Developmental biology is the study of the process by which animals and plants grow and develop. Developmental biology also encompasses the biology of regeneration, asexual reproduction, metamorphosis, and the growth and differentiation of stem cells in the adult organism.
In the late 20th century, the discipline largely transformed into evolutionary developmental biology.
The main processes involved in the embryonic development of animals are: regional specification, morphogenesis, cell differentiation, growth, and the overall control of timing explored in evolutionary developmental biology:
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 are different from those that control animal development.
Cell differentiation is the process whereby different functional cell types arise in development. For example, neurons, muscle fibers and hepatocytes (liver cells) are well known types of differentiated cell. Differentiated cells usually produce large amounts of a few proteins that are required for their specific function and this gives them the characteristic appearance that enables them to be recognized under the light microscope. The genes encoding these proteins are highly active. Typically their chromatin structure is very open, allowing access for the transcription enzymes, and specific transcription factors bind to regulatory sequences in the DNA in order to activate gene expression. For example, NeuroD is a key transcription factor for neuronal differentiation, myogenin for muscle differentiation, and HNF4 for hepatocyte differentiation.
Cell differentiation is usually the final stage of development, preceded by several states of commitment which are not visibly differentiated. A single tissue, formed from a single type of progenitor cell or stem cell, often consists of several differentiated cell types. Control of their formation involves a process of lateral inhibition, based on the properties of the Notch signaling pathway. For example, in the neural plate of the embryo this system operates to generate a population of neuronal precursor cells in which NeuroD is highly expressed.
Regeneration indicates the ability to regrow a missing part. This is very prevalent amongst plants, which show continuous growth, and also among colonial animals such as hydroids and ascidians. But most interest by developmental biologists has been shown in the regeneration of parts in free living animals. In particular four models have been the subject of much investigation. Two of these have the ability to regenerate whole bodies: Hydra, which can regenerate any part of the polyp from a small fragment, and planarian worms, which can usually regenerate both heads and tails. Both of these examples have continuous cell turnover fed by stem cells and, at least in planaria, at least some of the stem cells have been shown to be pluripotent. The other two models show only distal regeneration of appendages. These are the insect appendages, usually the legs of hemimetabolous insects such as the cricket, and the limbs of urodele amphibians. Considerable information is now available about amphibian limb regeneration and it is known that each cell type regenerates itself, except for connective tissues where there is considerable interconversion between cartilage, dermis and tendons. In terms of the pattern of structures, this is controlled by a re-activation of signals active in the embryo. There is still debate about the old question of whether regeneration is a "pristine" or an "adaptive" property. If the former is the case, with improved knowledge, we might expect to be able to improve regenerative ability in humans. If the latter, then each instance of regeneration is presumed to have arisen by natural selection in circumstances particular to the species, so no general rules would be expected.
The sperm and egg fuse in the process of fertilization to form a fertilized egg, or zygote. This undergoes a period of divisions to form a ball or sheet of similar cells called a blastula or blastoderm. These cell divisions are usually rapid with no growth so the daughter cells are half the size of the mother cell and the whole embryo stays about the same size. They are called cleavage divisions. Morphogenetic movements convert the cell mass into a three layered structure consisting of multicellular sheets called ectoderm, mesoderm and endoderm, which are known as germ layers. This is the process of gastrulation. During cleavage and gastrulation the first regional specification events occur. In addition to the formation of the three germ layers themselves, these often generate extraembryonic structures, such as the mammalian placenta, needed for support and nutrition of the embryo, and also establish differences of commitment along the anteroposterior axis (head, trunk and tail).
Regional specification is initiated by the presence of cytoplasmic determinants in one part of the zygote. The cells that contain the determinant become a signaling center and emit an inducing factor. Because the inducing factor is produced in one place, diffuses away, and decays, it forms a concentration gradient, high near the source cells and low further away. The remaining cells of the embryo, which do not contain the determinant, are competent to respond to different concentrations by upregulating specific developmental control genes. This results in a series of zones becoming set up, arranged at progressively greater distance from the signaling center. In each zone a different combination of developmental control genes is upregulated. These genes encode transcription factors which upregulate new combinations of gene activity in each region. Among other functions, these transcription factors control expression of genes conferring specific adhesive and motility properties on the cells in which they are active. Because of these different morphogenetic properties, the cells of each germ layer move to form sheets such that the ectoderm ends up on the outside, mesoderm in the middle, and endoderm on the inside. Morphogenetic movements not only change the shape and structure of the embryo, but by bringing cell sheets into new spatial relationships they also make possible new phases of signaling and response between them.
