all of the following anatomical features: A notochord, a stiff rod of cartilage that extends along the inside of the body. Among the vertebrate sub-group of chordates the notochord develops into the spine, in wholly aquatic species this helps the animal to swim by flexing its tail. A dorsal neural tube. In fish and other vertebrates, this develops into the spinal cord, the main communications trunk of the nervous system. Pharyngeal slits; the pharynx is the part of the throat behind the mouth. In fish, the slits are modified to form gills, but in some other chordates they are part of a filter-feeding system that extracts particles of food from the water in which the animals live. Post-anal tail. A muscular tail that extends backwards behind the anus. An endostyle; this is a groove in the ventral wall of the pharynx. In filter-feeding species it produces mucus to gather food particles, which helps in transporting food to the esophagus.
It stores iodine, may be a precursor of the vertebrate thyroid gland. There are soft constraints that separate chordates from certain other biological lineages, but are not part of the formal definition: All chordates are deuterostomes; this means. All chordates are based on a bilateral body plan. All chordates are coelomates, have a fluid filled body cavity called a coelom with a complete lining called peritoneum derived from mesoderm; the following schema is from the third edition of Vertebrate Palaeontology. The invertebrate chordate classes are from Fishes of the World. While it is structured so as to reflect evolutionary relationships, it retains the traditional ranks used in Linnaean taxonomy. Phylum Chordata †Vetulicolia? Subphylum Cephalochordata – Class Leptocardii Clade Olfactores Subphylum Tunicata – Class Ascidiacea Class Thaliacea Class Appendicularia Class Sorberacea Subphylum Vertebrata Infraphylum incertae sedis Cyclostomata Superclass'Agnatha' paraphyletic Class Myxini Class Petromyzontida or Hyperoartia Class †Conodonta Class †Myllokunmingiida Class †Pteraspidomorphi Class †Thelodonti Class †Anaspida Class †Cephalaspidomorphi Infraphylum Gnathostomata Class †Placodermi Class Chondrichthyes Class †Acanthodii Superclass Osteichthyes Class Actinopterygii Class Sarcopterygii Superclass Tetrapoda Class Amphibia Class Sauropsida Class Synapsida Craniates, one of the three subdivisions of chordates, all have distinct skulls.
They include the hagfish. Michael J. Benton commented that "craniates are characterized by their heads, just as chordates, or all deuterostomes, are by their tails". Most craniates are vertebrates; these consist of a series of bony or cartilaginous cylindrical vertebrae with neural arches that protect the spinal cord, with projections that link the vertebrae. However hagfish have incomplete braincases and no vertebrae, are therefore not regarded as vertebrates, but as members of the craniates, the group from which vertebrates are thought to have evolved; however the cladistic exclusion of hagfish from the vertebrates is controversial, as they ma
A motor neuron is a neuron whose cell body is located in the motor cortex, brainstem or the spinal cord, whose axon projects to the spinal cord or outside of the spinal cord to directly or indirectly control effector organs muscles and glands. There are two types of motor neuron -- lower motor neurons. Axons from upper motor neurons synapse onto interneurons in the spinal cord and directly onto lower motor neurons; the axons from the lower motor neurons are efferent nerve fibers that carry signals from the spinal cord to the effectors. Types of lower motor neurons are alpha motor neurons, beta motor neurons, gamma motor neurons. A single motor neuron may innervate many muscle fibres and a muscle fibre can undergo many action potentials in the time taken for a single muscle twitch; as a result, if an action potential arrives before a twitch has completed, the twitches can superimpose on one another, either through summation or a tetanic contraction. In summation, the muscle is stimulated repetitively such that additional action potentials coming from the somatic nervous system arrive before the end of the twitch.
The twitches thus superimpose on one another, leading to a force greater than that of a single twitch. A tetanic contraction is caused by constant high frequency stimulation - the action potentials come at such a rapid rate that individual twitches are indistinguishable, tension rises smoothly reaching a plateau. Motor neurons begin to develop early in embryonic development, motor function continues to develop well into childhood. In the neural tube cells are specified to either ventral-dorsal axis; the axons of motor neurons begin to appear in the fourth week of development from the ventral region of the ventral-dorsal axis. This homeodomain is known as the motor neural progenitor domain. Transcription factors here include Pax6, OLIG2, Nkx-6.1, Nkx-6.2, which are regulated by sonic hedgehog. The OLIG2 gene being the most important due to its role in promoting Ngn2 expression, a gene that causes cell cycle exiting as well as promoting further transcription factors associated with motor neuron development.
