The Info List - Developmental Biology

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Developmental biology
Developmental biology
is the study of the process by which animals and plants grow and develop. Developmental biology
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 has largely transformed into evolutionary developmental biology.


1 Perspectives 2 Developmental processes

2.1 Cell differentiation 2.2 Regeneration

3 Embryonic development of animals

3.1 Metamorphosis

4 Plant development

4.1 Growth 4.2 Morphological variation 4.3 Evolution of plant morphology

5 Developmental model organisms 6 See also 7 References 8 Further reading 9 External links

Perspectives[edit] 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. 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
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 are different from those that control animal development. Developmental processes[edit] Cell differentiation[edit]

The Notch-delta system in neurogenesis.(Slack Essential Dev Biol Fig 14.12a)

Cell differentiation
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.[1][2] For example, NeuroD is a key transcription factor for neuronal differentiation, myogenin for muscle differentiation, and HNF4 for hepatocyte differentiation. Cell 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,[3] based on the properties of the Notch signaling pathway.[4] 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[edit] Regeneration indicates the ability to regrow a missing part.[5] 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,[6] and planarian worms, which can usually regenerate both heads and tails.[7] 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.[8] 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,[9] and the limbs of urodele amphibians.[10] 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.[11] 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. Embryonic development of animals[edit] Main article: Embryogenesis

Generalized scheme of embryonic development. Slack "Essential Developmental Biology" Fig.2.8

The initial stages of human embryogenesis.

The sperm and egg fuse in the process of fertilization to form a fertilized egg, or zygote.[12] 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,[13] and also establish differences of commitment along the anteroposterior axis (head, trunk and tail).[14] 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.[15][16] 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.[17] 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.[18][19] 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.[20] 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.[21] Metamorphosis[edit] 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,[22][23] and the biology of the imaginal discs, which generate the adult body parts of the fly Drosophila melanogaster.[24][25] Plant development[edit] Further information: Plant development Plant development
Plant development
is the process by which structures originate and mature as a plant grows. It is studied in plant anatomy and plant physiology as well as plant morphology. Plants constantly produce new tissues and structures throughout their life from meristems[26] 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."[27] Growth[edit] 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.[28] 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.[29] 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).[30] 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. Morphological variation[edit] 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. Evolution of plant morphology[edit] 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.[31] Developmental model organisms[edit] 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
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
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. Plants:

Thale cress
Thale cress
(Arabidopsis thaliana)


Frog: Xenopus
(X.laevis and tropicalis).[32][33] Good embryo supply. Especially suitable for microsurgery. Zebrafish: Danio rerio.[34] Good embryo supply. Well developed genetics. Chicken: Gallus gallus.[35] Early stages similar to mammal, but microsurgery easier. Low cost. Mouse: Mus musculus.[36] A mammal with well developed genetics.


Fruit fly: Drosophila melanogaster.[37] Good embryo supply. Well developed genetics. Nematode: Caenorhabditis elegans.[38] Good embryo supply. Well developed genetics. Low cost.

Also popular for some purposes have been sea urchins[39] and ascidians.[40] For studies of regeneration urodele amphibians such as the axolotl Ambystoma mexicanum are used,[41] and also planarian worms such as Schmidtea mediterranea.[42] Organoids have also been demonstrated as an efficient model for development.[43] Plant development has focused on the thale cress Arabidopsis thaliana
Arabidopsis thaliana
as a model organism.[44] See also[edit]

Blastocyst Body plan Cell signaling Cell signaling
Cell signaling
networks Embryology Enhancer Fish development Gene regulatory network Ontogeny Plant evolutionary developmental biology Promoter (biology) Signal transduction Teratology


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Further reading[edit]

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.

