Limb developmentLimb 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. Limb formation begins in the morphogenetic limb field, as mesenchymal cells from the lateral plate mesoderm proliferate to the point that they cause the ectoderm above to bulge out, forming a limb bud. Fibroblast growth factor (FGF) induces the formation of an organizer at the end of the limb bud, called the apical ectodermal ridge (AER), which guides further development and controls cell death.
Limb budThe limb bud is a structure formed early in vertebrate limb development. As a result of interactions between the ectoderm and underlying mesoderm, formation occurs roughly around the fourth week of development. In the development of the human embryo the upper limb bud appears in the third week and the lower limb bud appears four days later. The limb bud consists of undifferentiated mesoderm cells that are sheathed in ectoderm.
Limb (anatomy)A limb is a jointed, muscled appendage of a tetrapod vertebrate animal used for weight-bearing and terrestrial locomotion. The distalmost portion of a limb is known as its extremity. The limbs' bony endoskeleton, known as the appendicular skeleton, is homologous among all tetrapods, who use their limbs for walking, running and jumping, swimming, grasping and climbing. All tetrapods have four limbs that are organized into two bilaterally symmetrical pairs, with one pair at each end of the torso, which phylogenetrically correspond to the four paired fins (pectoral and pelvic fins) of their fish ancestors.
Hox geneHox 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 (for example, legs, antennae, and wings in fruit flies), and Hox genes in vertebrates specify the types and shape of vertebrae that will form.
ExaptationExaptation and the related term co-option describe a shift in the function of a trait during evolution. For example, a trait can evolve because it served one particular function, but subsequently it may come to serve another. Exaptations are common in both anatomy and behaviour. Bird feathers are a classic example. Initially they may have evolved for temperature regulation, but later were adapted for flight. When feathers were first used to aid in flight, that was an exaptive use.
HomeoboxA homeobox is a DNA sequence, around 180 base pairs long, that regulates large-scale anatomical features in the early stages of embryonic development. Mutations in a homeobox may change large-scale anatomical features of the full-grown organism. Homeoboxes are found within genes that are involved in the regulation of patterns of anatomical development (morphogenesis) in animals, fungi, plants, and numerous single cell eukaryotes.
Homeotic geneHomeotic genes are genes which regulate the development of anatomical structures in various organisms such as echinoderms, insects, mammals, and plants. Homeotic genes often encode transcription factor proteins, and these proteins affect development by regulating downstream gene networks involved in body patterning. Mutations in homeotic genes cause displaced body parts (homeosis), such as antennae growing at the posterior of the fly instead of at the head. Mutations that lead to development of ectopic structures are usually lethal.
Gene regulatory networkA gene (or genetic) regulatory network (GRN) is a collection of molecular regulators that interact with each other and with other substances in the cell to govern the gene expression levels of mRNA and proteins which, in turn, determine the function of the cell. GRN also play a central role in morphogenesis, the creation of body structures, which in turn is central to evolutionary developmental biology (evo-devo).
Hedgehog signaling pathwayThe Hedgehog signaling pathway is a signaling pathway that transmits information to embryonic cells required for proper cell differentiation. Different parts of the embryo have different concentrations of hedgehog signaling proteins. The pathway also has roles in the adult. Diseases associated with the malfunction of this pathway include cancer. The Hedgehog signaling pathway is one of the key regulators of animal development and is present in all bilaterians.
TetrapodTetrapods ('tɛtrəˌpɒdz; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (tɛ'træpədə). It includes all extant and extinct amphibians, and the amniotes which in turn evolved into the sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids (extinct pelycosaurs, therapsids and all extant mammals). Some tetrapods such as snakes, legless lizards and caecilians had evolved to become limbless via mutations of the Hox gene, although some do still have a pair of vestigial spurs that are remnants of the hindlimbs.
AdaptationIn biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the population during that process. Thirdly, it is a phenotypic trait or adaptive trait, with a functional role in each individual organism, that is maintained and has evolved through natural selection. Historically, adaptation has been described from the time of the ancient Greek philosophers such as Empedocles and Aristotle.
Regulatory sequenceA regulatory sequence is a segment of a nucleic acid molecule which is capable of increasing or decreasing the expression of specific genes within an organism. Regulation of gene expression is an essential feature of all living organisms and viruses. In DNA, regulation of gene expression normally happens at the level of RNA biosynthesis (transcription). It is accomplished through the sequence-specific binding of proteins (transcription factors) that activate or inhibit transcription.
Upper limbThe upper limbs or upper extremities are the forelimbs of an upright-postured tetrapod vertebrate, extending from the scapulae and clavicles down to and including the digits, including all the musculatures and ligaments involved with the shoulder, elbow, wrist and knuckle joints. In humans, each upper limb is divided into the arm, forearm and hand, and is primarily used for climbing, lifting and manipulating objects. In formal usage, the term "arm" only refers to the structures from the shoulder to the elbow, explicitly excluding the forearm, and thus "upper limb" and "arm" are not synonymous.
Natural selectionNatural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Charles Darwin popularised the term "natural selection", contrasting it with artificial selection, which is intentional, whereas natural selection is not. Variation exists within all populations of organisms. This occurs partly because random mutations arise in the genome of an individual organism, and their offspring can inherit such mutations.
Cis-regulatory elementCis-regulatory elements (CREs) or Cis''-regulatory modules (CRMs) are regions of non-coding DNA which regulate the transcription of neighboring genes. CREs are vital components of genetic regulatory networks, which in turn control morphogenesis, the development of anatomy, and other aspects of embryonic development, studied in evolutionary developmental biology. CREs are found in the vicinity of the genes that they regulate. CREs typically regulate gene transcription by binding to transcription factors.
Teleology in biologyTeleology in biology is the use of the language of goal-directedness in accounts of evolutionary adaptation, which some biologists and philosophers of science find problematic. The term teleonomy has also been proposed. Before Darwin, organisms were seen as existing because God had designed and created them; their features such as eyes were taken by natural theology to have been made to enable them to carry out their functions, such as seeing.
Evo-devo gene toolkitThe 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. Toolkit genes are highly conserved among phyla, meaning that they are ancient, dating back to the last common ancestor of bilaterian animals.
Symmetry in biologySymmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, the face of a human being has a plane of symmetry down its centre, or a pine cone displays a clear symmetrical spiral pattern. Internal features can also show symmetry, for example the tubes in the human body (responsible for transporting gases, nutrients, and waste products) which are cylindrical and have several planes of symmetry.
Regulator geneA regulator gene, regulator, or regulatory gene is a gene involved in controlling the expression of one or more other genes. Regulatory sequences, which encode regulatory genes, are often at the five prime end (5') to the start site of transcription of the gene they regulate. In addition, these sequences can also be found at the three prime end (3') to the transcription start site. In both cases, whether the regulatory sequence occurs before (5') or after (3') the gene it regulates, the sequence is often many kilobases away from the transcription start site.
EvolutionIn biology, evolution is the change in heritable characteristics of biological populations over successive generations. Evolution occurs when evolutionary processes such as natural selection (including sexual selection) and genetic drift act on genetic variation, resulting in certain characteristics becoming more or less common within a population over successive generations. The process of evolution has given rise to biodiversity at every level of biological organisation.