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. [1] [2] 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. [3] 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. [4] 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. [4] Imaginal discs in insect larvae are examples of morphogenetic fields. [5]
The concept of the morphogenetic field, fundamental in the early twentieth century to the study of embryological development, was first introduced in 1910 by Alexander G. Gurwitsch. [6] Experimental support was provided by Ross Granville Harrison's experiments transplanting fragments of a newt embryo into different locations. [7]
Harrison was able to identify "fields" of cells producing organs such as limbs, tail and gills and to show that these fields could be fragmented or have undifferentiated cells added and a complete normal final structure would still result. It was thus considered that it was the "field" of cells, rather than individual cells, that were patterned for subsequent development of particular organs. The field concept was developed further by Harrison's friend Hans Spemann, and then by Paul Weiss and others. [3] The concept was similar to the meaning of the term entelechy of vitalists like Hans Adolf Eduard Driesch (1867–1941).
By the 1930s, however, the work of geneticists, especially Thomas Hunt Morgan, revealed the importance of chromosomes and genes for controlling development, and the rise of the new synthesis in evolutionary biology lessened the perceived importance of the field hypothesis. Morgan was a particularly harsh critic of fields since the gene and the field were perceived as competitors for recognition as the basic unit of ontogeny. [3] With the discovery and mapping of master control genes, such as the homeobox genes the pre-eminence of genes seemed assured. But in the late twentieth century the field concept was "rediscovered" as a useful part of developmental biology. It was found, for example, that different mutations could cause the same malformations, suggesting that the mutations were affecting a complex of structures as a unit, a unit that might correspond to the field of early 20th century embryology.
Scott Gilbert proposed that the morphogenetic field is a middle ground between genes and evolution. [3] That is, genes act upon fields, which then act upon the developing organism. [3] Jessica Bolker described morphogenetic fields not merely as incipient structures or organs, but as dynamic entities with their own localized development processes, which are central to the emerging field of Evolutionary developmental biology ("evo-devo"). [8] In 2005, Sean B. Carroll and colleagues mention morphogenetic fields only as a concept proposed by early embryologists to explain the finding that a forelimb bud could be transplanted and still give rise to a forelimb; they define "field" simply as "a discrete region" in an embryo. [9]
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.
Evolutionary developmental biology is a field of biological research that compares the developmental processes of different organisms to infer how developmental processes evolved.
In biology, homology is similarity due to shared ancestry between a pair of structures or genes in different taxa. A common example of homologous structures is the forelimbs of vertebrates, where the wings of bats and birds, the arms of primates, the front flippers of whales and the forelegs of four-legged vertebrates like dogs and crocodiles are all derived from the same ancestral tetrapod structure. Evolutionary biology explains homologous structures adapted to different purposes as the result of descent with modification from a common ancestor. The term was first applied to biology in a non-evolutionary context by the anatomist Richard Owen in 1843. Homology was later explained by Charles Darwin's theory of evolution in 1859, but had been observed before this, from Aristotle onwards, and it was explicitly analysed by Pierre Belon in 1555.
Evolutionary biology is the subfield of biology that studies the evolutionary processes that produced the diversity of life on Earth. Simply, it is also defined as the study of the history of life forms on Earth. Evolution is based on the theory that all species are related and they gradually change over time. In a population, the genetic variations affect the physical characteristics i.e. phenotypes of an organism. These changes in the phenotypes will be an advantage to some organisms, which will then be passed onto their offspring. Peppered Moth and Flightless birds are some examples of evolution in species over many generations. In the 1930s, the discipline of evolutionary biology emerged through what Julian Huxley called the modern synthesis of understanding, from previously unrelated fields of biological research, such as genetics and ecology, systematics, and paleontology. A person who studies Evolutionary biology is called an Evolutionary biologist. The importance of studying Evolutionary biology is mainly to understand the principles behind the origin and extinction of species.
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.
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.
