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Developmental biology is the study of the process by which animals and plants grow and develop, and is synonymous with ontogeny. In animals most development occurs in embryonic life, but it is also found in regeneration, asexual reproduction and metamorphosis, and in the growth and differentiation of stem cells in the adult organism. In plants, development occurs in embryos, during vegetative reproduction, and in the normal outgrowth of roots, shoots and flowers. Practical outcomes from the study of animal developmental biology have included in vitro fertilization, now widely used in fertility treatment, the understanding of risks from substances that can damage the fetus (teratogens), and the creation of various animal models for human disease which are useful in research. Developmental Biology has also help to generate modern stem cell biology which promises a number of important practical benefits for human health. Many of the processes of development are now well understood, and some major textbooks of the subject are  
The main processes involved in the embryonic development of animals are: regional specification, morphogenesis, cell differentiation, growth, and the overall control of timing. Regional specification refers to the processes that create spatial pattern in a ball or sheet of initially similar cells. This generally involves the action of cytoplasmic determinants, located within parts of the fertilized egg, and of inductive signals emitted from signaling centers in the embryo. The early stages of regional specification do not generate functional differentiated cells, but cell populations committed to develop to a specific region or part of the organism. These are defined by the expression of specific combinations of transcription factors. Morphogenesis relates to the formation of three-dimensional shape. It mainly involves the orchestrated movements of cell sheets and of individual cells. Morphogenesis is important for creating the three germ layers of the early embryo (ectoderm, mesoderm and endoderm) and for building up complex structures during organ development. Cell differentiation relates specifically to the formation of functional cell types such as nerve, muscle, secretory epithelia etc. Differentiated cells contain large amounts of specific proteins associated with the cell function. Growth involves both an overall increase in size, and also the differential growth of parts (allometry) which contributes to morphogenesis. Growth mostly occurs through cell division but also through changes of cell size and the deposition of extracellular materials. The control of timing of events and the integration of the various processes with one another is the least well understood area of the subject. It remains unclear whether animal embryos contain a master clock mechanism or not. The development of plants involves similar processes to that of animals. However plant cells are mostly immotile so morphogenesis is achieved by differential growth, without cell movements. Also, the inductive signals and the genes involved in plant development are different from those that control animal development.
Developmental model organisms
Much of developmental biology research in recent decades has focused on the use of a small number of model organisms. It has turned out that there is much conservation of developmental mechanisms across the animal kingdom. In early development different vertebrate species all use essentially the same inductive signals and the same genes encoding regional identity. Even invertebrates use a similar repertoire of signals and genes although the body parts formed are significantly different. Model organisms each have some particular experimental advantages which have enabled them to become popular among researchers. In one sense they are "models" for the whole animal kingdom, and in another sense they are "models" for human development, which is difficult to study directly for both ethical and practical reasons. Model organisms have been most useful for elucidating the broad nature of developmental mechanisms. The more detail is sought, the more they differ from each other and from humans.
Frog: Xenopus (X.laevis and tropicalis) . Good embryo supply. Especially suitable for microsurgery. Zebrafish: Danio rerio. Good embryo supply. Well developed genetics. Chicken: Gallus gallus. Early stages similar to mammal, but microsurgery easier. Low cost. Mouse: Mus musculus. A mammal with well developed genetics.
Also popular for some purposes have been sea urchins and ascidians. For studies of regeneration urodele amphibians such as the axolotl Ambystoma mexicanum are used, and also planarian worms such as Schmidtea mediterranea. Plant development has focused on the thale cress Arabidopsis thaliana as a model organism.
Differentiation is the formation of cell types, from what is originally one cell – the zygote or spore. The formation of cell types such as nerve cells occurs with a number of intermediary, less differentiated cell types. A cell stays a certain cell type by maintaining a particular pattern of gene expression. This depends on regulatory genes, e.g. for transcription factors and signaling proteins. These can take part in self-perpetuating circuits in the gene regulatory network, circuits that can involve several cells that communicate with each other. External signals can alter gene expression by activating a receptor, which triggers a signaling cascade that affects transcription factors. For example, the withdrawal of growth factors from myoblasts causes them to stop dividing and instead differentiate into muscle cells.
Embryogenesis is the step in the life cycle after fertilization – the development of the embryo, starting from the zygote (fertilized egg). Organisms can differ drastically in how the embryo develops, especially when they belong to different phyla. For example, embryonal development in placental mammals starts with cleavage of the zygote into eight uncommitted cells, which then form a ball (morula). The outer cells become the trophectoderm or trophoblast, which will form in combination with maternal uterine endometrial tissue the placenta, needed for fetal nurturing via maternal blood, while inner cells become the inner cell mass that will form all fetal organs (the bridge between these two parts eventually forms the umbilical cord). In contrast, the fruit fly zygote first forms a sausage-shaped syncytium, which is still one cell but with many cell nuclei.
