Hox gene - Knowledge (XXG)

1232:, something fundamental for them to be able to fly, being transformed into a wing, or an antenna turning into a leg; in the mouse, two to four genes must be simultaneously removed to get a similar complete transformation. Some researchers believe that, because of the redundancy of the vertebrate HOX cluster plan and more constrained compared to invertebrate HOX clusters, the evolvability of vertebrate HOX clusters is, for some structural or functional reason, far lower than their invertebrate counterparts. This rapid evolvability is in part because invertebrates experienced much more dramatic episodes of 557:

at the bottom of such hierarchies to ultimately form the tissues, structures, and organs of each segment. Segmentation involves such processes as morphogenesis (differentiation of precursor cells into their terminal specialized cells), the tight association of groups of cells with similar fates, the sculpting of structures and segment boundaries via programmed cell death, and the movement of cells from where they are first born to where they will ultimately function, so it is not surprising that the target genes of Hox genes promote cell division, cell adhesion,

160: 203:. This implies that Hox genes arose over 550 million years ago. In bilateria, Hox genes are often arranged in gene clusters, although there are many exceptions where the genes have been separated by chromosomal rearrangements. Comparing homeodomain sequences between Hox proteins often reveals greater similarity between species than within a species; this observation led to the conclusion that Hox gene clusters evolved early in animal evolution from a single Hox gene via tandem 232: 528:

identify the set of proteins between two different species that are most likely to be most similar in function, classification schemes are used. For Hox proteins, three different classification schemes exist: phylogenetic inference based, synteny-based, and sequence similarity-based. The three classification schemes provide conflicting information for Hox proteins expressed in the middle of the body axis (

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actual functional sites. Especially for Hox proteins, which produce such dramatic changes in morphology when misexpressed, this raises the question of how each transcription factor can produce such specific and different outcomes if they all bind the same sequence. One mechanism that introduces greater DNA sequence specificity to Hox proteins is to bind protein cofactors. Two such Hox cofactors are

1212:, but the location is different. For example, there are more HOX genes on the 5' side of the mouse segment compared to the invertebrates. These genes correspond to expression in the tail, which would make sense as flies would not have anything similar to the tail that all vertebrates have. Additionally, in most vertebrates there are 39 members segregated into four separate tightly clustered 775:. For example, high concentrations of one maternal protein and low concentrations of others will turn on a specific set of gap or pair-rule genes. In flies, stripe 2 in the embryo is activated by the maternal proteins Bicoid and Hunchback, but repressed by the gap proteins Giant and Kruppel. Thus, stripe 2 will only form wherever there is Bicoid and Hunchback, but 412: 214:, Hox genes are expressed in staggered domains along the head-to-tail axis of the embryo, suggesting that their role in specifying position is a shared, ancient feature. The functional conservation of Hox proteins can be demonstrated by the fact that a fly can function to a large degree with a chicken Hox protein in place of its own. So, despite having a 862: 719:

terminal T being the most important for binding. This sequence is conserved in nearly all sites recognized by homeodomains, and probably distinguishes such locations as DNA binding sites. The base pairs following this initial sequence are used to distinguish between homeodomain proteins, all of which

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Hox genes act at many levels within developmental gene hierarchies: at the "executive" level they regulate genes that in turn regulate large networks of other genes (like the gene pathway that forms an appendage). They also directly regulate what are called realisator genes or effector genes that act

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sequence known as the homeobox, of which the term "Hox" was originally a contraction. However, in current usage the term Hox is no longer equivalent to homeobox, because Hox genes are not the only genes to possess a homeobox sequence; for instance, humans have over 200 homeobox genes, of which 39 are

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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

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In some organisms, especially vertebrates, the various Hox genes are situated very close to one another on the chromosome in groups or clusters. The order of the genes on the chromosome is the same as the expression of the genes in the developing embryo, with the first gene being expressed in the

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An analogy for the Hox genes can be made to the role of a play director who calls which scene the actors should carry out next. If the play director calls the scenes in the wrong order, the overall play will be presented in the wrong order. Similarly, mutations in the Hox genes can result in body

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Studies on Hox genes in ciliated larvae have shown they are only expressed in future adult tissues. In larvae with gradual metamorphosis the Hox genes are activated in tissues of the larval body, generally in the trunk region, that will be maintained through metamorphosis. In larvae with complete

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Hox genes play critical roles in the development of structures such as limbs, lungs, the nervous system, and eyes. As T. R. Lappin and colleagues observed in 2006, "Evolutionary conservation provides unlimited scope for experimental investigation of the functional control of the Hox gene network

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The genetic studies by Morgan and others provided the foundation for the systematic analyses of Edward B. Lewis and Thomas Kaufman, which provided preliminary definitions of the many homeotic genes of the Bithorax and Antennapedia complexes, and also showed that the mutant phenotypes for most of

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Proteins with a high degree of sequence similarity are also generally assumed to exhibit a high degree of functional similarity, i.e. Hox proteins with identical homeodomains are assumed to have identical DNA-binding properties (unless additional sequences are known to influence DNA-binding). To

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In mouse embryos, the HOX10 genes, which is one of the genes that lie in the tail portion of the animal, turn the "rib-building" system off when the gene is activated. The genes are active in the lower back, where the vertebrae do not grow ribs, and inactive in the mid-back, allowing ribs to be

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However, all homeodomain-containing transcription factors bind essentially the same DNA sequence. The sequence bound by the homeodomain of a Hox protein is only six nucleotides long, and such a short sequence would be found at random many times throughout the genome, far more than the number of

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regulates differential expression of Hox genes along the anteroposterior axis. Genes in the 3' ends of Hox clusters are induced by retinoic acid resulting in expression domains that extend more anteriorly in the body compared to 5' Hox genes that are not induced by retinoic acid resulting in

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The Hox genes are named for the homeotic phenotypes that result when their function is disrupted, wherein one segment develops with the identity of another (e.g. legs where antennae should be). Hox genes in different phyla have been given different names, which has led to confusion about

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Helix 1 Helix 2 Helix 3/4 ______________ __________ _________________ RRRKRTAYTRYQLLELEKEFLFNRYLTRRRRIELAHSLNLTERHIKIWFQNRRMKWKKEN ....|....|....|....|....|....|....|....|....|....|....|....| 10 20 30 40 50

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are experimentally inactivated, the vertebrae of the lower back grow ribs. This research prompted an evolutionary search for these mutations among all animals. An example of this is in lizards and snakes. In snakes, HOX10 genes have lost their rib-blocking ability in that way.

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gene was initially so named because it disrupted the labial appendage; however, the lab gene is not expressed in the labial segment, and the labial appendage phenotype is likely a result of the broad disorganization resulting from the failure of head involution.

768:. This results in a transcription factor cascade: maternal factors activate gap or pair-rule genes; gap and pair-rule genes activate Hox genes; then, finally, Hox genes activate realisator genes that cause the segments in the developing embryo to differentiate. 1187:

of the back, and the skeletal muscles of the body wall and limbs. HOX genes help differentiate somite cells into more specific identities and direct them to develop differently depending on where they are in the body. A large difference between vertebrates and

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bodies are not segmented in the same way as insects; they are on average much more complex, leading to more infrastructure in their body plan compared to insects. HOX genes control the regulation and development of many key structures in the body, such as

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which occurred in their evolutionary history. In zebrafish, one of the eight Hox gene clusters (a Hoxd cluster) has lost all protein-coding genes, and just a single microRNA gene marks the location of the original cluster. In some teleost fish, such as

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through which they either activate or repress hundreds of other genes. The same Hox protein can act as a repressor at one gene and an activator at another. The ability of Hox proteins to bind DNA is conferred by a part of the protein referred to as the

540:). A combined approach used phylogenetic inference-based information of the different species and plotted the protein sequence types onto the phylogenetic tree of the species. The approach identified the proteins that best represent ancestral forms ( 833:

anterior end of the developing organism. The reason for this colinearity is not yet completely understood, but could be related to the activation of Hox genes in a temporal sequence by gradual unpacking of chromatin along a gene cluster.

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patterns T3 largely by repressing genes involved in wing formation. The wing blade is composed of two layers of cells that adhere tightly to one another, and are supplied with nutrient by several wing veins. One of the many genes that

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Kulakova M, Bakalenko N, Novikova E, Cook CE, Eliseeva E, Steinmetz PR, et al. (January 2007). "Hox gene expression in larval development of the polychaetes Nereis virens and Platynereis dumerilii (Annelida, Lophotrochozoa)".

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is the same as the order of their expression along the anterior-posterior axis of the developing animal, and are thus said to display colinearity. Production of Hox gene products at wrong location in the body is associated with

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in the embryo results in a failure of head involution (see labial gene), with a loss of larval head structures. Mutations in the adult have either deletions of parts of the head or transformations of head to thoracic identity.

