In avian gastrulation, Koller's sickle is a local thickening of cells at the posterior edge of the upper layer of the area pellucida called the epiblast. Koller's sickle is crucial for avian development, due to its critical role in inducing the differentiation of various avian body parts. Koller's sickle induces primitive streak and Hensen's node, which are major components of avian gastrulation. Avian gastrulation is a process by which developing cells in an avian embryo move relative to one another in order to form the three germ layers (endoderm, mesoderm, and ectoderm).
The thickening of the epiblast in Koller's sickle acts as a margin separating sheets of cells from posterior side of avian blastoderms from hypoblasts and area opaca endoderm. The blastoderm is a single layer of cells, and the hypoblast and area opaca endoderm cells lie directly below the blastoderm. Koller's sickle arises from the midpoint, between the hypoblast cells and the area opaca endoderm. As blastoderm cells migrate anteriorly they push primary hypoblast cells and form a secondary hypoblast known as the endoblast. Also during this migration, Koller's sickle prevents the hypoblast cells and the area opaca cells from making contact with the blastoderm, which allows the primitive streak to form. [2] [3] [4]
The primitive streak is induced by the posterior marginal zone (PMZ) of Koller's sickle, which can also induce Hensen's node. If cell movement in the PMZ is blocked, the primitive streak does not form. [5] Thus, the PMZ acts as an organizer. [6] Cells in marginal zones of the embryo, like the PMZ, are key to development and cell fate determination in chick embryos.
Avian gastrulation occurs as cells move though the primitive streak. Hence, primitive streak is analogous to the blastopore lip in amphibian gastrulation. [2] The primitive streak develops from Koller's sickle and the epiblast of the avian embryo. As the cells of Koller's sickle migrate during gastrulation, they form different portions of the primitive streak. The anterior cells of Koller's sickle become the anterior region of the primitive streak, known as Hensen's node. Similarly, the posterior cells of Koller's sickle form the posterior region of the primitive streak. [2] This differential movement is due to expression of different mesodermal marker genes among the cells located in different areas of Koller's sickle. Chordin is expressed in cells of the anterior streak, while Wnt8c is expressed in cells of the posterior streak. [1] The movement is coordinated by a Wnt signaling pathway which is activated by fibroblast growth factors from the hypoblast. [2]
The primitive streak is key in the development of the major body axes. The primitive groove forms as a depression in the primitive streak as it is developing, and allows a space for migrating cells to move into the deeper layers of the embryo. Cells migrate by entering through the dorsal side and moving toward the ventral side of the avian embryo, separating the left and right sections of the embryo. The primitive pit in Hensen's node, at the anterior end of the primitive streak, allows cells to enter which will form the notochord and prechordal plate. Cells that move through the center of the streak will become the heart and kidneys. The lateral plate and the extraembryonic mesoderm arise from the cells that enter at the posterior end of the primitive streak. Epiblast cells near the primitive streak form the neural plate and other dorsal structures, while the epiblast cells far from the streak become epidermis. [2]
Koller's sickle is one of two regions (the other being the caudal boundary region of the area opaca) where expression patterns for genes important for gastrulation are localized. For example, the gene Nodal is only expressed in Koller's sickle. [7]
While a single gene has not been isolated for the creation of Koller's sickle, there is evidence that the Homeobox gene Hex influences Koller's sickle development. The transcript cHex, which is a product of Hex, has been detected in Koller's sickle during chick embryogenesis. cHex is also involved with the formation of the hypoblast, the endoderm in an anterior arc that overlaps the cardiogenic region, pharyngeal endoderm immediately adjacent to the forming myocardium, in the endocardium, and in the liver and thyroid gland primordia. [8]
It is also possible that the Homeobox gene goosecoid (GSC) is involved in the formation of Koller's sickle, as Koller's sickle cells are the first to express the goosecoid transcript. In general, the goosecoid gene is thought to be involved in the development of the chicken organizer during gastrulation. [9]
Koller's sickle was originally described by August Rauber in 1876. Because of this Koller's sickle is sometimes referred to as Rauber's sickle. [10]
In 1926, Ludwig Graper first studied the involvement of Koller's sickle in the formation of the primitive streak. The cell movements reminded him of a dance called the Polonaise, in which dancers moved in parallel lines and in which they move from the back of the group to the center. It was not until 2007 that the mechanism for these movements was discovered, by Voiculescu and his associates. They determined that cells move to the center of the epiblast following the activation of the Wnt planar cell polarity pathway by fibroblast growth factors made by the hypoblast. [2]
There is still a lot that is unknown regarding Koller's sickle, but research is ongoing. By implanting a fragment of quail Koller's sickle into a chicken blastoderm, Drs. Callebaut and Van Nueten observed the formation of a normal secondary primitive streak, mesoderm, and definitive endoderm. This led them to the conclusion that Koller's sickle is the early avian representation of the organizer, and that there is homology between Koller's sickle in avians and the blastoporus in amphibians. [11] Drs. Callebaut and Van Nueten also optimized a method for preparation of unincubated avian eggs, and from this they demonstrated the fact that embryonic regulation is a result of the spatial distribution of Koller's sickle tissue. [12] Additionally, Drs. Callebaut and Van Nueten were able to determine that the differentiation of Koller' sickle cells to sickle endoblast is irreversible, and that the sickle endoblast induces early neurulation; they did this by implanting Koller's sickle tissue into different parts of unincubated chicken blastoderms and observing the effects. [13]
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.
Gastrulation is the stage in the early embryonic development of most animals, during which the blastula, or in mammals the blastocyst is reorganized into a multilayered structure known as the gastrula. Before gastrulation, the embryo is a continuous epithelial sheet of cells; by the end of gastrulation, the embryo has begun differentiation to establish distinct cell lineages, set up the basic axes of the body, and internalized one or more cell types including the prospective gut.