In evolutionary developmental biology, homeosis is the transformation of one organ into another, arising from mutation in or misexpression of certain developmentally critical genes, specifically homeotic genes. In animals, these developmental genes specifically control the development of organs on their anteroposterior axis. In plants, however, the developmental genes affected by homeosis may control anything from the development of a stamen or petals to the development of chlorophyll. Homeosis may be caused by mutations in Hox genes, found in animals, or others such as the MADS-box family in plants. Homeosis is a characteristic that has helped insects become as successful and diverse as they are.
A morphogen is a substance whose non-uniform distribution governs the pattern of tissue development in the process of morphogenesis or pattern formation, one of the core processes of developmental biology, establishing positions of the various specialized cell types within a tissue. More specifically, a morphogen is a signaling molecule that acts directly on cells to produce specific cellular responses depending on its local concentration.
Hox genes, a subset of homeobox genes, are a group of related genes that specify regions of the body plan of an embryo along the head-tail axis of animals. Hox proteins encode and specify the characteristics of 'position', ensuring that the correct structures form in the correct places of the body. For example, Hox genes in insects specify which appendages form on a segment, and Hox genes in vertebrates specify the types and shape of vertebrae that will form. In segmented animals, Hox proteins thus confer segmental or positional identity, but do not form the actual segments themselves.
A body plan, Bauplan, or ground plan is a set of morphological features common to many members of a phylum of animals. The vertebrates are one body plan: invertebrates have many.
Limb development in vertebrates is an area of active research in both developmental and evolutionary biology, with much of the latter work focused on the transition from fin to limb.
In evolutionary developmental biology, the concept of deep homology is used to describe cases where growth and differentiation processes are governed by genetic mechanisms that are homologous and deeply conserved across a wide range of species.
Gerd B. Müller is an Austrian biologist who is professor at the University of Vienna where he heads the Department of Theoretical Biology in the Center for Organismal Systems Biology. His research interests focus on vertebrate limb development, evolutionary novelties, evo-devo theory, and the Extended Evolutionary Synthesis. He is also concerned with the development of 3D based imaging tools in developmental biology.
Edward Michael De Robertis is an American embryologist and Professor at University of California, Los Angeles, whose work has contributed to the discovery of conserved molecular mechanisms of embryonic inductions that cause tissue differentiations during animal development.
Endless Forms Most Beautiful: The New Science of Evo Devo and the Making of the Animal Kingdom is a 2005 book by the molecular biologist Sean B. Carroll. It presents a summary of the emerging field of evolutionary developmental biology and the role of toolkit genes. It has won numerous awards for science communication.
Rudolf Albert Raff was an American biologist, and James H. Rudy Professor of Biology at Indiana University. He was known for research in, and promotion of, evolutionary developmental biology. He was also director of the Indiana Molecular Biology Institute.
Hox genes play a massive role in some amphibians and reptiles in their ability to regenerate lost limbs, especially HoxA and HoxD genes.
The extended evolutionary synthesis consists of a set of theoretical concepts argued to be more comprehensive than the earlier modern synthesis of evolutionary biology that took place between 1918 and 1942. The extended evolutionary synthesis was called for in the 1950s by C. H. Waddington, argued for on the basis of punctuated equilibrium by Stephen Jay Gould and Niles Eldredge in the 1980s, and was reconceptualized in 2007 by Massimo Pigliucci and Gerd B. Müller. Notably, Dr. Müller concluded from this research that Natural Selection has no way of explaining speciation, saying: “selection has no innovative capacity...the generative and the ordering aspects of morphological evolution are thus absent from evolutionary theory.”
Scott Frederick Gilbert is an American evolutionary developmental biologist and historian of biology.
The evo-devo gene toolkit is the small subset of genes in an organism's genome whose products control the organism's embryonic development. Toolkit genes are central to the synthesis of molecular genetics, palaeontology, evolution and developmental biology in the science of evolutionary developmental biology (evo-devo). Many of them are ancient and highly conserved among animal phyla.