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have similar recognition sites. For instance, the nucleotide following the TAAT sequence is recognized by the amino acid at position 9 of the homeodomain protein. In the maternal protein Bicoid, this position is occupied by

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embryo to internalize the mouth and head structures that initially develop on the outside of its body (a process called head involution). Failure of head involution disrupts or deletes the salivary glands and pharynx. The

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Vachon G, Cohen B, Pfeifle C, McGuffin ME, Botas J, Cohen SM (October 1992). "Homeotic genes of the Bithorax complex repress limb development in the abdomen of the Drosophila embryo through the target gene Distal-less".

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is made up of two clusters, the Antennapedia complex and the Bithorax complex, which together were historically referred to as the HOM-C (for Homeotic Complex). Although historically HOM-C genes have referred to

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is an important model for understanding body plan generation and evolution. The general principles of Hox gene function and logic elucidated in flies will apply to all bilaterian organisms, including humans.

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Due to the fact that the HOX genes are so highly conserved, most research has been done on much simpler model organisms, such as mice. One of the major differences that was noticed when comparing mice and

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Quantitative PCR has shown several trends regarding colinearity: the system is in equilibrium and the total number of transcripts depends on the number of genes present according to a linear relationship.

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is misexpressed in the second thoracic segment, such as occurs in flies with the "Cbx" enhancer mutation, it represses wing genes, and the wings develop as halteres, resulting in a four-haltered fly.

1360:). It transforms the third thoracic segment (T3) toward the second (T2). Bithorax arose spontaneously in the laboratory and has been maintained continuously as a laboratory stock ever since. 120:

The products of Hox genes are Hox proteins. Hox proteins are a subset of transcription factors, which are proteins that are capable of binding to specific nucleotide sequences on DNA called

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and mutations. More than 20 major clades of invertebrates differ so radically in body organization, partly due to a higher mutation rate, that they became formally classified as different

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which is providing important insights into human disease." In the future, more research can be done in investigating the roles of Hox genes in leukaemia and cancer (such as EOC).

252:, like all insects, has eight Hox genes. These are clustered into two complexes, both of which are located on chromosome 3. The Antennapedia complex (not to be confused with the 58:

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.

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Definitive evidence for a genetic basis of some homeotic transformations was obtained by isolating homeotic mutants. The first homeotic mutant was found by Calvin Bridges in

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and subsequent divergence, and that a prototypic Hox gene cluster containing at least seven different Hox genes was present in the common ancestor of all bilaterian animals.

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strands located in Hox clusters have been shown to inhibit more anterior hox genes ("posterior prevalence phenomenon"), possibly to better fine tune its expression pattern.

1446: 1349:'s genetic principles, Bateson and others realized that some examples of homeosis in floral organs and animal skeletons could be attributed to variation in genes. 145:

chain is shown below: Hox proteins often act in partnership with co-factors, such as PBC and Meis proteins encoded by very different types of homeobox gene.

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to be expressed in the antennal imaginal disc, so that, instead of forming an antenna, the disc makes a leg, resulting in a leg coming out of the fly's head.

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de Rosa R, Grenier JK, Andreeva T, Cook CE, Adoutte A, Akam M, et al. (June 1999). "Hox genes in brachiopods and priapulids and protostome evolution".

4830: 744:(Exd) and Homothorax (Hth). Exd and Hth bind to Hox proteins and appear to induce conformational changes in the Hox protein that increase its specificity. 78:

parts and limbs in the wrong place along the body. Like a play director, the Hox genes do not act in the play or participate in limb formation themselves.

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to give four Hox gene clusters: Hoxa, Hoxb, Hoxc and Hoxd. It is currently unclear whether these duplications occurred before or after the divergence of

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that lived over 550 million years ago, the chicken and fly version of the same Hox gene are similar enough to target the same downstream genes in flies.

2510:"Analysis of central Hox protein types across bilaterian clades: on the diversification of central Hox proteins from an Antennapedia/Hox7-like protein" 4820: 853:

homologues, while Hox genes referred to vertebrate homologues, this distinction is no longer made, and both HOM-C and Hox genes are called Hox genes.

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is the location and layering of HOX genes. The fundamental mechanisms of development are strongly conserved among vertebrates from fish to mammals.

3543:"Enigmatic orthology relationships between Hox clusters of the African butterfly fish and other teleosts following ancient whole-genome duplication" 1158:, an even more recent genome duplication occurred, doubling the seven or eight Hox gene clusters to give at least 13 clusters Another teleost, the 2136:"Rescue of Drosophila labial null mutant by the chicken ortholog Hoxb-1 demonstrates that the function of Hox genes is phylogenetically conserved" 2353:"The homeotic gene Sex combs reduced of Drosophila melanogaster is differentially regulated in the embryonic and imaginal stages of development" 434:

The third thoracic segment, or T3, bears a pair of legs and a pair of halteres (highly reduced wings that function in balancing during flight).

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Hox genes. Hox genes are thus a subset of the homeobox transcription factor genes. In many animals, the organization of the Hox genes in the

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represses is blistered, which activates proteins involved in cell-cell adhesion, and spalt, which patterns the placement of wing veins. In

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metamorphosis the Hox genes are mainly expressed in juvenile rudiments and are absent in the transient larval tissues. The larvae of the

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Capovilla M, Botas J (December 1998). "Functional dominance among Hox genes: repression dominates activation in the regulation of Dpp".

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identified and classified 15 genes of key importance in determining the body plan and the formation of body segments of the fruit fly

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no longer represses wing genes, and the halteres develop as a second pair of wings, resulting in the famous four-winged flies. When

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is ectopically expressed throughout the embryo, all segments anterior of A4 are transformed to an A4-like abdominal identity. The

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Lee PN, Callaerts P, De Couet HG, Martindale MQ (August 2003). "Cephalopod Hox genes and the origin of morphological novelties".

548:) and the proteins that represent new, derived versions (or were lost in an ancestor and are now missing in numerous species). 736:. If the lysine in Bicoid is replaced by glutamine, the resulting protein will recognize Antennapedia-binding enhancer sites. 396:

gene specifies this identity by promoting leg formation and allowing (but not directly activating) wing formation. A dominant

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gene is responsible for the formation of the maxillary and mandibular segments in the larval head. The mutant phenotypes of

4593: 3408:"Comparative phylogenomic analyses of teleost fish Hox gene clusters: lessons from the cichlid fish Astatotilapia burtoni" 1121:

The ancestors of vertebrates had a single Hox gene cluster, which was duplicated (twice) early in vertebrate evolution by

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abdomen. Both the regulatory protein and the morphogenic protein are involved in the development of the tail segment.

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Daniels LK (January 1976). "Rapid in-office and in-vivo desensitization of an injection phobia utilizing hypnosis".

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Hanes SD, Brent R (January 1991). "A genetic model for interaction of the homeodomain recognition helix with DNA".

1972:"Pre-bilaterian origins of the Hox cluster and the Hox code: evidence from the sea anemone, Nematostella vectensis" 1390: 276:). The Bithorax complex, named after the Ultrabithorax gene, consists of the remaining three genes: Ultrabithorax ( 3345:

Spagnuolo, A., Ristoratore, F., Di Gregorio, A., Aniello, F., Branno, M. & Di Lauro, R. (2003) Gene 309, 71–79

2636:"The Drosophila Hox gene deformed sculpts head morphology via direct regulation of the apoptosis activator reaper" 4445: 4188:

Nüsslein-Volhard C, Wieschaus E (October 1980). "Mutations affecting segment number and polarity in Drosophila".

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Pearson JC, Lemons D, McGinnis W (December 2005). "Modulating Hox gene functions during animal body patterning".

1356:'s laboratory in 1915. This mutant shows a partial duplication of the thorax and was therefore named Bithorax ( 1159: 1122: 483:

loss-of-function mutants, abdominal segments A2 through A8 are transformed into an identity more like A1. When

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changes for the most part. This is also one reason why homeotic mutations in vertebrates are so rarely seen.

3960:"Evolution of Antp-class genes and differential expression of Hydra Hox/paraHox genes in anterior patterning" 3057:

Fraser P, Bickmore W (May 2007). "Nuclear organization of the genome and the potential for gene regulation".

4686: 4623: 4598: 4481: 4401: 3357:"A new look at an old question: when did the second whole genome duplication occur in vertebrate evolution?" 796: 327: 243: 3457:

Woltering JM, Durston AJ (June 2006). "The zebrafish hoxDb cluster has been reduced to a single microRNA".

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Garcia-Fernández J, Holland PW (August 1994). "Archetypal organization of the amphioxus Hox gene cluster".

2304:"Developmental and molecular analysis of Deformed; a homeotic gene controlling Drosophila head development" 159: 4825: 4671: 4570: 4555: 4524: 4425: 2763:"DNA specificity of the bicoid activator protein is determined by homeodomain recognition helix residue 9" 1306: 1290: 791:(ncRNA) has been shown to be abundant in Hox clusters. In humans, 231 ncRNA may be present. One of these, 301: 103: 795:, silences in trans (it is transcribed from the HOXC cluster and inhibits late HOXD genes) by binding to 304:(an appendageless segment between the antenna and mandible), and also in the midgut. Loss of function of 4782: 4676: 4603: 4502: 4440: 4435: 4366: 4301: 3909:

Holland LZ, Albalat R, Azumi K, Benito-Gutiérrez E, Blow MJ, Bronner-Fraser M, et al. (July 2008).

