A developmental signaling center is defined as a group of cells that release various morphogens which can determine the fates, or destined cell types, of adjacent cells. This process in turn determines what tissues the adjacent cells will form. Throughout the years, various development signaling centers have been discovered. [1]
In 1924, Hans Spemann and Hilde Mangold discovered a region in the dorsal blastopore lip of an amphibian embryo that induced certain neighboring cells into becoming neural tissue. This Spemann-Mangold organizer was the first time that a developmental organizer region was identified and studied. [2] Since then many analogous organizers have been found in other organisms. The Spemann-Mangold organizer is important to developmental biology because it was the first proof that particular cell populations influenced the differentiation of other cells through signaling molecules. [3]
The Nieuwkoop center, named after the developmental biologist Pieter Nieuwkoop, [4] is a cluster of dorsal vegetal cells in a blastula which produce both mesoderm-inducing and dorsalizing signals. [5] Signals from the Nieuwkoop center induce the Spemann-Mangold organizer, thus the Nieuwkoop Center is known as the organizer of the organizer. Even with the BCNE center (Blastula chordin and noggin expression center) removed from the blastula, the Nieuwkoop Center is able to induce formation of the Spemann-Mangold organizer. [6] Transplant of the Nieuwkoop Center causes formation of an embryonic axis with an endodermal fate which contains dorsal mesoderm. [7]
Due to difficulty defining definitive Nieuwkoop regions, little is known about the molecular composition of the Nieuwkoop signal. However, cells from the Nieuwkoop Center express potent mesoderm inducers as well as the secreted protein, Cerberus (CER1), which contributes to the formation of the head, heart, and asymmetry of internal organs. [8] Furthermore, a homeobox gene, nieuwkoid, was named after the Nieuwkoop Center for its role in development. Nieuwkoid is expressed immediately following the mid-blastula transition to a pregastrula embryo on the dorsal side and mis-expression of nieuwkoid was found to be sufficient for induction of secondary axes. [9]
The BCNE center is the Blastula Chordin and Noggin Expressing center. The BCNE center is located in the dorsal region of the animal pole. It appears after the mid-blastula stage and is triggered by the expression of beta-catenin like the Nieuwkoop center. [8] This center is found to be distinct from the Nieuwkoop center, which secretes a different group of factors, due to expression of VegT and B1-Sox which prevents the BCNE center from extending into the vegetal pole of the blastula. [10] The BCNE center is found to secrete several factors: chordin, [8] noggin, [8] Xnr3, [8] siamois, [8] goosecoid, [11] twin, [10] Admp, [10] and FoxA4a. [10] This center predisposes cells in the blastula stage to become neural tissue. [8] The cells of the BCNE region give rise to the forebrain, most of the mid-brain and hind-brain, the notochord, and the floor plate. [8]
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 two-layered or three-layered embryo 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.
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.
Neurulation refers to the folding process in vertebrate embryos, which includes the transformation of the neural plate into the neural tube. The embryo at this stage is termed the neurula.
Somitogenesis is the process by which somites form. Somites are bilaterally paired blocks of paraxial mesoderm that form along the anterior-posterior axis of the developing embryo in segmented animals. In vertebrates, somites give rise to skeletal muscle, cartilage, tendons, endothelium, and dermis.
The neural plate is a key developmental structure that serves as the basis for the nervous system. Cranial to the primitive node of the embryonic primitive streak, ectodermal tissue thickens and flattens to become the neural plate. The region anterior to the primitive node can be generally referred to as the neural plate. Cells take on a columnar appearance in the process as they continue to lengthen and narrow. The ends of the neural plate, known as the neural folds, push the ends of the plate up and together, folding into the neural tube, a structure critical to brain and spinal cord development. This process as a whole is termed primary neurulation.
Neural crest cells are a temporary group of cells 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, peripheral and enteric neurons and glia.
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.
Bone morphogenetic protein 4 is a protein that in humans is encoded by BMP4 gene. BMP4 is found on chromosome 14q22-q23.
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.
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.
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.
The Nodal signaling pathway is a signal transduction pathway important in regional and cellular differentiation during embryonic development.
In Xenopus laevis, the specification of the three germ layers occurs at the blastula stage. Great efforts have been made to determine the factors that specify the endoderm and mesoderm. On the other hand, only a few examples of genes that are required for ectoderm specification have been described in the last decade. The first molecule identified to be required for the specification of ectoderm was the ubiquitin ligase Ectodermin ; later, it was found that the deubiquitinating enzyme, FAM/USP9x, is able to overcome the effects of ubiquitination made by Ectodermin in Smad4. Two transcription factors have been proposed to control gene expression of ectodermal specific genes: POU91/Oct3/4 and FoxIe1/Xema. A new factor specific for the ectoderm, XFDL156, has shown to be essential for suppression of mesoderm differentiation from pluripotent cells.
This article is about the role of Fibroblast Growth Factor Signaling in Mesoderm Formation.
Edward Michael De Robertis is an American embryologist and Professor at the University of California, Los Angeles. His work has contributed to the finding of conserved molecular processes of embryonic inductions that result in tissue differentiations during animal development. He was elected to the National Academy of Sciences in 2013, worked for the Howard Hughes Medical Institute for 26 years, and holds a Distinguished Professor at the University of California, Los Angeles. In 2009 Pope Benedict XVI appointed De Robertis to a lifetime position in the Pontifical Academy of Sciences, and in 2022 Pope Francis appointed him Councillor of the Academy for four years.
The Spemann-Mangold organizer is a group of cells that are responsible for the induction of the neural tissues during development in amphibian embryos. First described in 1924 by Hans Spemann and Hilde Mangold, the introduction of the organizer provided evidence that the fate of cells can be influenced by factors from other cell populations. This discovery significantly impacted the world of developmental biology and fundamentally changed the understanding of early development.
The dorsal lip of the blastopore is a structure that forms during early embryonic development and is important for its role in organizing the germ layers. The dorsal lip is formed during early gastrulation as folding of tissue along the involuting marginal zone of the blastocoel forms an opening known as the blastopore. It is particularly important for its role in neural induction through the default model, where signaling from the dorsal lip protects a region of the epiblast from becoming epidermis, thus allowing it to develop to its default neural tissue.