Further specification of motor neurons occurs when retinoic acid, fibroblast growth factor, TGFb, are integrated into the various Hox transcription factors. There are 13 Hox transcription factors and along with the signals, determine whether a motor neuron will be more rostral or caudal in character. In the spinal column, Hox 4-11 sort motor neurons to one of the five motor columns. Upper motor neurons originate in the motor cortex located in the precentral gyrus; the cells that make up the primary motor cortex are Betz cells. The axons of these cells descend from the cortex to form the corticospinal tract. Corticomotorneurons are neurons in the primary cortex which project directly to motor neurons in the ventral horn of the spinal cord. Axons of corticomotorneurons terminate on the spinal motor neurons of multiple muscles as well as on spinal interneurons, they are unique to primates and it has been suggested that their function is the adaptive control of the distal extremities including the independent control of individual fingers.
Corticomotorneurons have so far only been found in the primary motor cortex and not in secondary motor areas. Nerve tracts are bundles of axons as white matter. In the spinal cord these descending tracts carry impulses from different regions; these tracts serve as the place of origin for lower motor neurons. There are seven major descending motor tracts to be found in the spinal cord: Lateral corticospinal tract Rubrospinal tract Lateral reticulospinal tract Vestibulospinal tract Medial reticulospinal tract Tectospinal tract Anterior corticospinal tract Lower motor neurons are those that originate in the spinal cord and directly or indirectly innervate effector targets; the target of these neurons varies, but in the somatic nervous system the target will be some sort of muscle fiber. There are three primary categories lower motor neurons, which can be further divided in sub-categories. According to their targets, motor neurons are classified into three broad categories: Somatic motor neurons Special visceral motor neurons General visceral motor neurons Somatic motor neurons originate in the central nervous system, project their axons to skeletal muscles, which are involved in locomotion.
The three types of these neurons are the alpha efferent neurons, beta efferent neurons, gamma efferent neurons. They are called efferent to indicate the flow of information from the central nervous system to the periphery. Alpha motor neurons innervate extrafusal muscle fibers, which are the main force-generating component of a muscle, their cell bodies are in the ventral horn of the spinal cord and they are sometimes called ventral horn cells. A single motor neuron may synapse with 150 muscle fibers on average; the motor neuron and all of the muscle fibers to which it connects is a motor unit. Motor units are split up into 3 categories: Main Article: Motor Unit Slow motor units stimulate small muscle fibers, which contract slowly and provide small amounts of energy but are resistant to fatigue, so they are used to sustain muscular contraction, such as keeping the body upright, they gain their energy via oxidative hence require oxygen. They are called red fibers. Fast fatiguing motor units stimulate larger muscle groups, which apply large amounts of force but fatigue quickly.
They are used for tasks that require large brief bursts on energy, such as jumping or
In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring by sexual reproduction. Other ways of defining species include their karyotype, DNA sequence, behaviour or ecological niche. In addition, paleontologists use the concept of the chronospecies since fossil reproduction cannot be examined. While these definitions may seem adequate, when looked at more they represent problematic species concepts. For example, the boundaries between related species become unclear with hybridisation, in a species complex of hundreds of similar microspecies, in a ring species. Among organisms that reproduce only asexually, the concept of a reproductive species breaks down, each clone is a microspecies. All species are given a two-part name, a "binomial"; the first part of a binomial is the genus.
The second part is called the specific epithet. For example, Boa constrictor is one of four species of the genus Boa. None of these is satisfactory definitions, but scientists and conservationists need a species definition which allows them to work, regardless of the theoretical difficulties. If species were fixed and distinct from one another, there would be no problem, but evolutionary processes cause species to change continually, to grade into one another. Species were seen from the time of Aristotle until the 18th century as fixed kinds that could be arranged in a hierarchy, the great chain of being. In the 19th century, biologists grasped. Charles Darwin's 1859 book The Origin of Species explained how species could arise by natural selection; that understanding was extended in the 20th century through genetics and population ecology. Genetic variability arises from mutations and recombination, while organisms themselves are mobile, leading to geographical isolation and genetic drift with varying selection pressures.
Genes can sometimes be exchanged between species by horizontal gene transfer. Viruses are a special case, driven by a balance of mutation and selection, can be treated as quasispecies. Biologists and taxonomists have made many attempts to define species, beginning from morphology and moving towards genetics. Early taxonomists such as Linnaeus had no option but to describe what they saw: this was formalised as the typological or morphological species concept. Ernst Mayr emphasised reproductive isolation, but this, like other species concepts, is hard or impossible to test. Biologists have tried to refine Mayr's definition with the recognition and cohesion concepts, among others. Many of the concepts are quite similar or overlap, so they are not easy to count: the biologist R. L. Mayden recorded about 24 concepts, the philosopher of science John Wilkins counted 26. Wilkins further grouped the species concepts into seven basic kinds of concepts: agamospecies for asexual organisms biospecies for reproductively isolated sexual organisms ecospecies based on ecological niches evolutionary species based on lineage genetic species based on gene pool morphospecies based on form or phenotype and taxonomic species, a species as determined by a taxonomist.