Growth in embryos is mostly autonomous. For each territory of cells the growth rate is controlled by the combination of genes that are active. Free-living embryos do not grow in mass as they have no external food supply. But embryos fed by a placenta or extraembryonic yolk supply can grow very fast, and changes to relative growth rate between parts in these organisms help to produce the final overall anatomy.
The whole process needs to be coordinated in time and how this is controlled is not understood. There may be a master clock able to communicate with all parts of the embryo that controls the course of events, or timing may depend simply on local causal sequences of events.
Developmental processes are very evident during the process of metamorphosis. This occurs in various types of animal. Well-known are the examples of the frog, which usually hatches as a tadpole and metamorphoses to an adult frog, and certain insects which hatch as a larva and then become remodeled to the adult form during a pupal stage.
All the developmental processes listed above occur during metamorphosis. Examples that have been especially well studied include tail loss and other changes in the tadpole of the frog Xenopus, and the biology of the imaginal discs, which generate the adult body parts of the fly Drosophila melanogaster.
Plants constantly produce new tissues and structures throughout their life from meristems located at the tips of organs, or between mature tissues. Thus, a living plant always has embryonic tissues. By contrast, an animal embryo will very early produce all of the body parts that it will ever have in its life. When the animal is born (or hatches from its egg), it has all its body parts and from that point will only grow larger and more mature.
The properties of organization seen in a plant are emergent properties which are more than the sum of the individual parts. "The assembly of these tissues and functions into an integrated multicellular organism yields not only the characteristics of the separate parts and processes but also quite a new set of characteristics which would not have been predictable on the basis of examination of the separate parts."
A vascular plant begins from a single celled zygote, formed by fertilisation of an egg cell by a sperm cell. From that point, it begins to divide to form a plant embryo through the process of embryogenesis. As this happens, the resulting cells will organize so that one end becomes the first root, while the other end forms the tip of the shoot. In seed plants, the embryo will develop one or more "seed leaves" (cotyledons). By the end of embryogenesis, the young plant will have all the parts necessary to begin in its life.
Once the embryo germinates from its seed or parent plant, it begins to produce additional organs (leaves, stems, and roots) through the process of organogenesis. New roots grow from root meristems located at the tip of the root, and new stems and leaves grow from shoot meristems located at the tip of the shoot. Branching occurs when small clumps of cells left behind by the meristem, and which have not yet undergone cellular differentiation to form a specialized tissue, begin to grow as the tip of a new root or shoot. Growth from any such meristem at the tip of a root or shoot is termed primary growth and results in the lengthening of that root or shoot. Secondary growth results in widening of a root or shoot from divisions of cells in a cambium.
In addition to growth by cell division, a plant may grow through cell elongation. This occurs when individual cells or groups of cells grow longer. Not all plant cells will grow to the same length. When cells on one side of a stem grow longer and faster than cells on the other side, the stem will bend to the side of the slower growing cells as a result. This directional growth can occur via a plant's response to a particular stimulus, such as light (phototropism), gravity (gravitropism), water, (hydrotropism), and physical contact (thigmotropism).
Plant growth and development are mediated by specific plant hormones and plant growth regulators (PGRs) (Ross et al. 1983). Endogenous hormone levels are influenced by plant age, cold hardiness, dormancy, and other metabolic conditions; photoperiod, drought, temperature, and other external environmental conditions; and exogenous sources of PGRs, e.g., externally applied and of rhizospheric origin.
Plants exhibit natural variation in their form and structure. While all organisms vary from individual to individual, plants exhibit an additional type of variation. Within a single individual, parts are repeated which may differ in form and structure from other similar parts. This variation is most easily seen in the leaves of a plant, though other organs such as stems and flowers may show similar variation. There are three primary causes of this variation: positional effects, environmental effects, and juvenility.
Transcription factors and transcriptional regulatory networks play key roles in plant morphogenesis and their evolution. During plant landing, many novel transcription factor families emerged and are preferentially wired into the networks of multicellular development, reproduction, and organ development, contributing to more complex morphogenesis of land plants.
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.