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Just as Hox genes regulate realisator genes, they are in turn regulated themselves by other genes. In

4691: 4534: 4529: 4411: 4406: 4393: 4197: 4131: 4030: 3863: 3304: 3066: 2819: 2462: 2188: 1983: 1925: 1839: 1785: 1730: 1479: 807: 82: 3159:"Modeling Hox gene regulation in digits: reverse collinearity and the molecular origin of thumbness" 4787: 3010:"Functional demarcation of active and silent chromatin domains in human HOX loci by noncoding RNAs" 1402: 772: 511:

is transcribed in two different forms, a regulatory protein, and a morphogenic protein. Regulatory

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The DNA sequence bound by the homeodomain protein contains the nucleotide sequence TAAT, with the

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Lempradl A, Ringrose L (February 2008). "How does noncoding transcription regulate Hox genes?".

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Ryan JF, Mazza ME, Pang K, Matus DQ, Baxevanis AD, Martindale MQ, et al. (January 2007).

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structure is essential for transcription but it also requires the cluster to loop out of the

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Gaunt SJ (2018). "Hox cluster genes and collinearities throughout the tree of animal life".

2065: 2001: 1991: 1933: 1883: 1873: 1803: 1793: 1738: 1684: 1642: 1634: 1593: 1583: 1542: 1532: 1487: 1406: 1394: 1389:

In 1983, the homeobox was discovered independently by researchers in two labs: Ernst Hafen,

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is necessary to specify the identity of most of the abdominal segments. A major function of

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is expressed along most of the abdomen, from abdominal segments 1 (A1) to A8. Expression of

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DNA sequence, the homeobox). This amino acid sequence folds into a "helix-turn-helix" (i.e.

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Gauchat D, Mazet F, Berney C, Schummer M, Kreger S, Pawlowski J, et al. (April 2000).

4757: 4661: 4471: 3762:"Hox C cluster genes are dispensable for overall body plan of mouse embryonic development" 1772:

Janmaat VT, Nesteruk K, Spaander MC, Verhaar AP, Yu B, Silva RA, et al. (June 2021).

1718: 1367: 1342: 1184: 1162:, has instead seen a significant loss in HOX gene clusters, with only 5 clusters present. 300:

gene is the most anteriorly expressed gene. It is expressed in the head, primarily in the

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Hughes CL, Kaufman TC (2002). "Hox genes and the evolution of the arthropod body plan".

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suppress embryonic ventral epidermal structures in the eighth and ninth segments of the

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is responsible for the formation of the labial and maxillary palps. Some evidence shows

4055: 4010: 3935: 3910: 3886: 3851: 3827: 3802: 3737: 3710: 3651: 3626: 3567: 3542: 3434: 3407: 3383: 3356: 3229: 3212: 3183: 3158: 3134: 3109: 3034: 3009: 2933: 2889: 2854: 2485: 2450: 2426: 2402:"Developmental genetic analysis of Contrabithorax mutations in Drosophila melanogaster" 2401: 2377: 2352: 2319: 2006: 1971: 1888: 1861: 1808: 1773: 1647: 1622: 1598: 1571: 1547: 1516: 1398: 1213: 788: 761: 2942: 2917: 2870: 2709: 2684: 2652: 2635: 2328: 2303: 1831: 4814: 4725: 4639: 4507: 4341: 4318: 3986: 3959: 3265: 3008:

Rinn JL, Kertesz M, Wang JK, Squazzo SL, Xu X, Brugmann SA, et al. (June 2007).

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Hueber SD, Rauch J, Djordjevic MA, Gunter H, Weiller GF, Frickey T (November 2013).

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The second thoracic segment, or T2, develops a pair of legs and a pair of wings. The

4550: 4371: 4361: 4225: 4159: 4108: 3687: 3332: 3094: 2685:"p21 is a transcriptional target of HOXA10 in differentiating myelomonocytic cells" 1953: 1758: 1364:

these genes could be traced back to patterning defects in the embryonic body plan.

1221: 1189: 1138: 1074: 903: 416: 387: 168: 63: 38: 3727: 2288: 2177:"Origin of the metazoan phyla: molecular clocks confirm paleontological estimates" 1704: 1200:, in particular, has to do with the location and layering of HOX genes within the 231: 3911:"The amphioxus genome illuminates vertebrate origins and cephalochordate biology" 2994: 2475: 2417: 2368: 1996: 1382:

in 1980. For their work, Lewis, Nüsslein-Volhard, and Wieschaus were awarded the

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Proceedings of the National Academy of Sciences of the United States of America

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Proceedings of the National Academy of Sciences of the United States of America

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Proceedings of the National Academy of Sciences of the United States of America

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Ferrier DE, Minguillón C (2003). "Evolution of the Hox/ParaHox gene clusters".

1798: 1721:(August 1995). "Homeotic genes and the evolution of arthropods and chordates". 4730: 4430: 4383: 4092: 4045: 3373: 1638: 1537: 1492: 1467: 1217: 1197: 1171: 886: 753: 180: 130: 99: 94: 3500:

Mungpakdee S, Seo HC, Angotzi AR, Dong X, Akalin A, Chourrout D (July 2008).

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Discredited hypotheses for the Cambrian explosion (Section: Regulatory genes)

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probably had three body regions, each with a unique combination of Hox genes.

171:) by up to three segments. Segments with maxillopeds have Hox gene 7. Fossil 4681: 4649: 4416: 4322: 3876: 3558: 3518: 3501: 3424: 2831: 2612: 1265: 1245: 1142: 803: 729: 558: 211: 192: 184: 172: 164: 134: 4151: 4100: 4064: 3995: 3976: 3944: 3895: 3836: 3787: 3778: 3761: 3746: 3660: 3611: 3576: 3527: 3478: 3443: 3392: 3238: 3192: 3143: 3086: 3043: 2986: 2898: 2718: 2661: 2535: 2494: 2337: 2260: 2120: 2042: 2015: 1945: 1897: 1817: 1696: 1656: 1607: 1588: 1556: 1501: 167:. The Hox genes 7, 8, and 9 correspond in these groups but are shifted (by 4217: 3926: 3324: 3273: 2951: 2839: 2788: 2620: 2574: 2435: 2386: 2220: 2201: 2161: 2112: 2077: 1750: 411: 183:. The Hox genes, being a subset of homeobox genes, arose more recently in 179:

Homeobox genes, and thus the homeodomain protein motif, are found in most

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10.1002/(sici)1521-1878(200001)22:1<72::aid-bies12>3.0.co;2-s

2734:"The Origins of Anterior-Posterior Polarity: The Homeotic Selector Genes" 2152: 2135: 1338: 1326: 1322: 1311: 1295: 1229: 1225: 1134: 782: 757: 496: 492: 196: 71: 54: 34: 17: 4143: 3174: 3078: 4376: 3695: 3470: 2978: 2700: 2451:"Improving Hox protein classification across the major model organisms" 1130: 1126: 861: 733: 725: 188: 195:(animals with a clear head-to-tail axis), and have also been found in 4356: 4209: 4175:

Master Control Genes in Development and Evolution: The Homeobox Story

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and some insects (but not most animals), Hox genes are regulated by

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Expression of Hox genes in the body segments of different groups of

4035: 3252:

Scott MP (November 1992). "Vertebrate homeobox gene nomenclature".

1774:"HOXA13 in etiology and oncogenic potential of Barrett's esophagus" 771:

Regulation is achieved via protein concentration gradients, called

3157:

Montavon T, Le Garrec JF, Kerszberg M, Duboule D (February 2008).

3110:"Retinoic acid synthesis and signaling during early organogenesis" 1937: 1345:

in 1894, who coined the term "homeosis". After the rediscovery of

1096: 1092: 1088: 1084: 1080: 1043: 1039: 1035: 1031: 1027: 1002: 998: 994: 990: 986: 982: 978: 974: 970: 937: 933: 929: 925: 921: 917: 913: 909: 860: 410: 230: 158: 3852:"cis-regulatory change associated with snake body plan evolution" 2400:

González-Gaitán MA, Micol JL, García-Bellido A (September 1990).

1220:, whereas there are eight HOX genes in total for the Drosophila. 1149:, have seven or eight Hox gene clusters because of an additional 4654: 1341:. Homeotic transformations were first identified and studied by 1209: 4290: 4015:

gene expression in postmetamorphic juveniles of the brachiopod

3627:"Hox genes and regional patterning of the vertebrate body plan" 3502:"Differential evolution of the 13 Atlantic salmon Hox clusters" 2134:

Lutz B, Lu HC, Eichele G, Miller D, Kaufman TC (January 1996).

2056:

Krumlauf R (July 1994). "Hox genes in vertebrate development".

2918:"Regulation of even-skipped stripe 2 in the Drosophila embryo" 2855:"Hox specificity unique roles for cofactors and collaborators" 1208:

to those of the fly as it is one of the most highly conserved

191:. Within the animal kingdom, Hox genes are present across the 89: 3211:

Lappin TR, Grier DG, Thompson A, Halliday HL (January 2006).