A typological species is a group of organisms in which individuals conform to certain fixed properties, so that pre-literate people recognise the same taxon as do modern taxonomists. The clusters of variations or phenotypes within specimens would differentiate the species; this method was used as a "classical" method of determining species, such as with Linnaeus early in evolutionary theory. However, different phenotypes are not different species. Species named in this manner are called morphospecies. In the 1970s, Robert R. Sokal, Theodore J. Crovello and Peter Sneath proposed a variation on this, a phenetic species, defined as a set of organisms with a similar phenotype to each other, but a different phenotype from other sets of organisms, it differs from the morphological species concept in including a numerical measure of distance or similarity to cluster entities based on multivariate comparisons of a reasonably large number of phenotypic traits. A mate-recognition species is a group of sexually reproducing organisms that recognize one another as potential mates.
Expanding on this to allow for post-mating isolation, a cohesion species is the most inclusive population of individuals having the potential for phenotypic cohesion through intrinsic cohesion mechanisms. A further development of the recognition concept is provided by the biosemiotic concept of species. In microbiology, genes can move even between distantly related bacteria extending to the whole bacterial domain; as a rule of thumb, microbiologists have assumed that kinds of Bacteria or Archaea with 16S ribosomal RNA gene sequences more similar than 97% to each other need to be checked by DNA-DNA hybridisation to decide if they belong to the same species or not. This concept was narrowed in 2006 to a similarity of 98.7%. DNA-DNA hybri
The cell is the basic structural and biological unit of all known living organisms. A cell is the smallest unit of life. Cells are called the "building blocks of life"; the study of cells is called cellular biology. Cells consist of cytoplasm enclosed within a membrane, which contains many biomolecules such as proteins and nucleic acids. Organisms can be classified as multicellular; the number of cells in plants and animals varies from species to species, it has been estimated that humans contain somewhere around 40 trillion cells. Most plant and animal cells are visible only under a microscope, with dimensions between 1 and 100 micrometres. Cells were discovered by Robert Hooke in 1665, who named them for their resemblance to cells inhabited by Christian monks in a monastery. Cell theory, first developed in 1839 by Matthias Jakob Schleiden and Theodor Schwann, states that all organisms are composed of one or more cells, that cells are the fundamental unit of structure and function in all living organisms, that all cells come from pre-existing cells.
Cells emerged on Earth at least 3.5 billion years ago. Cells are of two types: eukaryotic, which contain a nucleus, prokaryotic, which do not. Prokaryotes are single-celled organisms, while eukaryotes can be either single-celled or multicellular. Prokaryotes include two of the three domains of life. Prokaryotic cells were the first form of life on Earth, characterised by having vital biological processes including cell signaling, they are simpler and smaller than eukaryotic cells, lack membrane-bound organelles such as a nucleus. The DNA of a prokaryotic cell consists of a single chromosome, in direct contact with the cytoplasm; the nuclear region in the cytoplasm is called the nucleoid. Most prokaryotes are the smallest of all organisms ranging from 0.5 to 2.0 µm in diameter. A prokaryotic cell has three architectural regions: Enclosing the cell is the cell envelope – consisting of a plasma membrane covered by a cell wall which, for some bacteria, may be further covered by a third layer called a capsule.
Though most prokaryotes have both a cell membrane and a cell wall, there are exceptions such as Mycoplasma and Thermoplasma which only possess the cell membrane layer. The envelope gives rigidity to the cell and separates the interior of the cell from its environment, serving as a protective filter; the cell wall consists of peptidoglycan in bacteria, acts as an additional barrier against exterior forces. It prevents the cell from expanding and bursting from osmotic pressure due to a hypotonic environment; some eukaryotic cells have a cell wall. Inside the cell is the cytoplasmic region that contains the genome and various sorts of inclusions; the genetic material is found in the cytoplasm. Prokaryotes can carry extrachromosomal DNA elements called plasmids, which are circular. Linear bacterial plasmids have been identified in several species of spirochete bacteria, including members of the genus Borrelia notably Borrelia burgdorferi, which causes Lyme disease. Though not forming a nucleus, the DNA is condensed in a nucleoid.