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. Organoids have also been demonstrated as an efficient model for development. Plant development has focused on the thale cress Arabidopsis thaliana as a model organism.
Developmental Biology is a peer-reviewed scientific journal. It was established in 1959 and is the official journal of the Society for Developmental Biology. It publishes research on the mechanisms of development, differentiation, and growth in animals and plants at the molecular, cellular, and genetic levels. The journal is published twice a month by Elsevier.Diencephalon
The diencephalon is a division of the forebrain (embryonic prosencephalon), and is situated between the telencephalon and the midbrain (embryonic mesencephalon). It consists of structures that are on either side of the third ventricle, including the thalamus, the hypothalamus, the epithalamus and the subthalamus.
The diencephalon is one of the main vesicles of the brain formed during embryogenesis. During the third week of development a neural tube is created from the ectoderm, one of the three primary germ layers. The tube forms three main vesicles during the third week of development: the prosencephalon, the mesencephalon and the rhombencephalon. The prosencephlon gradually divides into the telencephalon and the diencephalon.Embryo
An embryo is an early stage of development of a multicellular diploid eukaryotic organism. In general, in organisms that reproduce sexually, an embryo develops from a zygote, the single cell resulting from the fertilization of the female egg cell by the male sperm cell. The zygote possesses half the DNA from each of its two parents. In plants, animals, and some protists, the zygote will begin to divide by mitosis to produce a multicellular organism. The result of this process is an embryo.
In human pregnancy, a developing fetus is considered as an embryo until the ninth week, fertilization age, or eleventh-week gestational age. After this time the embryo is referred to as a fetus.Endoderm
Endoderm is one of the three primary germ layers in the very early embryo. The other two layers are the ectoderm (outside layer) and mesoderm (middle layer), with the endoderm being the innermost layer. Cells migrating inward along the archenteron form the inner layer of the gastrula, which develops into the endoderm.The endoderm consists at first of flattened cells, which subsequently become columnar. It forms the epithelial lining of multiple systems.In plant biology, endoderm corresponds to the innermost part of the cortex (bark) in young shoots and young roots often consisting of a single cell layer. As the plant becomes older, more endoderm will lignify.Evolutionary developmental biology
Evolutionary developmental biology (informally, evo-devo) is a field of biological research that compares the developmental processes of different organisms to infer the ancestral relationships between them and how developmental processes evolved.
The field grew from 19th-century beginnings, where embryology faced a mystery: zoologists did not know how embryonic development was controlled at the molecular level. Charles Darwin noted that having similar embryos implied common ancestry, but little progress was made until the 1970s. Then, recombinant DNA technology at last brought embryology together with molecular genetics. A key early discovery was of homeotic genes that regulate development in a wide range of eukaryotes.
The field is characterised by some key concepts, which took evolutionary biologists by surprise. One is deep homology, the finding that dissimilar organs such as the eyes of insects, vertebrates and cephalopod molluscs, long thought to have evolved separately, are controlled by similar genes such as pax-6, from the evo-devo gene toolkit. These genes are ancient, being highly conserved among phyla; they generate the patterns in time and space which shape the embryo, and ultimately form the body plan of the organism. Another is that species do not differ much in their structural genes, such as those coding for enzymes; what does differ is the way that gene expression is regulated by the toolkit genes. These genes are reused, unchanged, many times in different parts of the embryo and at different stages of development, forming a complex cascade of control, switching other regulatory genes as well as structural genes on and off in a precise pattern. This multiple pleiotropic reuse explains why these genes are highly conserved, as any change would have many adverse consequences which natural selection would oppose.
New morphological features and ultimately new species are produced by variations in the toolkit, either when genes are expressed in a new pattern, or when toolkit genes acquire additional functions. Another possibility is the Neo-Lamarckian theory that epigenetic changes are later consolidated at gene level, something that may have been important early in the history of multicellular life.Germ layer
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 (animals more complex than the sponge) produce two or three primary germ layers. Some animals, like cnidarians, produce two germ layers (the ectoderm and endoderm) making them diploblastic. Other animals such as chordates 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.Imago
In biology, the imago is the last stage an insect attains during its metamorphosis, its process of growth and development; it also is called the imaginal stage, the stage in which the insect attains maturity. It follows the final ecdysis of the immature instars.In a member of the Ametabola or Hemimetabola, in which metamorphosis is "incomplete", the final ecdysis follows the last immature or nymphal stage.