Drosophila embryogenesis, the process by which Drosophila embryos form, is a favorite model system for genetics and developmental biology. The study of its embryogenesis unlocked the century-long puzzle of how development was controlled, creating the field of evolutionary developmental biology. The small size, short generation time, and large brood size make it ideal for genetic studies. Transparent embryos facilitate developmental studies. Drosophila melanogaster was introduced into the field of genetic experiments by Thomas Hunt Morgan in 1909.
The blastocoel, also spelled blastocoele and blastocele, and also called cleavage cavity, or segmentation cavity is a fluid-filled or yolk-filled cavity that forms in the blastula during very early embryonic development. At this stage in mammals the blastula develops into the blastocyst containing an inner cell mass, and outer trophectoderm.
A germ layer is a primary layer of cells that forms during embryonic development. The three germ layers in vertebrates are particularly pronounced; however, all eumetazoans produce two or three primary germ layers. Some animals, like cnidarians, produce two germ layers making them diploblastic. Other animals such as bilaterians produce a third layer 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.
The archenteron, also called the gastrocoel or the primitive digestive tube, is the internal cavity of the primitive gastrointestinal tract that forms during gastrulation in a developing animal embryo. It develops into the endoderm and mesoderm of the animal.
In developmental biology, animal embryonic development, also known as animal embryogenesis, is the developmental stage of an animal embryo. Embryonic development starts with the fertilization of an egg cell (ovum) by a sperm cell, (spermatozoon). Once fertilized, the ovum becomes a single diploid cell known as a zygote. The zygote undergoes mitotic divisions with no significant growth and cellular differentiation, leading to development of a multicellular embryo after passing through an organizational checkpoint during mid-embryogenesis. In mammals, the term refers chiefly to the early stages of prenatal development, whereas the terms fetus and fetal development describe later stages.
The primitive node is the organizer for gastrulation in most amniote embryos. In birds it is known as Hensen's node, and in amphibians it is known as the Spemann-Mangold organizer. It is induced by the Nieuwkoop center in amphibians, or by the posterior marginal zone in amniotes including birds.
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 broad end towards the anterior, and the narrower region projected to the posterior. The primitive streak forms a longitudinal midline structure in the narrower posterior (caudal) region of the developing embryo on its dorsal side. At first formation the primitive streak extends for half the length of the embryo. In the human embryo this appears by stage 6, about 17 days.
In amniote embryonic development, the epiblast 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 the hypoblast. It derives the embryo proper through its differentiation into the three primary germ layers, ectoderm, mesoderm and endoderm, during gastrulation. The amnionic ectoderm and extraembryonic mesoderm also originate from the epiblast.
Chordin is a protein with a prominent role in dorsal–ventral patterning during early embryonic development. In humans it is encoded for by the CHRD gene.
The bilaminar embryonic disc, bilaminar blastoderm or embryonic disc is the distinct two-layered structure of cells formed by day eight of the development of a human embryo. It is formed when the inner cell mass, also known as the embryoblast, forms a bilaminar disc of two layers, an upper layer called the epiblast and a lower layer called the hypoblast, which will eventually form into fetus. These two layers of cells are stretched between two fluid-filled cavities at either end: the primitive yolk sac and the amniotic sac.
In the field of developmental biology, regional differentiation is the process by which different areas are identified in the development of the early embryo. The process by which the cells become specified differs between organisms.
The development of fishes is unique in some specific aspects compared to the development of other animals.
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 development. In biological terms, the development of the human body entails growth from a one-celled zygote to an adult human being. Fertilization occurs when the sperm cell successfully enters and fuses with an egg cell (ovum). The genetic material of the sperm and egg then combine to form the single cell zygote and the germinal stage of development commences. Embryonic development in the human, covers the first eight weeks of development; at the beginning of the ninth week the embryo is termed a fetus. The eight weeks has 23 stages.
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 the chorion.
Nodal homolog is a secretory protein that in humans is encoded by the NODAL gene which is located on chromosome 10q22.1. It belongs to the transforming growth factor beta superfamily. Like many other members of this superfamily it is involved in cell differentiation in early embryogenesis, playing a key role in signal transfer from the primitive node, in the anterior primitive streak, to lateral plate mesoderm (LPM).
Homeobox protein goosecoid(GSC) is a homeobox protein that is encoded in humans by the GSC gene. Like other homeobox proteins, goosecoid functions as a transcription factor involved in morphogenesis. In Xenopus, GSC is thought to play a crucial role in the phenomenon of the Spemann-Mangold organizer. Through lineage tracing and timelapse microscopy, the effects of GSC on neighboring cell fates could be observed. In an experiment that injected cells with GSC and observed the effects of uninjected cells, GSC recruited neighboring uninjected cells in the dorsal blastopore lip of the Xenopus gastrula to form a twinned dorsal axis, suggesting that the goosecoid protein plays a role in the regulation and migration of cells during gastrulation.
Rosa Susan Penelope Beddington FRS was a British biologist whose career had a major impact on developmental biology.
Evx1 is a mammalian gene located downstream of the HoxA cluster, which encodes for a homeobox transcription factor. Evx1 is a homolog of even-skipped (eve), which is a pair-rule gene that regulates body segmentation in Drosophila. The expression of Evx1 is developmentally regulated, displaying a biphasic expression pattern with peak expression in the primitive streak during gastrulation and in interneurons during neural development. Evx1 has been shown to regulate anterior-posterior patterning during gastrulation by acting as a downstream effector of the Wnt and BMP signalling pathways. It is also a critical regulator of interneuron identity.