2302:

Regulski M, McGinnis N, Chadwick R, McGinnis W (March 1987).

1572:"Classification and nomenclature of all human homeobox genes" 374:

gene is responsible for cephalic and thoracic development in

2449:

Hueber SD, Weiller GF, Djordjevic MA, Frickey T (May 2010).

3590:

Dale KJ, Pourquié O (January 2000). "A clock-work somite".

2634:

Lohmann I, McGinnis N, Bodmer M, McGinnis W (August 2002).

1337:

The Hox genes are so named because mutations in them cause

764:, which are in their turn regulated by maternally-supplied 141:) motif that is stabilized by a third helix. The consensus 133:-long DNA-binding domain (encoded by its corresponding 180- 1244:

genes need to be knocked out in order for there to be any

680:

distal limb that will form digit, carpal and tarsal bones

106:

is the result of altered Hox coding and is a precursor to

1517:"Hox gene expression during development of the phoronid 695:

Cell cycle: differentiation of myelomonocyte cells into

3406:

Hoegg S, Boore JL, Kuehl JV, Meyer A (September 2007).

2282: 2280: 2278: 697:

monocytes (white blood cells), with cell cycle arrest

641:

boundary between the maxilla and mandible of the head

639:

Apoptosis: localized cell death creates the segmental

491:

also affects the pattern of cuticle generation in the

3213:"HOX genes: seductive science, mysterious mechanisms" 1670: 1668: 1666: 1228:. In flies, one gene can be mutated, resulting in a 876:

Mice and humans have 39 Hox genes in four clusters:

678:

Cell adhesion: causes tight association of cells in

357:

are similar to those of labial. Loss of function of

4775: 4704: 4632: 4586: 4579: 4543: 4495: 4459: 4392: 4329: 1862:"The TALE face of Hox proteins in animal evolution" 1224:are far more redundant and less likely to generate 2101:The International Journal of Developmental Biology 2031:The International Journal of Developmental Biology 1272:have a single Hox cluster with 15 genes, known as 656:prevents the above cell changes in more posterior 3206: 3204: 3202: 1570:Holland PW, Booth HA, Bruford EA (October 2007). 4240:"The Nobel Prize in Physiology or Medicine 1995" 728:. In Antennapedia, this position is occupied by 622:triggers cell shape changes in the gut that are 3625:Mallo M, Wellik DM, Deschamps J (August 2010). 2175:Ayala FJ, Rzhetsky A, Ayala FJ (January 1998). 1314:, have a few Hox/ParaHox-like homeobox genes. 818:expression domains that remain more posterior. 724:, which recognizes and binds to the nucleotide 4302: 2594: 2592: 2547: 2545: 2234: 2232: 2230: 1317:Hox gene expression has also been studied in 1288:Six Hox genes are dispersed in the genome of 844:nomenclature. The complement of Hox genes in 102:and predisposes to oncological disease, e.g. 8: 1911: 1909: 1907: 1515:Gąsiorowski L, Hejnol A (10 February 2020). 479:in insects is to repress limb formation. In 3355:Holland LZ, Ocampo Daza D (November 2018). 1838:. Department of Biosciences and Nutrition, 1409:and Amy Weiner (in Thomas Kaufman's lab at 4583: 4309: 4295: 4287: 2683:Bromleigh VC, Freedman LP (October 2000). 2351:Pattatucci AM, Kaufman TC (October 1991). 608:activates gene pathway for limb formation 594:activates gene pathway for limb formation 495:, and pattern of muscle generation in the 81:The protein product of each Hox gene is a 4054: 4044: 4034: 3985: 3975: 3934: 3885: 3875: 3826: 3777: 3736: 3726: 3676:The American Journal of Clinical Hypnosis 3650: 3566: 3517: 3433: 3423: 3382: 3372: 3228: 3182: 3133: 3033: 2941: 2888: 2778: 2708: 2651: 2525: 2484: 2474: 2425: 2376: 2327: 2210: 2200: 2151: 2005: 1995: 1887: 1877: 1807: 1797: 1646: 1597: 1587: 1546: 1536: 1491: 1204:. Vertebrates do have HOX genes that are 1179:, which form the vertebrae and ribs, the 1468:"Larval Evolution: I'll Tail You Later…" 878: 711:Enhancer sequences bound by homeodomains 624:required for normal visceral morphology 563: 2859:Current Topics in Developmental Biology 1965: 1963: 1466:Hejnol A, Vellutini BC (January 2017). 1458: 4799:Index of evolutionary biology articles 3541:Martin KJ, Holland PW (October 2014). 256:gene) consists of five genes: labial ( 1384:Nobel Prize in Physiology or Medicine 7: 2740:(6th ed.). Sinauer Associates. 1621:Bürglin TR, Affolter M (June 2016). 1437:Hox genes in amphibians and reptiles 1167:Hox genes in amphibians and reptiles 813:In higher animals including humans, 3760:Suemori H, Noguchi S (April 2000). 3709:Hrycaj SM, Wellik DM (2016-05-10). 2853:Mann RS, Lelli KM, Joshi R (2009). 1137:have four HOX clusters, while most 4831:Evolutionary developmental biology 4609:Evolutionary developmental biology 2934:10.1002/j.1460-2075.1992.tb05498.x 2320:10.1002/j.1460-2075.1987.tb04819.x 779:where there is Giant and Kruppel. 235:Homeobox (Hox) gene expression in 25: 2818:(4992). New York, N.Y.: 426–430. 1623:"Homeodomain proteins: an update" 1411:Indiana University in Bloomington 4821:Developmental genes and proteins 4566:Evolution of sexual reproduction 4009:Gąsiorowski L, Hejnol A (2018). 2253:10.1046/j.1525-142x.2002.02034.x 732:, which recognizes and binds to 43:specify regions of the body plan 4081:Development Genes and Evolution 3547:Molecular Biology and Evolution 3506:Molecular Biology and Evolution 2761:Hanes SD, Brent R (June 1989). 576:Normal function of target gene 4337:Genotype–phenotype distinction 3688:10.1080/00029157.1976.10403798 523:Classification of Hox proteins 308:results in the failure of the 1: 4594:Regulation of gene expression 3728:10.12688/f1000research.7663.1 2871:10.1016/S0070-2153(09)88003-4 2653:10.1016/s0092-8674(02)00871-1 1860:Merabet S, Galliot B (2015). 1832:"Helix 1, Helix 2, Helix 3/4" 1133:from other vertebrates. Most 187:within the animal kingdom or 4764:Endless Forms Most Beautiful 4544:Evolution of genetic systems 4352:Gene–environment correlation 4347:Gene–environment interaction 3266:10.1016/0092-8674(92)90588-4 3108:Duester G (September 2008). 2780:10.1016/0092-8674(89)90063-9 2567:10.1016/0092-8674(92)90513-C 2476:10.1371/journal.pone.0010820 2070:10.1016/0092-8674(94)90290-9 1997:10.1371/journal.pone.0000153 865:Hox genes in various species 664:(represses decapentaplegic) 630:(activates decapentaplegic) 4743:Christiane Nüsslein-Volhard 4263:"The Function of Hox Genes" 3643:10.1016/j.ydbio.2010.04.024 2527:10.1016/j.ydbio.2013.09.009 2241:Evolution & Development 1372:Christiane Nüsslein-Volhard 85:. Each Hox gene contains a 4847: 4619:Hedgehog signaling pathway 4496:Developmental architecture 3850:Mansfield JH (June 2013). 3819:10.2174/138920212800793302 3801:Woltering JM (June 2012). 3217:The Ulster Medical Journal 3126:10.1016/j.cell.2008.09.002 3026:10.1016/j.cell.2007.05.022 2418:10.1093/genetics/126.1.139 2369:10.1093/genetics/129.2.443 1799:10.1038/s41467-021-23641-8 1164: 447:loss-of-function mutants, 427: 385: 129:. The homeodomain is a 60- 74:do not express Hox genes. 70:and the pilidium larva of 68:Schizocardium californicum 4796: 4446:Transgressive segregation 4093:10.1007/s00427-006-0119-y 4046:10.1186/s13227-018-0114-1 3711:"Hox genes and evolution" 3374:10.1186/s13059-018-1592-0 1639:10.1007/s00412-015-0543-8 1538:10.1186/s13227-020-0148-z 1493:10.1016/j.cub.2016.10.057 1123:whole genome duplications 669: 583: 565:Examples of target genes 4177:. Yale University Press. 1879:10.3389/fgene.2015.00267 1677:Nature Reviews. Genetics 1339:homeotic transformations 1252:formed. When the HOX10 1183:of the dorsal skin, the 1160:freshwater butterflyfish 613:(represses distal-less) 599:(represses distal-less) 4624:Notch signaling pathway 4599:Gene regulatory network 4482:Dual inheritance theory 4017:Terebratalia transversa 3877:10.1073/pnas.1307778110 3425:10.1186/1471-2164-8-317 3163:Genes & Development 2832:10.1126/science.1671176 2689:Genes & Development 2613:10.1242/dev.125.24.4949 2140:Genes & Development 1216:(A–D) on four separate 797:Polycomb-group proteins 748:Regulation of Hox genes 244:Drosophila melanogaster 237:Drosophila melanogaster 4672:cis-regulatory element 4580:Control of development 4460:Non-genetic influences 4426:evolutionary landscape 4265:. In Bittar EE (ed.). 3977:10.1073/pnas.97.9.4493 3779:10.1006/dbio.2000.9651 1589:10.1186/1741-7007-5-47 1307:Nematostella vectensis 1291:Caenorhabditis elegans 1270:Branchiostoma floridae 866: 561:, and cell migration. 420: 400:mutation, caused by a 268:), sex combs reduced ( 239: 176: 39:group of related genes 4783:Nature versus nurture 4687:Cell surface receptor 4604:Evo-devo gene toolkit 4503:Developmental biology 4441:Polygenic inheritance 4367:Quantitative genetics 4267:Developmental Biology 3927:10.1101/gr.073676.107 3766:Developmental Biology 3631:Developmental Biology 3559:10.1093/molbev/msu202 3519:10.1093/molbev/msn097 2738:Developmental Biology 2514:Developmental Biology 2289:"The Interactive Fly" 2202:10.1073/pnas.95.2.606 2113:10.1387/ijdb.180162sg 1866:Frontiers in Genetics 1778:Nature Communications 1310:, both in the phylum 864: 414: 402:chromosomal inversion 272:), and Antennapedia ( 234: 162: 4692:Transcription factor 4407:Genetic assimilation 4394:Genetic architecture 2153:10.1101/gad.10.2.176 1840:Karolinska Institute 808:chromosome territory 216:last common ancestor 83:transcription factor 4788:Morphogenetic field 4705:Influential figures 4202:1980Natur.287..795N 4173:Gehring WJ (1998). 