In members of the Holometabola, in which there is a pupal stage, the final ecdysis follows emergence from the pupa, after which the metamorphosis is complete, although there is a prolonged period of maturation in some species.The imago is the only stage during which the insect is sexually mature and, if it is a winged species, has functional wings. The imago often is referred to as the adult stage.Members of the order Ephemeroptera (mayflies) do not have a pupal stage, but they briefly pass through an extra winged stage called the subimago. Insects at this stage have functional wings but are not yet sexually mature.The Latin plural of imago is imagines, and this is the term generally used by entomologists –
however, imagoes is also acceptable.Juvenile (organism)
A juvenile is an individual organism that has not yet reached its adult form, sexual maturity or size. Juveniles sometimes look very different from the adult form, particularly in colour. In many organisms the juvenile has a different name from the adult (see also List of animal names).
Some organisms reach sexual maturity in a short metamorphosis, such as eclosion in many insects. For others, the transition from juvenile to fully mature is a more prolonged process—puberty, for example. In such cases, juveniles during this transformation are sometimes called subadults.
Many invertebrates, on reaching the adult stage, are fully mature and their development and growth stops. Their juveniles are larvae or nymphs.
In vertebrates and some invertebrates (e.g. spiders), larval forms (e.g. tadpoles) are usually considered a development stage of their own, and "juvenile" refers to a post-larval stage that is not fully grown and not sexually mature. In amniotes and most plants, the embryo represents the larval stage. Here, a "juvenile" is an individual in the time between hatching/birth/germination and reaching maturity.Maggot
A maggot is the larva of a fly (order Diptera); it is applied in particular to the larvae of Brachycera flies, such as houseflies, cheese flies, and blowflies, rather than larvae of the Nematocera, such as mosquitoes and Crane flies. A 2012 study estimated the population of maggots in North America alone to be in excess of 3×105 trillion.Max Planck Institute for Developmental Biology
The Max Planck Institute for Developmental Biology is located in Tübingen, Germany; it was founded in 1954 as an offshoot of the Tübingen-based Max Planck Institute for Biology. The main topics of scientific research conducted by the Max Planck Institute for Developmental Biology focus on the molecular mechanisms underlying spatial information within the embryo, communication between cells in the induction process, as well as the formation and differentiation of tissues and organs.Morphogenesis
Morphogenesis (from the Greek morphê shape and genesis creation, literally, "beginning of the shape") is the biological process that causes an organism to develop its shape. It is one of three fundamental aspects of developmental biology along with the control of cell growth and cellular differentiation, unified in evolutionary developmental biology (evo-devo).
The process controls the organized spatial distribution of cells during the embryonic development of an organism. Morphogenesis can take place also in a mature organism, in cell culture or inside tumor cell masses. Morphogenesis also describes the development of unicellular life forms that do not have an embryonic stage in their life cycle, or describes the evolution of a body structure within a taxonomic group.
Morphogenetic responses may be induced in organisms by hormones, by environmental chemicals ranging from substances produced by other organisms to toxic chemicals or radionuclides released as pollutants, and other plants, or by mechanical stresses induced by spatial patterning of the cells.Morphogenetic field
This article is about the concept in developmental biology. For Rupert Sheldrake's concept of morphic fields, see the bibliography in his article.
In the developmental biology of the early twentieth century, a morphogenetic field is a group of cells able to respond to discrete, localized biochemical signals leading to the development of specific morphological structures or organs. The spatial and temporal extents of the embryonic field are dynamic, and within the field is a collection of interacting cells out of which a particular organ is formed. As a group, the cells within a given morphogenetic field are constrained: thus, cells in a limb field will become a limb tissue, those in a cardiac field will become heart tissue. However, specific cellular programming of individual cells in a field is flexible: an individual cell in a cardiac field can be redirected via cell-to-cell signaling to replace specific damaged or missing cells. Imaginal discs in insect larvae are examples of morphogenetic fields.Neurogenesis
Neurogenesis is the process by which nervous system cells, known as neurons, are produced by neural stem cells (NSC)s, and it occurs in all species of animals except the porifera (sponges) and placozoans. Types of NSCs include neuroepithelial cells (NECs), radial glial cells (RGCs), basal progenitors (BPs), intermediate neuronal precursors (INP)s, subventricular zone astrocytes, and subgranular zone radial astrocytes, among others. Neurogenesis is most active during embryonic development, and is responsible for producing all the various types of neurons of the organism, but continues throughout adult life in a variety of organisms. Once born, neurons do not divide (see mitosis), and many will live the lifetime of the animal.Nidicolous
A nidicolous animal ( ny-DIK-ə-ləs; from Latin nidus "nest" and -colus "inhabiting") is an animal that stays at its birthplace for a long time because it depends on the parents for food, protection, and the learning of survival skills. They are the opposite of nidifugous species, which leave their parents more quickly and survive independently.
