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Bat wing development

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is expressed in bat interdigit tissue during a time when apoptosis occurs which does not occur in mice. Thus, FGFs may play a role in blocking the apoptotic effects of BMPs in the bat wing interdigit. Finally, applying ectopic BMPs and FGF antagonists to developing bat wings results in apoptosis of
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to the skeletal components found in other tetrapod forelimbs. Through adaptive evolution these structures in bats have undergone many morphological changes, such as webbed digits, elongation of the forelimb, and reduction in bone thickness. Recently, there have been comparative studies of mouse and
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Although many of the molecular mechanisms involved in limb development are conserved between mouse and bat, there are a number of differences primarily seen in gene expression patterns. Surprisingly, the coding regions of many of these genes with different expression domains are highly conserved
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is a structure found at the distal most tip which becomes a key signaling center for the developing limb. Surprisingly many of the same signaling pathways known to play a role in tetrapod limb development have been found to play a role in bat forelimb development but the timing, intensity, and
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One major difference in bat forearms is that their skeletal limb structures are elongated. This elongation of the forelimb skeleton is required to support the wing membrane. Comparative in situ hybridization studies have revealed that the expression domain of
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genes have changed. Since mice are also mammals, it is convenient to compare morphology and development of forelimbs between mice and bats; these comparisons may elucidate the genetic basis of adaptive bat wing development.
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may also explain reduced size and density of the ulna found in bats. Overall, these studies suggest that the molecular changes responsible for the evolution of wings in bats is due to genetic regulatory changes.
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Another major difference in bat forelimbs is in the density of their skeletal limbs. The bones found in their forelimbs are reduced to achieve a light body weight required for flight. In particular, their
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occurs to separate the digits. BMP signals are most likely responsible for the interdigit apoptosis as they are expressed in the interdigit tissue and blocking BMP signaling will prevent interdigital
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enhancer with the endogenous enhancer found in mice, these transgenic mice had slightly increased forelimbs. Comparative studies have established that bat digits undergo a more rapid rate of
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is reduced in width and fused to the other zeugopod element, the radius. One of the possible molecular pathways involved in reduction of bat skeletal forelimb thickness is differences in
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Simplified diagram showing expanded gene expression domains in developing bat forelimb potentially contributing to the morphological changes resulting in the bat wing.
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shows increased expression in the digits of bats compared with mice, suggesting that a change in the BMP pathway has occurred to give rise to longer bat digits.
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Chen CH, Cretekos CJ, Rasweiler JJ, Behringer RR (2005). "Hoxd13 expression in the developing limbs of the short-tailed fruit bat, Carollia perspicillata".
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may contribute to the larger size of the bat forelimb. Because the mouse and bat orthologs are conserved, there is likely to be a regulatory change in
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expression domain in bat limbs has been shifted posteriorly in comparison to mouse. This observed difference in the expression pattern of
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proliferation. In addition to interdigit apoptosis, BMPs have been shown to affect chondrocyte proliferation and digit length in mice.
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bat forelimb development to understand the genetic basis of morphological evolution. Consequently, the bat wing is a valuable
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Saunders JW; Benthe, HF; Haberland, G (1972). "Developmental control of three-dimensional polarity in the avian limb".
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in bat forelimb AER are expanded in comparison to the mouse forelimb, suggesting that expanded expression of
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Zou H, Niswander L (1996). "Requirement for BMP Signaling in Interdigital Apoptosis and Scale Formation".
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Eswarakumar V, Lax I, Schlessinger J (2005). "Cellular signaling by fibroblast growth factor receptors".
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between mouse and bat. Thus, it is likely that this major morphological transition was a consequence of
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has an expanded expression domain and is upregulated. Researchers found that the coding region of
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changes. Researchers can study the genetic basis of bat wing development by using comparative
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null mutations lose their ulna structure. Another good candidate for bat bone reduction is
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De Luca F, Barnes KM, Uyeda JA, De-Levi S, Abad V, Palese T, Mericq V, Baron J (2001).
