The origin and evolution of segmentation.

Arthropods, annelids and chordates all possess segments. It remains unclear, however, whether the segments of these animals evolved independently or instead were derived from a common ancestor. Considering this question involves examining not only the similarities and differences in the process of segmentation between these phyla, but also how this process varies within phyla, where the homology of segments is generally accepted. This article reviews what is known about the segmentation process and considers various proposals to explain its evolution.

[1]  J. Rossant,et al.  Notch1 is required for the coordinate segmentation of somites. , 1995, Development.

[2]  J. Ahringer,et al.  Posterior patterning by the Caenorhabditis elegans even-skipped homolog vab-7. , 1996, Genes & development.

[3]  D. Tautz,et al.  A conserved mode of head segmentation in arthropods revealed by the expression pattern of Hox genes in a spider. , 1998, Proceedings of the National Academy of Sciences of the United States of America.

[4]  D. Weisblat,et al.  Segmental expression of an engrailed-class gene during early development and neurogenesis in an annelid. , 1991, Development.

[5]  E. D. De Robertis Evolutionary biology. The ancestry of segmentation. , 1997, Nature.

[6]  R. W. Padgett,et al.  TGF-beta signaling, Smads, and tumor suppressors. , 1998, BioEssays : news and reviews in molecular, cellular and developmental biology.

[7]  Susan J. Brown,et al.  The beetle Tribolium castaneum has a fushi tarazu homolog expressed in stripes during segmentation. , 1994, Proceedings of the National Academy of Sciences of the United States of America.

[8]  K. G. Coleman,et al.  Expression of engrailed proteins in arthropods, annelids, and chordates , 1989, Cell.

[9]  N. Patel The evolution of arthropod segmentation: insights from comparisons of gene expression patterns. , 1994, Development (Cambridge, England). Supplement.

[10]  D. Melton,et al.  Bimodal and graded expression of the Xenopus homeobox gene Xhox3 during embryonic development. , 1989, Development.

[11]  D. Hirsh,et al.  Number and organization of collagen genes in Caenorhabditis elegans , 1984, Molecular and cellular biology.

[12]  N. Williams,et al.  Sequence and embryonic expression of the amphioxus engrailed gene (AmphiEn): the metameric pattern of transcription resembles that of its segment-polarity homolog in Drosophila. , 1997, Development.

[13]  M. Biggin,et al.  Eve and ftz regulate a wide array of genes in blastoderm embryos: the selector homeoproteins directly or indirectly regulate most genes in Drosophila. , 1998, Development.

[14]  R. Pennell Cell walls: structures and signals. , 1998, Current opinion in plant biology.

[15]  O. Pourquié,et al.  Avian hairy Gene Expression Identifies a Molecular Clock Linked to Vertebrate Segmentation and Somitogenesis , 1997, Cell.

[16]  Nipam H. Patel,et al.  Pair-rule expression patterns of even-skipped are found in both short- and long-germ beetles , 1994, Nature.

[17]  K. Kanyuka,et al.  A higher plant seven-transmembrane receptor that influences sensitivity to cytokinins , 2001, Current Biology.

[18]  M. D'Esposito,et al.  EVX2, a human homeobox gene homologous to the even-skipped segmentation gene, is localized at the 5' end of HOX4 locus on chromosome 2. , 1991, Genomics.

[19]  D. Weisblat,et al.  Cell lineage analysis of the expression of an engrailed homolog in leech embryos. , 1993, Development.

[20]  Gregor Bucher,et al.  Pair-rule and gap gene mutants in the flour beetle Tribolium castaneum , 1998, Development Genes and Evolution.

[21]  M. H. Angelis,et al.  Maintenance of somite borders in mice requires the Delta homologue Dll1 , 1997, Nature.

[22]  R. B. Clark Locomotion and the phylogeny of the Metazoa , 1981 .

[23]  Susan J. Brown,et al.  A deficiency of the homeotic complex of the beetle Tribolium , 1991, Nature.

[24]  K. Sander Specification of the Basic Body Pattern in Insect Embryogenesis1 , 1976 .

[25]  Nipam H. Patel,et al.  Changing role of even-skipped during the evolution of insect pattern formation , 1992, Nature.

[26]  C. Kimmel,et al.  Was Urbilateria segmented? , 1996, Trends in genetics : TIG.

[27]  N. Lartillot,et al.  Animal evolution. The end of the intermediate taxa? , 1999, Trends in genetics : TIG.

[28]  R. Raff,et al.  Evidence for a clade of nematodes, arthropods and other moulting animals , 1997, Nature.

[29]  Susan J. Brown,et al.  Molecular characterization and embryonic expression of the even-skipped ortholog of Tribolium castaneum , 1997, Mechanisms of Development.

[30]  M. Shankland Leech segmentation: cell lineage and the formation of complex body patterns. , 1991, Developmental biology.

[31]  D. Weisblat,et al.  Gangliogenesis in leech: morphogenetic processes leading to segmentation in the central nervous system , 1998, Development Genes and Evolution.

[32]  J. Campos-Ortega,et al.  Expression domains of a zebrafish homologue of the Drosophila pair-rule gene hairy correspond to primordia of alternating somites. , 1996, Development.

[33]  K. Irie,et al.  Isolation of ATMEKK1 (a MAP kinase kinase kinase)-interacting proteins and analysis of a MAP kinase cascade in Arabidopsis. , 1998, Biochemical and biophysical research communications.

[34]  J. Kramer Genetic analysis of extracellular matrix in C. elegans. , 1994, Annual review of genetics.

[35]  E. Robertis,et al.  The ancestry of segmentation , 1997, nature.

[36]  P. Gruss,et al.  A murine even‐skipped homologue, Evx 1, is expressed during early embryogenesis and neurogenesis in a biphasic manner. , 1990, The EMBO journal.

[37]  C. Nüsslein-Volhard,et al.  Zebrafish segmentation and pair-rule patterning. , 1998, Developmental genetics.

[38]  F. Falciani,et al.  Dax, a locust Hox gene related to fushi-tarazu but showing no pair-rule expression. , 1994, Development.

[39]  D. Eernisse,et al.  Annelida and Arthropoda are Not Sister Taxa: A Phylogenetic Analysis of Spiralian Metazoan Morphology , 1992 .

[40]  M. Goulding,et al.  Regulation of Pax-3 expression in the dermomyotome and its role in muscle development. , 1994, Development.

[41]  Ralf J. Sommer,et al.  Involvement of an orthologue of the Drosophila pair-rule gene hairy in segment formation of the short germ-band embryo of Tribolium (Coleoptera) , 1993, Nature.

[42]  T. Gridley,et al.  Defects in somite formation in lunatic fringe-deficient mice , 1998, Nature.

[43]  Sean B. Carroll,et al.  Hox genes in brachiopods and priapulids and protostome evolution , 1999, Nature.

[44]  D. Weisblat,et al.  Segmentation in leech development. , 1988, Development.