4144:10.1038/nature01872 4136:2003Natur.424.1061L 4130:(6952): 1061–1065. 3868:2013PNAS..11010473M 3862:(26): 10473–10474. 3309:1994Natur.370..563G 3175:10.1101/gad.1631708 3079:10.1038/nature05916 3071:2007Natur.447..413F 2824:1991Sci...251..426H 2732:Gilbert SF (2000). 2467:2010PLoSO...510820H 2193:1998PNAS...95..606J 1988:2007PLoSO...2..153R 1930:1999Natur.399..772D 1790:2021NatCo..12.3354J 1735:1995Natur.376..479C 1519:Phoronopsis harmeri 1484:2017CBio...27..R21H 1405:, Switzerland) and 1403:University of Basel 1284:Other invertebrates 703:(activates Cdkn1a) 647:(activates reaper) 566: 302:intercalary segment 284:) and abdominal-B ( 104:Barrett's esophagus 4477:Genomic imprinting 3471:10.1038/ng0606-601 2979:10.1002/bies.20704 2912:Small S, Blair A, 2701:10.1101/gad.817100 2107:(11–12): 673–683. 1354:Thomas Hunt Morgan 1325:, and a suite of 1234:adaptive radiation 1151:genome duplication 867: 773:morphogenic fields 686:(activates EphA7) 564: 421: 415:Wild type (left), 378:embryo and adult. 260:), proboscipedia ( 240: 212:bilaterian animals 177: 4805: 4804: 4738:Eric F. Wieschaus 4700: 4699: 4518:Pattern formation 4422:Fitness landscape 4276:978-1-55938-816-0 4196:(5785): 795–801. 3553:(10): 2592–2611. 3303:(6490): 563–566. 3065:(7143): 413–417. 2928:(11): 4047–4057. 2916:(November 1992). 2880:978-0-12-374529-3 2695:(20): 2581–2586. 2607:(24): 4949–4957. 1924:(6738): 772–776. 1729:(6540): 479–485. 1376:Eric F. Wieschaus 1119: 1118: 708: 707: 366:Sex combs reduced 108:esophageal cancer 16:(Redirected from 4838: 4748:William McGinnis 4717:Richard Lewontin 4712:C. H. Waddington 4584: 4561:Neutral networks 4311: 4304: 4297: 4288: 4280: 4248: 4247: 4236: 4230: 4229: 4210:10.1038/287795a0 4185: 4179: 4178: 4170: 4164: 4163: 4119: 4113: 4112: 4075: 4069: 4068: 4058: 4048: 4038: 4006: 4000: 3999: 3989: 3979: 3970:(9): 4493–4498. 3955: 3949: 3948: 3938: 3921:(7): 1100–1111. 3906: 3900: 3899: 3889: 3879: 3847: 3841: 3840: 3830: 3807:Current Genomics 3798: 3792: 3791: 3781: 3757: 3751: 3750: 3740: 3730: 3706: 3700: 3699: 3671: 3665: 3664: 3654: 3622: 3616: 3615: 3587: 3581: 3580: 3570: 3538: 3532: 3531: 3521: 3512:(7): 1333–1343. 3497: 3491: 3490: 3454: 3448: 3447: 3437: 3427: 3403: 3397: 3396: 3386: 3376: 3352: 3346: 3343: 3337: 3336: 3317:10.1038/370563a0 3292: 3286: 3285: 3249: 3243: 3242: 3232: 3208: 3197: 3196: 3186: 3154: 3148: 3147: 3137: 3105: 3099: 3098: 3054: 3048: 3047: 3037: 3020:(7): 1311–1323. 3005: 2999: 2998: 2962: 2956: 2955: 2945: 2922:The EMBO Journal 2909: 2903: 2902: 2892: 2850: 2844: 2843: 2807: 2801: 2800: 2782: 2758: 2752: 2751: 2729: 2723: 2722: 2712: 2680: 2674: 2673: 2655: 2631: 2625: 2624: 2596: 2587: 2586: 2549: 2540: 2539: 2529: 2505: 2499: 2498: 2488: 2478: 2446: 2440: 2439: 2429: 2397: 2391: 2390: 2380: 2348: 2342: 2341: 2331: 2308:The EMBO Journal 2299: 2293: 2292: 2287:Brody T (1996). 2284: 2273: 2272: 2236: 2225: 2224: 2214: 2204: 2172: 2166: 2165: 2155: 2131: 2125: 2124: 2096: 2090: 2089: 2053: 2047: 2046: 2037:(7–8): 605–611. 2026: 2020: 2019: 2009: 1999: 1967: 1958: 1957: 1913: 1902: 1901: 1891: 1881: 1857: 1851: 1850: 1848: 1846: 1828: 1822: 1821: 1811: 1801: 1769: 1763: 1762: 1743:10.1038/376479a0 1715: 1709: 1708: 1672: 1661: 1660: 1650: 1618: 1612: 1611: 1601: 1591: 1567: 1561: 1560: 1550: 1540: 1512: 1506: 1505: 1495: 1463: 1407:Matthew P. Scott 1395:William McGinnis 1185:skeletal muscles 879: 857:In other species 653:decapentaplegic 619:decapentaplegic 567: 280:), abdominal-A ( 139:homeodomain fold 53:of animals. Hox 21: 4846: 4845: 4841: 4840: 4839: 4837: 4836: 4835: 4811: 4810: 4808: 4806: 4801: 4792: 4771: 4758:Sean B. Carroll 4696: 4628: 4575: 4539: 4491: 4472:Maternal effect 4455: 4388: 4325: 4315: 4277: 4261:Hunt P (1998). 4260: 4257: 4255:Further reading 4252: 4251: 4238: 4237: 4233: 4187: 4186: 4182: 4172: 4171: 4167: 4121: 4120: 4116: 4077: 4076: 4072: 4008: 4007: 4003: 3957: 3956: 3952: 3915:Genome Research 3908: 3907: 3903: 3849: 3848: 3844: 3800: 3799: 3795: 3759: 3758: 3754: 3708: 3707: 3703: 3673: 3672: 3668: 3624: 3623: 3619: 3589: 3588: 3584: 3540: 3539: 3535: 3499: 3498: 3494: 3459:Nature Genetics 3456: 3455: 3451: 3405: 3404: 3400: 3354: 3353: 3349: 3344: 3340: 3294: 3293: 3289: 3251: 3250: 3246: 3210: 3209: 3200: 3156: 3155: 3151: 3107: 3106: 3102: 3056: 3055: 3051: 3007: 3006: 3002: 2964: 2963: 2959: 2911: 2910: 2906: 2881: 2852: 2851: 2847: 2809: 2808: 2804: 2760: 2759: 2755: 2748: 2731: 2730: 2726: 2682: 2681: 2677: 2633: 2632: 2628: 2598: 2597: 2590: 2551: 2550: 2543: 2507: 2506: 2502: 2448: 2447: 2443: 2399: 2398: 2394: 2350: 2349: 2345: 2301: 2300: 2296: 2286: 2285: 2276: 2238: 2237: 2228: 2174: 2173: 2169: 2133: 2132: 2128: 2098: 2097: 2093: 2055: 2054: 2050: 2028: 2027: 2023: 1969: 1968: 1961: 1915: 1914: 1905: 1859: 1858: 1854: 1844: 1842: 1830: 1829: 1825: 1771: 1770: 1766: 1717: 1716: 1712: 1689:10.1038/nrg1726 1683:(12): 893–904. 1674: 1673: 1664: 1620: 1619: 1615: 1569: 1568: 1564: 1514: 1513: 1509: 1472:Current Biology 1465: 1464: 1460: 1455: 1428: 1419: 1380:D. melanogaster 1343:William Bateson 1335: 1286: 1263: 1169: 872: 859: 839: 828: 762:pair-rule genes 750: 713: 554: 552:Regulated genes 525: 505: 461: 432: 426: 390: 384: 368: 347: 337:interacts with 324: 294: 227: 155: 150: 116: 28: 23: 22: 15: 12: 11: 5: 4844: 4842: 4834: 4833: 4828: 4823: 4813: 4812: 4803: 4802: 4797: 4794: 4793: 4791: 4790: 4785: 4779: 4777: 4773: 4772: 4770: 4769: 4768: 4767: 4755: 4750: 4745: 4740: 4735: 4734: 4733: 4722:François Jacob 4719: 4714: 4708: 4706: 4702: 4701: 4698: 4697: 4695: 4694: 4689: 4684: 4679: 4674: 4669: 4664: 4659: 4658: 4657: 4647: 4642: 4636: 4634: 4630: 4629: 4627: 4626: 4621: 4616: 4611: 4606: 4601: 4596: 4590: 4588: 4581: 4577: 4576: 4574: 4573: 4568: 4563: 4558: 4553: 4547: 4545: 4541: 4540: 4538: 4537: 4532: 4527: 4522: 4521: 4520: 4515: 4505: 4499: 4497: 4493: 4492: 4490: 4489: 4484: 4479: 4474: 4469: 4463: 4461: 4457: 4456: 4454: 4453: 4451:Sequence space 4448: 4443: 4438: 4433: 4428: 4419: 4414: 4409: 4404: 4398: 4396: 4390: 4389: 4387: 4386: 4381: 4380: 4379: 4369: 4364: 4359: 4354: 4349: 4344: 4339: 4333: 4331: 4327: 4326: 4316: 4314: 4313: 4306: 4299: 4291: 4285: 4284: 4281: 4275: 4256: 4253: 4250: 4249: 4244:Nobelprize.org 4231: 4180: 4165: 4114: 4070: 4036:10.1101/449488 4001: 3950: 3901: 3842: 3813:(4): 289–299. 3793: 3772:(2): 333–342. 3752: 3701: 3682:(3): 200–203. 3666: 3617: 3582: 3533: 3492: 3465:(6): 601–602. 3449: 3398: 3361:Genome Biology 3347: 3338: 3287: 3260:(4): 551–553. 3244: 3198: 3169:(3): 346–359. 3149: 3120:(6): 921–931. 3100: 3049: 3000: 2973:(2): 110–121. 2957: 2904: 2879: 2845: 2802: 2773:(7): 1275–83. 2753: 2746: 2724: 2675: 2646:(4): 457–466. 2626: 2588: 2561:(3): 437–450. 2541: 2520:(2): 175–185. 2500: 2441: 2412:(1): 139–155. 2392: 2363:(2): 443–461. 2343: 2314:(3): 767–777. 2294: 2274: 2247:(6): 459–499. 2226: 2187:(2): 606–611. 2167: 2146:(2): 176–184. 2126: 2091: 2064:(2): 191–201. 2048: 2021: 1959: 1903: 1852: 1823: 1764: 1710: 1662: 1633:(3): 497–521. 1613: 1562: 1507: 1478:(1): R21–R24. 