External links[edit]

Wikibooks has more on the topic of: Developmental biology

Society for Developmental Biology Collaborative resources Developmental Biology
- 10th edition Essential Developmental Biology
3rd edition

v t e

Developmental biology

Ageing Compartment Embryogenesis

Human Drosophila Fish polarity

Embryology Metamerism Metamorphosis

Indirect development: holometabolism, hemimetabolism Direct development: ametabolism

Model organisms Ontogeny Puberty Regeneration



Evolutionary developmental biology

v t e

Human embryogenesis
Human embryogenesis
in the first three weeks

Week 1

Fertilization Oocyte activation Zygote Cleavage Blastomere Morula Blastocoele Blastocyst Blastula Inner cell mass Trophoblast

Week 2 (Bilaminar)

Hypoblast Epiblast

Week 3 (Trilaminar)

Germ layers

Archenteron/Primitive streak

Primitive pit Primitive knot/Blastopore Primitive groove



Regional specification Embryonic disc


Surface ectoderm Neuroectoderm Somatopleuric mesenchyme Neurulation Neural crest


Splanchnopleuric mesenchyme


Axial mesoderm Paraxial

Somite Somitomere

Intermediate Lateral plate

Intraembryonic coelom Splanchnopleuric mesenchyme Somatopleuric mesenchyme

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Human biological and psychological development

Before birth

Embryo Fetus

After birth

Infant Toddler Early childhood Childhood


Preadolescence Adolescence Adulthood Middle age Old age


Minor Age of majority

Events and phases

Gestational age Prenatal development Birth Child


Cognitive development of infants Human development Adult
development Puberty Ageing Senescence Death

Developmental psychology

Antenatal Positive youth development Young adult Positive adult development Maturity

Theorists and theories

Freud (1856–1939) (Psychosexual development) Piaget (1896–1980) (Theory of cognitive development) Vygotsky (1896–1934) (Cultural-historical psychology) Erikson (1902–1994) (Psychosocial development) Bowlby (1907–1990) (Attachment theory) Bronfenbrenner (1917–2005) (Ecological systems theory) Kohlberg (1927–1987) (Stages of moral development) Commons (b. 1939), Fischer (b. 1943), Kegan (b. 1946), Demetriou (b. 1950), and others (Neo-Piagetian theories of cognitive development) Evolutionary developmental psychology

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Branches of life science and biology

Anatomy Astrobiology Biochemistry Biogeography Biohistory Biomechanics Biophysics Bioinformatics Biostatistics Botany Cell biology Cellular microbiology Chemical biology Chronobiology Computational biology Conservation biology Cytogenetics Developmental biology Ecology Embryology Epidemiology Epigenetics Evolutionary biology Freshwater biology Geobiology Genetics Genomics Histology Human biology Immunology Marine biology Mathematical biology Microbiology Molecular biology Mycology Neontology Neuroscience Nutrition Origin of life Paleontology Parasitology Pathology Pharmacology Phylogenetics Physiology Quantum biology Sociobiology Structural biology Systematics Systems biology Taxonomy Teratology Toxicology Virology Virophysics Zoology

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The development of phenotype

Key concepts

Genotype–phenotype distinction Norms of reaction Gene–environment interaction Gene–environment correlation Operon Heritability Quantitative genetics Heterochrony



Genetic architecture

Canalisation Genetic assimilation Dominance Epistasis Fitness landscape/evolutionary landscape Pleiotropy Plasticity Polygenic inheritance Transgressive segregation Sequence space

Non-genetic influences

Epigenetics Maternal effect Dual inheritance theory Polyphenism

Developmental architecture

Developmental biology Morphogenesis

Eyespot Pattern formation

Segmentation Modularity

Evolution of genetic systems

Evolvability Mutational robustness Neutral networks Evolution of sexual reproduction

Control of development


Regulation of gene expression Gene regulatory network Developmental-genetic toolkit Evolutionary developmental biology Homeobox Hedgehog signaling pathway Notch signaling pathway


Homeotic gene Hox gene Pax genes

eyeless gene

Distal-less Engrailed cis-regulatory element Ligand Morphogen Cell surface receptor Transcription factor

Influential figures

C. H. Waddington Richard Lewontin François Jacob
François Jacob
+ Jacques Monod

Lac operon

Eric F. Wieschaus Christiane Nüsslein-Volhard William McGinnis Mike Levine Sean B. Carroll

Endless Forms Most Beautiful


Nature versus nurture Morphogenetic field

Index of evolutionary biology articles

Authority control