Plasmids encode additional genes, such as antibiotic resistance genes. On the outside and pili project from the cell's surface; these are structures made of proteins that facilitate communication between cells. Plants, fungi, slime moulds and algae are all eukaryotic; these cells are about fifteen times wider than a typical prokaryote and can be as much as a thousand times greater in volume. The main distinguishing feature of eukaryotes as compared to prokaryotes is compartmentalization: the presence of membrane-bound organelles in which specific activities take place. Most important among these is a cell nucleus, an organelle that houses the cell's DNA; this nucleus gives the eukaryote its name, which means "true kernel". Other differences include: The plasma membrane resembles that of prokaryotes in function, with minor differences in the setup. Cell walls may not be present; the eukaryotic DNA is organized in one or more linear molecules, called chromosomes, which are associated with histone proteins.
All chromosomal DNA is stored in the cell nucleus, separated from the cytoplasm by a membrane. Some eukaryotic organelles such as mitochondria contain some DNA. Many eukaryotic cells are ciliated with primary cilia. Primary cilia play important roles in chemosensation and thermosensation. Cilia may thus be "viewed as a sensory cellular antennae that coordinates a large number of cellular signaling pathways, sometimes coupling the signaling to ciliary motility or alternatively to cell division and differentiation." Motile eukaryotes can move using motile flagella. Motile cells are absent in flowering plants. Eukaryotic flagella are more complex than those of prokaryotes. All cells, whether prokaryotic or eukaryotic, have a membrane that envelops the cell, regulates what moves in and out, maintains the electric potential of the cell. Inside the membrane, the cytoplasm takes up most of the cell's volume. All cells possess DNA, the hereditary material of genes, RNA, containing the information necessary to build various proteins such as enzymes, the cell's primary machinery.
There are other kinds of biomolecules in cells. This article lists these primary cellular components briefly
In amniote animal embryology, the epiblast is one of two distinct layers arising from the inner cell mass in the mammalian blastocyst or from the blastodisc in reptiles and birds. It derives the embryo proper through its differentiation into the three primary germ layers, ectoderm and endoderm, during gastrulation; the amnionic ectoderm and extraembryonic mesoderm originate from the epiblast. In mammalian embryogenesis and segregation of cells composing the inner cell mass of the blastocyst yields two distinct layers—the epiblast and the hypoblast. While the cuboidal hypoblast cells delaminate ventrally, away from the embryonic pole, to line the blastocoele, the remaining cells of the inner cell mass, situated between the hypoblast and the polar trophoblast, become the epiblast and comprise columnar cells. In the mouse, primordial germ cells are specified from epiblast cells; this specification is accompanied by extensive epigenetic reprogramming that involves global DNA demethylation, chromatin reorganization and imprint erasure leading to totipotency.
The DNA base excision repair pathway has a central role in the process of genome-wide demethylation. Upon commencement of gastrulation, the primitive streak, a visible, morphological groove, appears on the posterior epiblast and orients along the anterior-posterior embryo axis. Initiated by signals from the underlying hypoblast, formation of the primitive streak is predicated on epiblast cell migration, mediated by Nodal, from the lateral-posterior regions of the epiblast to the center midline; the primitive knot is situated at the anterior end of the primitive streak and serves as the organizer for gastrulation, determining epiblast cell fate by inducing the differentiation of migrating epiblast cells during gastrulation. During gastrulation, migrating epiblast cells undergo epithelial-mesenchymal transition in order to lose cell-cell adhesion, delaminate from the epiblast layer and migrate over the dorsal surface of the epiblast down through the primitive streak; the first wave of epiblast cells to invaginate through the primitive streak invades and displaces the hypoblast to become the embryonic endoderm.
The mesoderm layer is established next as migrating epiblast cells move through the primitive streak spread out within the space between the endoderm and remaining epiblast, which once the mesoderm layer has formed becomes the definitive ectoderm. The process of gastrulation results in a trilaminar germ disc, consisting of the ectoderm and endoderm layers. Epiblasts exhibit diverse structure across species as a result of early embryo morphogenesis; the human epiblast assumes a disc shape. Gastrulation occurs in the epiblast of avian embryos. A local thickening of the epiblast, known as Koller's sickle, is key in inducing the primitive streak, the structure through which gastrulation occurs. Embryogenesis Human embryogenesis Hypoblast http://www.embryology.ch/allemand/iperiodembry/carnegie02.html http://www.med.umich.edu/lrc/coursepages/M1/embryology/embryo/04secondweek.htm https://web.archive.org/web/20080103050509/http://isc.temple.edu/marino/embryology/EMBII97/sld005.htm