Evo-devo gene toolkit

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Expression of all 8 Hox genes in the fruit fly Drosophila melanogaster Hoxgenesoffruitfly.svg
Expression of all 8 Hox genes in the fruit fly Drosophila melanogaster

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.

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Toolkit

Toolkit genes are highly conserved among phyla, meaning that they are ancient, dating back to the last common ancestor of bilaterian animals. For example, that ancestor had at least 7 Pax genes for transcription factors. [1]

Differences in deployment of toolkit genes affect the body plan and the number, identity, and pattern of body parts. The majority of toolkit genes are components of signaling pathways and encode for the production of transcription factors, cell adhesion proteins, cell surface receptor proteins (and signalling ligands that bind to them), and secreted morphogens; all of these participate in defining the fate of undifferentiated cells, generating spatial and temporal patterns that, in turn, form the body plan of the organism. Among the most important of the toolkit genes are those of the Hox gene cluster, or complex. Hox genes, transcription factors containing the more broadly distributed homeobox protein-binding DNA motif, function in patterning the body axis. Thus, by combinatorially specifying the identity of particular body regions, Hox genes determine where limbs and other body segments will grow in a developing embryo or larva. A paradigmatic toolkit gene is Pax6/eyeless, which controls eye formation in all animals. It has been found to produce eyes in mice and Drosophila , even if mouse Pax6/eyeless was expressed in Drosophila. [2]

This means that a big part of the morphological evolution undergone by organisms is a product of variation in the genetic toolkit, either by the genes changing their expression pattern or acquiring new functions. A good example of the first is the enlargement of the beak in Darwin's large ground-finch (Geospiza magnirostris), in which the gene BMP is responsible for the larger beak of this bird, relative to the other finches. [3]

The loss of legs in snakes and other squamates is another good example of genes changing their expression pattern. In this case the gene Distal-less is very under-expressed, or not expressed at all, in the regions where limbs would form in other tetrapods. [4] In 1994, Sean B. Carroll's team made the "groundbreaking" discovery that this same gene determines the eyespot pattern in butterfly wings, showing that toolbox genes can change their function. [5] [6] [7]

Toolkit genes, as well as being highly conserved, also tend to evolve the same function convergently or in parallel. Classic examples of this are the already mentioned Distal-less gene, which is responsible for appendage formation in both tetrapods and insects, or, at a finer scale, the generation of wing patterns in the butterflies Heliconius erato and Heliconius melpomene . These butterflies are Müllerian mimics whose coloration pattern arose in different evolutionary events, but is controlled by the same genes. [8] This supports Marc Kirschner and John C. Gerhart's theory of Facilitated Variation, which states that morphological evolutionary novelty is generated by regulatory changes in various members of a large set of conserved mechanisms of development and physiology. [9]

See also

Related Research Articles

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Evolutionary developmental biology is a field of biological research that compares the developmental processes of different organisms to infer how developmental processes evolved.

<span class="mw-page-title-main">Homeobox</span> DNA pattern affecting anatomy development

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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.

<i>Antennapedia</i> Hox gene

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<span class="mw-page-title-main">Facilitated variation</span>

The theory of facilitated variation demonstrates how seemingly complex biological systems can arise through a limited number of regulatory genetic changes, through the differential re-use of pre-existing developmental components. The theory was presented in 2005 by Marc W. Kirschner and John C. Gerhart.

<span class="mw-page-title-main">Pax genes</span> Family of transcription factors

In evolutionary developmental biology, Paired box (Pax) genes are a family of genes coding for tissue specific transcription factors containing an N-terminal paired domain and usually a partial, or in the case of four family members, a complete homeodomain to the C-terminus. An octapeptide as well as a Pro-Ser-Thr-rich C terminus may also be present. Pax proteins are important in early animal development for the specification of specific tissues, as well as during epimorphic limb regeneration in animals capable of such.

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Abdominal pigmentation in Drosophila melanogaster is a morphologically simple but highly variable trait that often has adaptive significance. Pigmentation has extensively been studied in Drosophila melanogaster. It has been used as a model for understanding the development and evolution of morphological phenotypes.

References

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