Patterning is important for determining which cells develop into which organs. This is mediated by signaling between adjacent cells by proteins on their surfaces, and by gradients of signaling secreted molecules. An example is retinoic acid, which forms a gradient in the head to tail direction in animals. Retinoic acid enters cells and activates Hox genes in a concentration-dependent manner – Hox genes differ in how much retinoic acid they require for activation and will thus show differential rostral expression boundaries, in a colinear fashion with their genomic order. As Hox genes code for transcription factors, this causes different activated combinations of both Hox and other genes in discrete anteroposterior transverse segments of the neural tube (neuromeres) and related patterns in surrounding tissues, such as branchial arches, lateral mesoderm, neural crest, skin and endoderm, in the head to tail direction. This is important for e.g. the segmentation of the spine in vertebrates.
Embryonic development does not always proceed correctly, and errors can result in birth defects or miscarriage. Often the reason is genetic (mutation or chromosome abnormality), but there can be environmental influence (like teratogens) or stochastic events. Abnormal development caused by mutation is also of evolutionary interest as it provides a mechanism for changes in body plan (see evolutionary developmental biology).
Growth is the enlargement of a tissue or organism. Growth continues after the embryonal stage, and occurs through cell proliferation, enlargement of cells or accumulation of extracellular material. In plants, growth results in an adult organism that is strikingly different from the embryo. The proliferating cells tend to be distinct from differentiated cells (see stem cell and progenitor cell). In some tissues proliferating cells are restricted to specialised areas, such as the growth plates of bones. But some stem cells migrate to where they are needed, such as mesenchymal stem cells which can migrate from the bone marrow to form e.g. muscle, bone or adipose tissue. The size of an organ frequently determines its growth, as in the case of the liver which grows back to its previous size if a part is removed. Growth factors, such as fibroblast growth factors in the animal embryo and growth hormone in juvenile mammals, also control the extent of growth.
Most animals have a larval stage, with a body plan different from that of the adult organism. The larva abruptly develops into an adult in a process called metamorphosis. For example, caterpillars (butterfly larvae) are specialized for feeding whereas adult butterflies (imagos) are specialised for flight and reproduction. When the caterpillar has grown enough, it turns into an immobile pupa. Here, the imago develops from imaginal discs found inside the larva.
Regeneration is the reactivation of development so that a missing body part grows back. This phenomenon has been studied particularly in salamanders, where the adults can reconstruct a whole limb after it has been amputated. Researchers hope to one day be able to induce regeneration in humans. There is little spontaneous regeneration in adult humans, although the liver is a notable exception. Like for salamanders, the regeneration of the liver involves dedifferentiation of some cells to a more embryonal state. Regeneration can also be studied in planaria. Cutting planaria causes an accumulation of epidermal cells at the site of the cut; a regeneration blastema is then formed from cells lacking differentiation, and differentiation occurs to replace absent parts within one week of the initial cut.
Developmental systems biology
Computer simulation of multicellular development is a research methodology to understand the function of the very complex processes involved in the development of organisms. This includes simulation of cell signaling, multicell interactions and regulatory genomic networks in development of multicellular structures and processes (see French flag model or Category:Developmental biology journals for literature). Minimal genomes for minimal multicellular organisms may pave the way to understand such complex processes in vivo.
- Altricial and Precocial
- Body plan
- Cell signaling
- Cell signaling networks
- Developmental noise
- Evolutionary developmental biology
- Fish development
- Gene regulatory network
- Homologous chromosome
- Plant evolutionary developmental biology
- Promoter (biology)
- Signal transduction
- Transcription factor
- Gilbert, S. F. (2013). Developmental Biology. Sunderland, Mass.: Sinauer Associates Inc.
- Slack, J. M. W. (2013). Essential Developmental Biology. Oxford: Wiley-Blackwell.
- Wolpert, L. and Tickle, C. (2011). Principles of Development. Oxford and New York: Oxford University Press.
- Nieuwkoop, P.D. and Faber, J. (1967) Normal table of Xenopus laevis (Daudin). North-Holland, Amsterdam.
- Harland, R.M. and Grainger, R.M. (2011) Xenopus research: metamorphosed by genetics and genomics. Trends in Genetics 27, 507-515.
- Lawson, N. D. and Wolfe, S. A. (2011) Forward and Reverse Genetic Approaches for the Analysis of Vertebrate Development in the Zebrafish. Developmental Cell 21, 48-64.