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to examine gene expression domains and using experimental embryology in mice and bats.
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Chiang C, Litingtung Y, Harris MP, Simandl BK, Li Y, Beachy PA, Fallon JF (2001).
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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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Cretekos CJ, Deng JM, Green ED, Rasweiler JJ, Behringer RR (2007).
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Sears KE (2008). "Molecular determinants of bat wing development".
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Weatherbee SD, Behringer RR, Rasweiler JJ, Niswander L (2006).
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model for studying the evolution of vertebrate limb diversity.
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in bats is nearly identical to mice but found a bat-specific
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Sears KE, Behringer RR, Rasweiler JJ, Niswander LA (2006).
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family. In situ hybridization studies have found that the
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(2008). 36:structures in their wings are morphologically 2144: 849: 8: 2151: 2137: 2129: 856: 842: 834: 326:Annals of the New York Academy of Sciences 69:involves many signaling molecules such as 2507:Tradeoffs for locomotion in air and water 771: 730: 689: 679: 627: 578: 537: 496: 486: 2090:Transgenerational epigenetic inheritance 649: 647: 276: 274: 204: 123:Formation of the bat wing membrane (the 47: 561:Martin JF, Bradley A, Olson EN (1995). 270: 1853:Dialogues Concerning Natural Religion 61:Comparisons to mouse limb development 7: 428:Cytokine & Growth Factor Reviews 56:skeletal structures of bat and mouse 1258:Evolutionary developmental biology 346:10.1111/j.1749-6632.1972.tb27821.x 173:in bats differs from mice in that 14: 2543:Vertebrate developmental biology 2247: 2112: 2103: 2102: 809:10.1111/j.1525-142X.2005.05015.x 1915:Extended evolutionary synthesis 1104:Gene-centered view of evolution 2043:Hologenome theory of evolution 1910:History of molecular evolution 1136:Evolutionarily stable strategy 1025:Last universal common ancestor 188:. When they replaced the bat 1: 1837:Renaissance and Enlightenment 440:10.1016/j.cytogfr.2005.01.001 28:forelimbs. Because bats are 2048:Missing heritability problem 1675:Gamete differentiation/sexes 397:10.1126/science.272.5262.738 797:Evolution & Development 214:Reduction in bone thickness 2559: 2363:Flying and gliding animals 2199:Fin and flipper locomotion 1680:Life cycles/nuclear phases 1232:Trivers–Willard hypothesis 237:, a gene belonging to the 24:. Bat wings are modified 2245: 2098: 1178:Parent–offspring conflict 983:Earliest known life forms 871: 119:Presence of webbed digits 2031:Cultural group selection 1895:The eclipse of Darwinism 1867:On the Origin of Species 1842:Transmutation of species 