1457: 1456: 1454: 1451: 1450: 1449: 1444: 1439: 1434: 1427: 1424: 1418: 1415: 1401:'s lab at the 1399:Walter Gehring 1391:Michael Levine 1334: 1331: 1285: 1282: 1262: 1259: 1240:. All of the 1165:Main article: 1117: 1116: 1077: 1072: 1064: 1063: 1024: 1019: 1011: 1010: 967: 962: 954: 953: 906: 901: 893: 892: 889: 883: 871: 868: 858: 855: 838: 835: 827: 824: 789:Non-coding RNA 749: 746: 712: 709: 706: 705: 699: 693: 689: 688: 682: 676: 673: 667: 666: 660: 654: 650: 649: 643: 637: 633: 632: 628:ULTRABITHORAX 626: 620: 616: 615: 609: 606: 602: 601: 597:ULTRABITHORAX 595: 592: 589: 581: 580: 577: 574: 571: 553: 550: 524: 521: 504: 501: 460: 457: 428:Main article: 425: 422: 419:mutant (right) 386:Main article: 383: 380: 367: 364: 346: 343: 323: 320: 293: 290: 226: 220: 154: 151: 147: 115: 112: 87:well-conserved 51:head-tail axis 35:homeobox genes 33:, a subset of 27:Group of genes 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 4843: 4832: 4829: 4827: 4826:Gene clusters 4824: 4822: 4819: 4818: 4816: 4809: 4800: 4795: 4789: 4786: 4784: 4781: 4780: 4778: 4774: 4766: 4765: 4761: 4760: 4759: 4756: 4754: 4751: 4749: 4746: 4744: 4741: 4739: 4736: 4732: 4729: 4728: 4727: 4726:Jacques Monod 4723: 4720: 4718: 4715: 4713: 4710: 4709: 4707: 4703: 4693: 4690: 4688: 4685: 4683: 4680: 4678: 4675: 4673: 4670: 4668: 4665: 4663: 4660: 4656: 4653: 4652: 4651: 4648: 4646: 4643: 4641: 4640:Homeotic gene 4638: 4637: 4635: 4631: 4625: 4622: 4620: 4617: 4615: 4612: 4610: 4607: 4605: 4602: 4600: 4597: 4595: 4592: 4591: 4589: 4585: 4582: 4578: 4572: 4569: 4567: 4564: 4562: 4559: 4557: 4554: 4552: 4549: 4548: 4546: 4542: 4536: 4533: 4531: 4528: 4526: 4523: 4519: 4516: 4514: 4511: 4510: 4509: 4508:Morphogenesis 4506: 4504: 4501: 4500: 4498: 4494: 4488: 4485: 4483: 4480: 4478: 4475: 4473: 4470: 4468: 4465: 4464: 4462: 4458: 4452: 4449: 4447: 4444: 4442: 4439: 4437: 4434: 4432: 4429: 4427: 4423: 4420: 4418: 4415: 4413: 4410: 4408: 4405: 4403: 4400: 4399: 4397: 4395: 4391: 4385: 4382: 4378: 4375: 4374: 4373: 4370: 4368: 4365: 4363: 4360: 4358: 4355: 4353: 4350: 4348: 4345: 4343: 4342:Reaction norm 4340: 4338: 4335: 4334: 4332: 4328: 4324: 4320: 4312: 4307: 4305: 4300: 4298: 4293: 4292: 4289: 4282: 4278: 4272: 4268: 4264: 4259: 4258: 4254: 4245: 4241: 4235: 4232: 4227: 4223: 4219: 4215: 4211: 4207: 4203: 4199: 4195: 4191: 4184: 4181: 4176: 4169: 4166: 4161: 4157: 4153: 4149: 4145: 4141: 4137: 4133: 4129: 4125: 4118: 4115: 4110: 4106: 4102: 4098: 4094: 4090: 4086: 4082: 4074: 4071: 4066: 4062: 4057: 4052: 4047: 4042: 4037: 4032: 4028: 4024: 4020: 4018: 4014: 4005: 4002: 3997: 3993: 3988: 3983: 3978: 3973: 3969: 3965: 3961: 3954: 3951: 3946: 3942: 3937: 3932: 3928: 3924: 3920: 3916: 3912: 3905: 3902: 3897: 3893: 3888: 3883: 3878: 3873: 3869: 3865: 3861: 3857: 3853: 3846: 3843: 3838: 3834: 3829: 3824: 3820: 3816: 3812: 3808: 3804: 3797: 3794: 3789: 3785: 3780: 3775: 3771: 3767: 3763: 3756: 3753: 3748: 3744: 3739: 3734: 3729: 3724: 3720: 3716: 3715:F1000Research 3712: 3705: 3702: 3697: 3693: 3689: 3685: 3681: 3677: 3670: 3667: 3662: 3658: 3653: 3648: 3644: 3640: 3636: 3632: 3628: 3621: 3618: 3613: 3609: 3605: 3601: 3597: 3593: 3586: 3583: 3578: 3574: 3569: 3564: 3560: 3556: 3552: 3548: 3544: 3537: 3534: 3529: 3525: 3520: 3515: 3511: 3507: 3503: 3496: 3493: 3488: 3484: 3480: 3476: 3472: 3468: 3464: 3460: 3453: 3450: 3445: 3441: 3436: 3431: 3426: 3421: 3417: 3413: 3409: 3402: 3399: 3394: 3390: 3385: 3380: 3375: 3370: 3366: 3362: 3358: 3351: 3348: 3342: 3339: 3334: 3330: 3326: 3322: 3318: 3314: 3310: 3306: 3302: 3298: 3291: 3288: 3283: 3279: 3275: 3271: 3267: 3263: 3259: 3255: 3248: 3245: 3240: 3236: 3231: 3226: 3222: 3218: 3214: 3207: 3205: 3203: 3199: 3194: 3190: 3185: 3180: 3176: 3172: 3168: 3164: 3160: 3153: 3150: 3145: 3141: 3136: 3131: 3127: 3123: 3119: 3115: 3111: 3104: 3101: 3096: 3092: 3088: 3084: 3080: 3076: 3072: 3068: 3064: 3060: 3053: 3050: 3045: 3041: 3036: 3031: 3027: 3023: 3019: 3015: 3011: 3004: 3001: 2996: 2992: 2988: 2984: 2980: 2976: 2972: 2968: 2961: 2958: 2953: 2949: 2944: 2939: 2935: 2931: 2927: 2923: 2919: 2915: 2908: 2905: 2900: 2896: 2891: 2886: 2882: 2876: 2872: 2868: 2864: 2860: 2856: 2849: 2846: 2841: 2837: 2833: 2829: 2825: 2821: 2817: 2813: 2806: 2803: 2798: 2794: 2790: 2786: 2781: 2776: 2772: 2768: 2764: 2757: 2754: 2749: 2747:0-87893-243-7 2743: 2739: 2735: 2728: 2725: 2720: 2716: 2711: 2706: 2702: 2698: 2694: 2690: 2686: 2679: 2676: 2671: 2667: 2663: 2659: 2654: 2649: 2645: 2641: 2637: 2630: 2627: 2622: 2618: 2614: 2610: 2606: 2602: 2595: 2593: 2589: 2584: 2580: 2576: 2572: 2568: 2564: 2560: 2556: 2548: 2546: 2542: 2537: 2533: 2528: 2523: 2519: 2515: 2511: 2504: 2501: 2496: 2492: 2487: 2482: 2477: 2472: 2468: 2464: 2461:(5): e10820. 2460: 2456: 2452: 2445: 2442: 2437: 2433: 2428: 2423: 2419: 2415: 2411: 2407: 2403: 2396: 2393: 2388: 2384: 2379: 2374: 2370: 2366: 2362: 2358: 2354: 2347: 2344: 2339: 2335: 2330: 2325: 2321: 2317: 2313: 2309: 2305: 2298: 2295: 2290: 2283: 2281: 2279: 2275: 2270: 2266: 2262: 2258: 2254: 2250: 2246: 2242: 2235: 2233: 2231: 2227: 2222: 2218: 2213: 2208: 2203: 2198: 2194: 2190: 2186: 2182: 2178: 2171: 2168: 2163: 2159: 2154: 2149: 2145: 2141: 2137: 2130: 2127: 2122: 2118: 2114: 2110: 2106: 2102: 2095: 2092: 2087: 2083: 2079: 2075: 2071: 2067: 2063: 2059: 2052: 2049: 2044: 2040: 2036: 2032: 2025: 2022: 2017: 2013: 2008: 2003: 1998: 1993: 1989: 1985: 1981: 1977: 1973: 1966: 1964: 1960: 1955: 1951: 1947: 1943: 1939: 1938:10.1038/21631 1935: 1931: 1927: 1923: 1919: 1912: 1910: 1908: 1904: 1899: 1895: 1890: 1885: 1880: 1875: 1871: 1867: 1863: 1856: 1853: 1841: 1837: 1833: 1827: 1824: 1819: 1815: 1810: 1805: 1800: 1795: 1791: 1787: 1783: 1779: 1775: 1768: 1765: 1760: 1756: 1752: 1748: 1744: 1740: 1736: 1732: 1728: 1724: 1720: 1714: 1711: 1706: 1702: 1698: 1694: 1690: 1686: 1682: 1678: 1671: 1669: 1667: 1663: 1658: 1654: 1649: 1644: 1640: 1636: 1632: 1628: 1624: 1617: 1614: 1609: 1605: 1600: 1595: 1590: 1585: 1581: 1577: 1573: 1566: 1563: 1558: 1554: 1549: 1544: 1539: 1534: 1530: 1526: 1522: 1520: 1511: 1508: 1503: 1499: 1494: 1489: 1485: 1481: 1477: 1473: 1469: 1462: 1459: 1452: 1448: 1445: 1443: 1442:Morphogenesis 1440: 1438: 1435: 1433: 1432:Homeotic gene 1430: 1429: 1425: 1423: 1416: 1414: 1412: 1408: 1404: 1400: 1396: 1392: 1387: 1385: 1381: 1377: 1373: 1369: 1365: 1361: 1359: 1355: 1350: 1348: 1344: 1340: 1332: 1330: 1328: 1324: 1320: 1315: 1313: 1309: 1308: 1303: 1302: 1297: 1293: 1292: 1283: 1281: 1279: 1275: 1271: 1267: 1260: 1258: 1255: 1249: 1247: 1243: 1239: 1235: 1231: 1227: 1223: 1219: 1215: 1211: 1207: 1203: 1199: 1193: 1191: 1190:invertebrates 1186: 1182: 1178: 1173: 1168: 1163: 1161: 1157: 1152: 1148: 1144: 1140: 1136: 1132: 1128: 1124: 1115: 1114: 1110: 1106: 1102: 1098: 1094: 1090: 1086: 1082: 1078: 1076: 1073: 1071: 1070: 1066: 1065: 1062: 1061: 1057: 1053: 1049: 1045: 1041: 1037: 1033: 1029: 1025: 1023: 1022:chromosome 12 1020: 1018: 1017: 1013: 1012: 1009: 1008: 1004: 1000: 996: 992: 988: 984: 980: 976: 972: 968: 966: 965:chromosome 17 963: 961: 960: 956: 955: 952: 951: 947: 943: 939: 935: 931: 927: 923: 919: 915: 911: 907: 905: 902: 900: 899: 895: 894: 890: 888: 884: 881: 880: 877: 874: 869: 863: 856: 854: 852: 847: 841: 836: 834: 830: 825: 823: 819: 816: 815:retinoic acid 811: 809: 805: 800: 798: 794: 790: 786: 784: 780: 778: 774: 769: 767: 763: 759: 755: 747: 745: 743: 742:Extradenticle 737: 735: 731: 727: 723: 718: 710: 704: 700: 698: 694: 691: 690: 687: 683: 681: 677: 674: 672: 668: 665: 661: 659: 655: 652: 651: 648: 644: 642: 638: 635: 634: 631: 627: 625: 621: 618: 617: 614: 610: 607: 604: 603: 600: 596: 593: 590: 588: 587: 582: 579:Regulated by 578: 575: 572: 569: 568: 562: 560: 551: 549: 547: 543: 539: 535: 531: 522: 520: 518: 514: 510: 502: 500: 498: 494: 490: 486: 482: 478: 474: 470: 466: 458: 456: 454: 450: 446: 442: 437: 431: 430:Ultrabithorax 424:Ultrabithorax 423: 418: 413: 409: 407: 403: 399: 395: 389: 381: 379: 377: 373: 365: 363: 360: 356: 352: 344: 342: 340: 336: 332: 330: 322:Proboscipedia 321: 319: 316: 311: 307: 303: 299: 291: 289: 287: 283: 279: 275: 271: 267: 264:), deformed ( 263: 259: 255: 251: 246: 245: 238: 233: 229: 225: 221: 219: 217: 213: 208: 206: 202: 198: 194: 190: 186: 182: 174: 170: 166: 161: 157: 152: 146: 144: 140: 136: 132: 128: 123: 118: 113: 111: 109: 105: 101: 96: 91: 88: 84: 79: 75: 73: 69: 65: 59: 56: 52: 48: 44: 40: 36: 32: 19: 4807: 4762: 4655:eyeless gene 4644: 4551:Evolvability 4525:Segmentation 4402:Canalisation 4372:Heterochrony 4362:Heritability 4330:Key concepts 4269:. Elsevier. 