- Hassan Rashidi, V.S. (2009) The chick embryo: hatching a model for contemporary biomedical research. BioEssays 31, 459-465.
- Behringer, R., Gertsenstein, M, Vintersten, K. and Nagy, M. (2014) Manipulating the Mouse Embryo. A Laboratory Manual, Fourth Edition. Cold Spring Harbor, NY: Cold Spring Harbor Laboratory Press.
- St Johnston, D. (2002) The art and design of genetic screens: Drosophila melanogaster. Nat Rev Genet 3, 176-188.
- Riddle, D.L., Blumenthal, T., Meyer, B.J. and Priess, J.R. (1997) C.elegans II. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, NY.
- Ettensohn, C.A. and Sweet, H.C. (2000) Patterning the early sea urchin embryo. Curr. Top. Dev. Biol. 50, 1-44.
- Lemaire, P. (2011) Evolutionary crossroads in developmental biology: the tunicates. Development 138, 2143-2152.
- Nacu, E. and Tanaka, E.M. (2011) Limb Regeneration: A New Development? Annual Review of Cell and Developmental Biology 27, 409-440.
- Reddien, P.W. and Alvarado, A.S. (2004) Fundamentals of planarian regeneration. Annual Review of Cell and Developmental Biology 20, 725-757.
- Weigel, D. and Glazebrook, J. (2002) Arabidopsis. A Laboratory Manual. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, NY.
- Wolpert L, Beddington R, Jessell T, Lawrence P, Meyerowitz E, Smith J (2002). Principles of development (2nd ed.). Oxford university press. pp. 293–295. ISBN 0-19-879291-3.
- Ben-Tabou de-Leon S; Davidson EH (2007). "Gene regulation: gene control network in development". Annu Rev Biophys Biomol Struct 36: 191–212. doi:10.1146/annurev.biophys.35.040405.102002. PMID 17291181.
- Wolpert L, Beddington R, Jessell T, Lawrence P, Meyerowitz E, Smith J (2002). Principles of development (2nd ed.). Oxford university press. pp. 304–307. ISBN 0-19-879291-3.
- Wolpert L, Beddington R, Jessell T, Lawrence P, Meyerowitz E, Smith J (2002). Principles of development (2nd ed.). Oxford university press. pp. 41–50, 493. ISBN 0-19-879291-3.
- Christ B; Schmidt C; Huang R; Wilting J; Brand-Saberi B (January 1998). "Segmentation of the vertebrate body". Anat. Embryol. 197 (1): 1–8. doi:10.1007/s004290050116. PMID 9462855.
- Marshall H; Morrison A; Studer M; Pöpperl H; Krumlauf R (July 1996). "Retinoids and Hox genes". FASEB J. 10 (9): 969–78. PMID 8801179.
- Holtzman NA; Khoury MJ (1986). "Monitoring for congenital malformations". Annu Rev Public Health 7: 237–66. doi:10.1146/annurev.pu.07.050186.001321. PMID 3521645.
- Wolf U (1997). "Identical mutations and phenotypic variation". Hum Genet 100 (3–4): 305–21. doi:10.1007/s004390050509. PMID 9272148.
- Fujimoto K; Ishihara S; Kaneko K (2008). Hogeweg, Paulien, ed. "Network Evolution of Body Plans". PLoS ONE 3 (7): e2772. doi:10.1371/journal.pone.0002772. PMC 2464711. PMID 18648662.
- Wolpert L, Beddington R, Jessell T, Lawrence P, Meyerowitz E, Smith J (2002). Principles of development (2nd ed.). Oxford university press. pp. 467–482. ISBN 0-19-879291-3.
- Chamberlain G; Fox J; Ashton B; Middleton J (November 2007). "Concise review: mesenchymal stem cells: their phenotype, differentiation capacity, immunological features, and potential for homing". Stem Cells 25 (11): 2739–49. doi:10.1634/stemcells.2007-0197. PMID 17656645.
- Gilbert SF (2003). Developmental biology (7th ed.). Sinauer. pp. 575–585. ISBN 0-87893-258-5.
- Gilbert SF (2003). Developmental biology (7th ed.). Sinauer. pp. 592–601. ISBN 0-87893-258-5.
- Stocum DL (December 2002). "Development. A tail of transdifferentiation". Science 298 (5600): 1901–3. doi:10.1126/science.1079853. PMID 12471238.
- Newmark, Alvarado (2001). "Regeneration in Planaria". Wiley eLS. Nature Publishing Group. Retrieved 29 January 2014.
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