2036:Dual inheritance theory 1875:History of paleontology 732:10.1210/endo.142.1.7901 681:10.1073/pnas.0509716103 488:10.1073/pnas.0604934103 227:expression. Mice with 87:apical ectodermal ridge 2502:Terrestrial locomotion 2446:Evolution of cetaceans 2441:Origin of avian flight 2426:Evolution of tetrapods 1724:Punctuated equilibrium 1045:Non-adaptive radiation 993:Evolutionary arms race 773:10.1006/dbio.2001.0346 210: 146:Elongation of forelimb 57: 2517:Undulatory locomotion 2466:Homologous structures 2016:Evolutionary medicine 1890:Mendelian inheritance 1598:Biological complexity 1586:Programmed cell death 1278:Phenotypic plasticity 998:Evolutionary pressure 988:Evidence of evolution 886:Timeline of evolution 580:10.1101/gad.9.10.1237 539:10.1387/ijdb.062257cc 208: 113:in situ hybridization 51: 2461:Analogous structures 2456:Convergent evolution 1990:Teleology in biology 1885:Blending inheritance 1263:Genetic assimilation 1126:Artificial selection 865:Evolutionary biology 283:Cells Tissues Organs 260:Evolution of mammals 2512:Rotating locomotion 2451:Comparative anatomy 2053:Molecular evolution 2011:Ecological genetics 1880:Transitional fossil 1670:Sexual reproduction 1510:endomembrane system 1439:pollinator-mediated 1395:dolphins and whales 1173:Parental investment 672:2006PNAS..103.6581S 620:10.1101/gad.1620408 479:2006PNAS..10315103W 473:(41): 15103–15107. 389:1996Sci...272..738Z 338:1972NYASA.193...29S 2431:Evolution of birds 2184:Aquatic locomotion 2026:Cultural evolution 1141:Fisher's principle 1070:Handicap principle 1060:Parallel evolution 924:Adaptive radiation 211: 58: 2525: 2524: 2482:Animal locomotion 2421:Evolution of fish 2301:facultative biped 2126: 2125: 1742:Uniformitarianism 1695:Sex-determination 1200:Sexual dimorphism 1195:Natural selection 1099:Unit of selection 1065:Signalling theory 573:(10): 1237–1249. 295:10.1159/000109959 2550: 2492:Robot locomotion 2266:Limb development 2251: 2224:Lobe-finned fish 2153: 2146: 2139: 2130: 2116: 2106: 2105: 1905:Modern synthesis 1665:Multicellularity 1660:Mosaic evolution 1545:auditory ossicle 1227:Social selection 1210:Flowering plants 1205:Sexual selection 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1221: 1218: 1216: 1213: 1211: 1208: 1207: 1206: 1203: 1201: 1198: 1196: 1193: 1191: 1188: 1186: 1183: 1179: 1176: 1174: 1171: 1170: 1169: 1168:Kin selection 1166: 1164: 1163:Genetic drift 1161: 1159: 1156: 1152: 1149: 1148: 1147: 1144: 1142: 1139: 1137: 1134: 1132: 1129: 1127: 1124: 1123: 1121: 1119: 1113: 1105: 1102: 1101: 1100: 1097: 1095: 1092: 1088: 1085: 1083: 1080: 1079: 1078: 1075: 1071: 1068: 1067: 1066: 1063: 1061: 1058: 1056: 1053: 1051: 1048: 1046: 1043: 1041: 1038: 1036: 1033: 1031: 1028: 1026: 1023: 1021: 1018: 1014: 1011: 1010: 1009: 1006: 1004: 1001: 999: 996: 994: 991: 989: 986: 984: 981: 979: 976: 974: 971: 969: 966: 962: 959: 958: 957: 954: 952: 949: 947: 944: 940: 937: 935: 932: 931: 930: 927: 925: 922: 920: 917: 915: 912: 911: 909: 907: 903: 897: 894: 892: 889: 887: 884: 882: 879: 