4266: 4243: 4234: 4193: 4189: 4183: 4174: 4168: 4127: 4123: 4117: 4087:(1): 39–54. 4084: 4080: 4073: 4026: 4022: 4016: 4012: 4004: 3967: 3963: 3953: 3918: 3914: 3904: 3859: 3855: 3845: 3810: 3806: 3796: 3769: 3765: 3755: 3718: 3714: 3704: 3679: 3675: 3669: 3634: 3630: 3620: 3598:(1): 72–83. 3595: 3591: 3585: 3550: 3546: 3536: 3509: 3505: 3495: 3462: 3458: 3452: 3415: 3412:BMC Genomics 3411: 3401: 3364: 3360: 3350: 3341: 3300: 3296: 3290: 3257: 3253: 3247: 3223:(1): 23–31. 3220: 3216: 3166: 3162: 3152: 3117: 3113: 3103: 3062: 3058: 3052: 3017: 3013: 3003: 2970: 2966: 2960: 2925: 2921: 2907: 2862: 2858: 2848: 2815: 2811: 2805: 2770: 2766: 2756: 2737: 2727: 2692: 2688: 2678: 2643: 2639: 2629: 2604: 2600: 2558: 2554: 2517: 2513: 2503: 2458: 2454: 2444: 2409: 2405: 2395: 2360: 2356: 2346: 2311: 2307: 2297: 2244: 2240: 2184: 2180: 2170: 2143: 2139: 2129: 2104: 2100: 2094: 2061: 2057: 2051: 2034: 2030: 2024: 1979: 1975: 1921: 1917: 1869: 1865: 1855: 1843:. Retrieved 1835: 1826: 1781: 1777: 1767: 1726: 1722: 1713: 1680: 1676: 1630: 1626: 1616: 1579: 1575: 1565: 1528: 1524: 1518: 1510: 1475: 1471: 1461: 1420: 1388: 1379: 1366: 1362: 1357: 1351: 1336: 1316: 1305: 1299: 1289: 1287: 1277: 1273: 1269: 1264: 1250: 1194: 1170: 1141:, including 1139:teleost fish 1120: 1079: 1075:chromosome 2 1067: 1026: 1014: 969: 957: 908: 904:chromosome 7 896: 875: 873: 850: 845: 842: 840: 837:Nomenclature 831: 829: 820: 812: 801: 787: 781: 776: 770: 751: 738: 714: 702: 696: 685: 679: 670: 663: 662:ABDOMINAL-B 657: 646: 640: 629: 623: 612: 611:ABDOMINAL-A 605:distal-less 598: 591:distal-less 585: 584: 573:Target gene 555: 545: 541: 537: 533: 529: 526: 516: 512: 508: 506: 488: 484: 480: 476: 472: 468: 464: 462: 452: 448: 444: 440: 435: 433: 417:Antennapedia 405: 397: 393: 391: 388:Antennapedia 382:Antennapedia 375: 371: 369: 358: 354: 350: 348: 338: 334: 328: 325: 314: 309: 305: 297: 295: 285: 281: 277: 273: 269: 265: 261: 257: 253: 249: 242: 241: 236: 228: 223: 209: 201:sea anemones 178: 169:heterochrony 156: 119: 117: 80: 76: 67: 64:hemichordate 60: 30: 29: 4753:Mike Levine 4662:Distal-less 4487:Polyphenism 4467:Epigenetics 4319:development 3637:(1): 7–15. 2601:Development 1982:(1): e153. 1784:(1): 3354. 1576:BMC Biology 1319:brachiopods 1218:chromosomes 1214:gene arrays 870:Vertebrates 826:Colinearity 503:Abdominal-B 459:Abdominal-A 205:duplication 143:polypeptide 127:homeodomain 4815:Categories 4731:Lac operon 4556:Robustness 4535:Modularity 4530:Metamerism 4436:Plasticity 4431:Pleiotropy 4384:Heterotopy 3418:(1): 317. 3367:(1): 209. 2865:: 63–101. 1719:Carroll SB 1627:Chromosoma 1453:References 1278:AmphiHox15 1246:phenotypic 1242:paralogous 1206:homologous 1198:drosophila 1172:Vertebrate 887:Chromosome 851:Drosophila 846:Drosophila 754:Drosophila 658:positions 586:Drosophila 517:Drosophila 489:abd-A gene 465:Drosophila 376:Drosophila 310:Drosophila 250:Drosophila 224:Drosophila 181:eukaryotes 173:trilobites 131:amino-acid 100:metaplasia 95:chromosome 49:along the 4682:Morphogen 4667:Engrailed 4650:Pax genes 4571:Tinkering 4417:Epistasis 4412:Dominance 4323:phenotype 3592:BioEssays 2967:BioEssays 1836:csb.ki.se 1386:in 1995. 1296:roundworm 1274:AmphiHox1 1266:Amphioxus 1261:Amphioxus 1226:mutations 1143:zebrafish 1135:tetrapods 804:chromatin 758:gap genes 730:glutamine 645:DEFORMED 570:Organism 559:apoptosis 534:Antp, Ubx 404:, causes 193:bilateria 185:evolution 165:arthropod 153:Evolution 135:base-pair 122:enhancers 31:Hox genes 18:Hox genes 4645:Hox gene 4633:Elements 4614:Homeobox 4152:12944969 4101:17180685 4065:30637095 3996:10781050 3945:18562680 3896:23749870 3837:23204918 3788:10753520 3747:27239281 3661:20435029 3612:10649293 3577:24974377 3528:18424774 3487:41211603 3479:16736008 3444:17845724 3393:30486862 3282:13370372 3239:16457401 3193:18245448 3144:18805086 3087:17522674 3044:17604720 2987:18200528 2914:Levine M 2899:19651302 2797:22478337 2719:11040212 2670:17464919 2662:12202035 2583:10977579 2536:24055174 2495:20520839 2455:PLOS ONE 2406:Genetics 2357:Genetics 2338:16453752 2269:46085797 2261:12492146 2121:30604837 2086:39168197 2043:14756336 2016:17252055 1976:PLOS ONE 1946:10391241 1898:26347770 1845:March 9, 1818:34099670 1697:16341070 1657:26464018 1608:17963489 1557:32064072 1531:(2): 2. 1502:28073016 1426:See also 1417:Research 1368:Ed Lewis 1327:molluscs 1323:annelids 1312:Cnidaria 1268:such as 1254:paralogs 1222:Clusters 1127:lampreys 799:(PRC2). 783:MicroRNA 701:Hox-A10 684:HOX-A13 497:mesoderm 493:ectoderm 345:Deformed 210:In most 199:such as 197:Cnidaria 114:Function 72:Nemertea 66:species 55:proteins 37:, are a 4776:Debates 4587:Systems 4513:Eyespot 4377:Neoteny 4226:4337658 4218:6776413 4198:Bibcode 4160:4317828 4132:Bibcode 4109:7314266 4056:6325747 4031:bioRxiv 4023:EvoDevo 3936:2493399 3887:3696806 3864:Bibcode 3828:3394116 3738:4863668 3721:: 859. 3652:2909379 3568:4166920 3435:2080641 3384:6260733 3333:4329696 3325:7914353 3305:Bibcode 3274:1358459 3230:1891803 3184:2216694 3135:2632951 3095:4388060 3067:Bibcode 3035:2084369 2952:1327756 2890:2810641 2840:1671176 2820:Bibcode 2812:Science 2789:2500253 2621:9811579 2575:1358457 2486:2876039 2463:Bibcode 2436:1977655 2427:1204118 2387:1683847 2378:1204635 2221:9435239 2189:Bibcode 2162:8566751 2078:7913880 2007:1779807 1984:Bibcode 1954:4413694 1926:Bibcode 1889:4539518 1872:: 267. 1809:8184780 1786:Bibcode 1759:4230019 1751:7637779 1731:Bibcode 1648:4901127 1599:2211742 1548:7011278 1525:EvoDevo 1480:Bibcode 1333:History 1230:haltere 1177:somites 1131:hagfish 882:Cluster 734:adenine 726:guanine 692:Cdkn1a 636:reaper 189:Metazoa 4677:Ligand 4357:Operon 4273: 4224: 4216: 4190:Nature 4158: 4150: 4124:Nature 4107: 4099: 4063: 4053: 4033: 3994: 3984: 3943: 3933: 3894: 3884: 3835: 3825: 3786: 3745: 3735: 3694: 3659: 3649: 3610: 3575: 3565: 3526: 3485: 3477: 3442: 3432: 3391: 3381: 3331: 3323: 3297:Nature 3280: 3272: 3237: 3227: 3191: 3181: 3142: 3132: 3093: 3085: 3059:Nature 3042: 3032: 2993: 2985: 2950: 2943:556915 2940: 2897: 2887: 2877: 2838: 2795: 2787: 2744: 2717: 2710:317001 2707: 2668: 2660: 2619: 2581: 2573: 2534: 2493: 2483: 2434: 2424: 2385: 2375: 2336: 2329:553462 2326: 2267: 2259: 2219: 2209: 2160: 2119: 2084: 2076: 2041: 2014: 2004: 1952: 1944: 1918:Nature 1896: 1886: 1816: 1806: 1757: 1749: 1723:Nature 1705:256216 1703: 1695: 1655: 1645: 1606: 1596: 1582:: 47. 1555: 1545: 1500: 1393:, and 1347:Mendel 1202:genome 1181:dermis 1156:salmon 1147:medaka 1113:HOXD13 1109:HOXD12 1105:HOXD11 1101:HOXD10 1060:HOXC13 1056:HOXC12 1052:HOXC11 1048:HOXC10 1007:HOXB13 950:HOXA13 946:HOXA11 942:HOXA10 891:Genes 885:Human 793:HOTAIR 722:lysine 675:EphA7 530:Hox6-8 292:Labial 47:embryo 45:of an 4283:Fatim 4222:S2CID 4156:S2CID 4105:S2CID 4029:: 1. 3987:18262 3483:S2CID 3329:S2CID 3278:S2CID 3091:S2CID 2995:53138 2991:S2CID 2793:S2CID 2666:S2CID 2579:S2CID 2265:S2CID 2212:18467 2082:S2CID 1950:S2CID 1755:S2CID 1701:S2CID 1301:Hydra 1238:phyla 1210:genes 1097:HOXD9 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