Two other terms are also used by scientists for related developmental phenomena: altricial (relatively undeveloped at birth or hatching; helpless, blind, without feathers or hair, and unable to fend for themselves) and precocial (relatively developed at birth or hatching; able to fend for themselves). Although there is much overlap between altricial and nidicolous species, the terms are not identical. All altricial animals are nidicolous by necessity, but an animal may be nidicolous, such as staying at the nest, even if it is precocial and fully capable of leaving if needed. Examples of precocious but nidicolous species include many gulls and terns.Examples of nidicolous species are most mammals and many species of birds. The majority of nidicolous animals are altricial. During the life span, the brain of a nidicolous animal expands 8–10 times its initial size; in nidifugous animals, from 1.5 to 2.5 times.Ontogeny
Ontogeny (also ontogenesis or morphogenesis) is the origination and development of an organism, usually from the time of fertilization of the egg to the organism's mature form—although the term can be used to refer to the study of the entirety of an organism's lifespan.
Ontogeny is the developmental history of an organism within its own lifetime, as distinct from phylogeny, which refers to the evolutionary history of a species. In practice, writers on evolution often speak of species as "developing" traits or characteristics. This can be misleading. While developmental (i.e., ontogenetic) processes can influence subsequent evolutionary (e.g., phylogenetic) processes (see evolutionary developmental biology), individual organisms develop (ontogeny), while species evolve (phylogeny).
Ontogeny, embryology and developmental biology are closely related studies and the terms are sometimes used interchangeably. The term ontogeny has also been used in cell biology to describe the development of various cell types within an organism.Ontogeny is a useful field of study in many disciplines, including developmental biology, developmental psychology, developmental cognitive neuroscience, and developmental psychobiology.
Ontogeny is also a concept used in anthropology as "the process through which each of us embodies the history of our own making".Pattern formation
The science of pattern formation deals with the visible, (statistically) orderly outcomes of self-organization and the common principles behind similar patterns in nature.
In developmental biology, pattern formation refers to the generation of complex organizations of cell fates in space and time. Pattern formation is controlled by genes. The role of genes in pattern formation is an aspect of morphogenesis, the creation of diverse anatomies from similar genes, now being explored in the science of evolutionary developmental biology or evo-devo. The mechanisms involved are well seen in the anterior-posterior patterning of embryos from the model organism Drosophila melanogaster (a fruit fly), one of the first organisms to have its morphogenesis studied, and in the eyespots of butterflies, whose development is a variant of the standard (fruit fly) mechanism.Somatic (biology)
The term somatic is often used in biology to refer to the cells of the body in contrast to the germ line cells which usually give rise to the gametes (ovum or sperm). These somatic cells are diploid containing two copies of each chromosome, whereas the germ cells are haploid as they only contain one copy of each chromosome. Although under normal circumstances all somatic cells in an organism contain identical DNA, they develop a variety of tissue-specific characteristics. This process is called differentiation, through epigenetic and regulatory alterations. The grouping of like cells and tissues creates the foundation for organs.
Somatic mutations are changes to the genetics of a multi-cellular organism which are not passed on to its offspring through the germ-line. Many cancers are somatic mutations.
Somatic is also defined as relating to the wall of the body cavity, particularly as distinguished from the head, limbs or viscera.
It is also used in the term somatic nervous system which is the portion of the vertebrate nervous system which regulates voluntary movements of the body.Zygote
A zygote (from Greek ζυγωτός zygōtos "joined" or "yoked", from ζυγοῦν zygoun "to join" or "to yoke") 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 necessary to form a new individual. In multicellular organisms, the zygote is the earliest developmental stage. In single-celled organisms, the zygote can divide asexually by mitosis to produce identical offspring.
Oscar Hertwig and Richard Hertwig made some of the first discoveries on animal zygote formation.