877: 874: 873: 870: 866: 859: 854: 852: 847: 845: 840: 839: 836: 826: 822: 818: 814: 810: 806: 802: 798: 791: 788: 783: 779: 774: 769: 765: 761: 757: 750: 747: 742: 738: 733: 728: 724: 720: 719:Endocrinology 716: 709: 706: 701: 697: 692: 687: 682: 677: 673: 669: 665: 661: 657: 650: 648: 644: 639: 635: 630: 625: 621: 617: 613: 609: 605: 598: 595: 590: 586: 581: 576: 572: 568: 564: 557: 554: 549: 545: 540: 535: 531: 527: 523: 516: 513: 508: 504: 499: 494: 489: 484: 480: 476: 472: 468: 464: 457: 454: 449: 445: 441: 437: 433: 429: 422: 419: 414: 410: 406: 402: 398: 394: 390: 386: 382: 378: 371: 368: 363: 359: 355: 351: 347: 343: 339: 335: 332:(17): 29–42. 331: 327: 320: 317: 312: 308: 304: 300: 296: 292: 288: 284: 277: 275: 271: 265: 261: 258: 257: 253: 251: 248: 244: 240: 236: 235: 230: 226: 222: 213: 207: 203: 201: 200: 195: 191: 187: 184: 180: 176: 172: 168: 167: 162: 158: 154: 145: 143: 140: 139: 134: 130: 126: 118: 116: 114: 110: 106: 100: 97: 93: 88: 84: 80: 76: 72: 68: 60: 55: 50: 46: 44: 39: 35: 31: 27: 23: 19: 2367: 2075:Polymorphism 2058:Astrobiology 2006:Biogeography 1961:Saltationism 1951:Orthogenesis 1936:Alternatives 1865: 1851: 1784:Cospeciation 1779:Cladogenesis 1728:Saltationism 1685:Mating types 1608:Color vision 1593:Avian flight 1515:mitochondria 1253:Canalisation 1131:Biodiversity 876:Introduction 800: 796: 790: 763: 759: 749: 722: 718: 708: 663: 659: 611: 607: 597: 570: 566: 556: 532:(4): 333–8. 529: 525: 515: 470: 466: 456: 431: 427: 421: 380: 376: 370: 329: 325: 319: 286: 282: 246: 242: 232: 228: 217: 197: 189: 182: 178: 174: 170: 164: 160: 156: 152: 149: 136: 122: 101: 64: 15: 2395:Insect wing 2345:Webbed foot 2286:unguligrade 2281:plantigrade 2276:digitigrade 2085:Systematics 1956:Mutationism 1774:Catagenesis 1702:Snake venom 1635:Eusociality 1613:in primates 1603:Cooperation 1531:In animals 1351:butterflies 1324:Cephalopods 1314:Brachiopods 1246:Development 1220:Mate choice 973:Convergence 956:Coevolution 914:Abiogenesis 289:(1): 6–12. 194:chondrocyte 96:orthologous 2532:Categories 2323:Cephalopod 2239:Pelvic fin 2209:Dorsal fin 2204:Caudal fin 1946:Lamarckism 1924:Philosophy 1847:David Hume 1809:Peripatric 1804:Parapatric 1789:Ecological 1769:Anagenesis 1764:Allopatric 1756:Speciation 1720:Gradualism 1645:Metabolism 1505:chromosome 1495:Eukaryotes 1273:Modularity 1190:Population 1116:Population 1077:Speciation 1055:Panspermia 1008:Extinction 1003:Exaptation 978:Divergence 951:Cladistics 939:Reciprocal 919:Adaptation 266:References 109:regulatory 54:homologous 38:homologous 18:Chiroptera 16:The order 2414:Evolution 2373:Bird wing 2318:Arthropod 2311:quadruped 2080:Protocell 1931:Darwinism 1819:Sympatric 1568:processes 1456:Tetrapods 1405:Kangaroos 1331:Dinosaurs 1268:Inversion 1237:Variation 1158:Gene flow 1151:Inclusive 961:Mutualism 906:Evolution 760:Dev. Biol 608:Genes Dev 567:Genes Dev 133:apoptosis 129:apoptosis 65:Tetrapod 2405:Wingspan 2388:feathers 2383:skeleton 2368:Bat wing 2328:Tetrapod 2214:Fish fin 2108:Category 1983:Vitalism 1978:Theistic 1971:Spandrel 1655:Morality 1650:Monogamy 1525:plastids 1490:Flagella 1446:Reptiles 1427:sea cows 1410:primates 1319:Molluscs 1297:Bacteria 1185:Mutation 1118:genetics 1094:Taxonomy 1040:Mismatch 1020:Homology 934:Cheating 929:Altruism 825:12204277 817:15733311 782:11476582 741:11145607 700:16618938 638:18198333 548:17554686 507:17015842 448:15863030 413:27174863 362:30365208 303:18160799 254:See also 239:Hox gene 186:enhancer 125:patagium 94:of some 90:spatial 43:evo-devo 34:skeletal 26:tetrapod 2475:Related 2333:dactyly 2219:Flipper 1999:Related 1829:History 1690:Meiosis 1625:Empathy 1620:Emotion 1520:nucleus 1461:Viruses 1451:Spiders 1363:Mammals 1346:Insects 1146:Fitness 1082:Species 881:Outline 691:1458926 668:Bibcode 629:2192750 589:7758948 498:1622783 475:Bibcode 405:8614838 385:Bibcode 377:Science 354:4561097 334:Bibcode 311:9969396 234:Hox-d13 85:. The 30:mammals 2497:Samara 2306:triped 2291:uniped 2118:Portal 1794:Hybrid 1630:Ethics 1472:organs 1434:Plants 1420:lemurs 1415:humans 1400:horses 1390:hyenas 1378:wolves 1373:canids 1307:origin 823:  815:  780:  739:  698:  688:  636:  626:  587:  546:  505:  495:  446:  411:  403:  360:  352:  309:  301:  247:Hoxd13 243:Hoxd13 32:, the 22:flight 2355:Wings 2340:Digit 2296:biped 2258:Limbs 2168:wings 2164:limbs 1581:Death 1576:Aging 1555:brain 1341:Fungi 1302:Birds 1215:Fungi 1013:Event 896:Index 821:S2CID 409:S2CID 358:S2CID 307:S2CID 199:Bmp-2 2487:Gait 2378:keel 2176:Fins 2166:and 2160:Fins 2068:Tree 1540:hair 1480:Cell 1383:dogs 1368:cats 1358:Life 1336:Fish 1289:taxa 813:PMID 778:PMID 737:PMID 696:PMID 634:PMID 585:PMID 544:PMID 503:PMID 444:PMID 401:PMID 350:PMID 299:PMID 221:ulna 190:prx1 183:prx1 179:prx1 175:prx1 171:prx1 166:prx1 161:fgf8 157:fgf8 153:fgf8 138:Fgf8 81:and 1566:Of 1535:eye 1485:DNA 1470:Of 1287:Of 805:doi 768:doi 764:236 727:doi 723:142 686:PMC 676:doi 664:103 624:PMC 616:doi 575:doi 534:doi 493:PMC 483:doi 471:103 436:doi 393:doi 381:272 342:doi 330:193 291:doi 287:187 229:shh 225:SHH 105:cis 83:WNT 79:SHH 75:BMP 71:FGF 2534:: 2162:, 819:. 811:. 799:. 776:. 762:. 758:. 735:. 721:. 717:. 694:. 684:. 674:. 662:. 658:. 646:^ 632:. 622:. 612:22 610:. 606:. 583:. 569:. 565:. 542:. 530:51 528:. 524:. 501:. 491:. 481:. 469:. 465:. 442:. 432:16 430:. 407:. 399:. 391:. 379:. 356:. 348:. 340:. 328:. 305:. 297:. 285:. 273:^ 77:, 73:, 2152:e 2145:t 2138:v 1744:/ 1735:/ 1726:/ 1722:/ 857:e 850:t 843:v 827:. 807:: 801:7 784:. 770:: 743:. 729:: 702:. 678:: 670:: 640:. 618:: 591:. 577:: 571:9 550:. 536:: 509:. 485:: 477:: 450:. 438:: 415:. 395:: 387:: 364:. 344:: 336:: 313:. 293:: 107:-

Index

Chiroptera
flight
tetrapod
mammals
skeletal
homologous
evo-devo
skeletal forelimbs
homologous
limb development
FGF
BMP
SHH
WNT
apical ectodermal ridge
gene expression
orthologous
cis
regulatory
in situ hybridization
patagium
apoptosis
apoptosis
Fgf8
prx1
enhancer
chondrocyte
Bmp-2
developing bat forelimb showing gene expression patterns
ulna

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