2249:
206:
2104:
2114:
49:
141:
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
40:
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
102:
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
89:
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
150:
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
98:
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.
249:
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.
218:
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
131:
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
192:
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
223:
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
1836:
135:. However, in bats, BMP genes are still expressed in the interdigits and yet interdigit apoptosis is repressed. FGF signaling has been associated with blocking cell death.
209:
Simplified diagram showing expanded gene expression domains in developing bat forelimb potentially contributing to the morphological changes resulting in the bat wing.
2150:
202:
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.
885:
795:
Chen CH, Cretekos CJ, Rasweiler JJ, Behringer RR (2005). "Hoxd13 expression in the developing limbs of the short-tailed fruit bat, Carollia perspicillata".
159:
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
2542:
2506:
127:) allowed a greater surface area of the wing necessary for flight. All vertebrate limb formation initially has tissue between the digits after which
2089:
1544:
895:
2511:
1852:
1612:
1209:
245:
expression domain in bat limbs has been shifted posteriorly in comparison to mouse. This observed difference in the expression pattern of
196:
proliferation. In addition to interdigit apoptosis, BMPs have been shown to affect chondrocyte proliferation and digit length in mice.
2143:
1935:
855:
1597:
1257:
42:
1669:
1267:
1914:
1904:
1828:
1103:
41:
bat forelimb development to understand the genetic basis of morphological evolution. Consequently, the bat wing is a valuable
1231:
2136:
2042:
1909:
1689:
1433:
1135:
1024:
1177:
2248:
2107:
1438:
2047:
1549:
198:
324:
Saunders JW; Benthe, HF; Haberland, G (1972). "Developmental control of three-dimensional polarity in the avian limb".
2270:
2117:
1711:
2362:
2198:
1813:
987:
74:
1607:
1404:
1214:
982:
875:
463:"Interdigital webbing retention in bat wings illustrates genetic changes underlying amniote limb diversification"
2030:
1894:
1866:
1841:
1798:
1701:
1694:
1634:
1382:
1350:
1323:
1313:
137:
70:
522:"genomic structure and developmental expression of Fgf8 in the short-tailed fruit bat, Carollia perspicillata"
155:
in bat forelimb AER are expanded in comparison to the mouse forelimb, suggesting that expanded expression of
2035:
1965:
1874:
1602:
1484:
1252:
848:
86:
2537:
2501:
2445:
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1723:
1664:
1592:
1554:
1455:
1426:
1399:
1394:
992:
375:
Zou H, Niswander L (1996). "Requirement for BMP Signaling in
Interdigital Apoptosis and Scale Formation".
426:
Eswarakumar V, Lax I, Schlessinger J (2005). "Cellular signaling by fibroblast growth factor receptors".
103:
between mouse and bat. Thus, it is likely that this major morphological transition was a consequence of
2516:
2015:
1889:
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1318:
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997:
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82:
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1989:
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1362:
1345:
1272:
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972:
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667:
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259:
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has an expanded expression domain and is upregulated. Researchers found that the coding region of
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2010:
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1125:
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933:
928:
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841:
820:
756:"Manifestation of the limb prepattern: limb development in the absence of sonic hedgehog pattern"
408:
357:
306:
53:
37:
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changes. Researchers can study the genetic basis of bat wing development by using comparative
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290:
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null mutations lose their ulna structure. Another good candidate for bat bone reduction is
66:
2435:
2339:
2322:
2193:
2020:
1471:
1460:
1414:
1357:
1306:
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De Luca F, Barnes KM, Uyeda JA, De-Levi S, Abad V, Palese T, Mericq V, Baron J (2001).
690:
655:
628:
603:
497:
462:
345:
224:
115:
to examine gene expression domains and using experimental embryology in mice and bats.
78:
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2317:
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808:
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412:
361:
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2057:
2005:
1950:
1783:
1778:
1519:
1130:
656:"Development of bat flight: morphologic and molecular evolution of bat wing digits"
310:
48:
754:
Chiang C, Litingtung Y, Harris MP, Simandl BK, Li Y, Beachy PA, Fallon JF (2001).
439:
396:
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2344:
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2275:
2084:
1955:
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1684:
1219:
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913:
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Proceedings of the
National Academy of Sciences of the United States of America
467:
Proceedings of the
National Academy of Sciences of the United States of America
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1945:
1846:
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17:
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1157:
905:
731:
714:
680:
487:
132:
128:
816:
781:
772:
755:
740:
715:"Regulation of growth plate chondrogenesis by bone morphogenetic protein-2"
699:
637:
547:
506:
447:
302:
588:
579:
563:"MHox is required for early events of skeletogenesis in multiple lineages"
562:
538:
521:
404:
353:
2404:
2327:
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2213:
1982:
1330:
1184:
238:
124:
95:
33:
25:
2128:
619:
2332:
1624:
1524:
1372:
1367:
1081:
833:
29:
294:
20:, comprising all bats, has evolved the unique mammalian adaptation of
2290:
233:
21:
169:, which encodes a transcription factor. The expression patterns of
520:
Cretekos CJ, Deng JM, Green ED, Rasweiler JJ, Behringer RR (2007).
281:
Sears KE (2008). "Molecular determinants of bat wing development".
1580:
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2354:
2167:
1539:
220:
2132:
837:
461:
Weatherbee SD, Behringer RR, Rasweiler JJ, Niswander L (2006).
45:
model for studying the evolution of vertebrate limb diversity.
2175:
2159:
181:
in bats is nearly identical to mice but found a bat-specific
604:"Regulatory divergence modifies limb length between mammals"
654:
Sears KE, Behringer RR, Rasweiler JJ, Niswander LA (2006).
241:
family. In situ hybridization studies have found that the
163:. In mice, one gene known to regulate limb growth is
2474:
2413:
2353:
2256:
2174:
1998:
1923:
1827:
1754:
1710:
1565:
1469:
1286:
1245:
1114:
904:
602:Cretekos CJ, Wang Y, Green ED, et al. (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
858:
851:
844:
835:
829:
828:
792:
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383:(5262): 738–41.
372:
366:
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321:
315:
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278:
67:limb development
52:diagram showing
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2557:
2553:
2552:
2551:
2549:
2548:
2547:
2528:
2527:
2526:
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2470:
2436:Origin of birds
2409:
2349:
2271:Limb morphology
2252:
2243:
2229:Ray-finned fish
2194:Fish locomotion
2170:
2157:
2127:
2122:
2094:
2021:Group selection
1994:
1919:
1823:
1750:
1712:Tempo and modes
1706:
1561:
1465:
1282:
1241:
1117:
1110:
1087:Species complex
900:
891:History of life
867:
862:
832:
794:
793:
789:
753:
752:
748:
712:
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92:gene expression
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2400:Pterosaur wing
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2201:
2196:
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2189:Cephalopod fin
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2018:
2013:
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1987:
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1877:
1872:
1871:
1870:
1861:Charles Darwin
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1856:
1844:
1839:
1833:
1831:
1825:
1824:
1822:
1821:
1816:
1811:
1806:
1801:
1799:Non-ecological
1796:
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1781:
1776:
1771:
1766:
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1572:
1570:
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1550:nervous system
1547:
1542:
1537:
1529:
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1507:
1502:
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1407:
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1397:
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1387:
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1316:
1311:
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1255:
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1234:
1229:
1224:
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1202:
1197:
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1187:
1182:
1181:
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1175:
1165:
1160:
1155:
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1153:
1143:
1138:
1133:
1128:
1122:
1120:
1112:
1111:
1109:
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1106:
1096:
1091:
1090:
1089:
1084:
1074:
1073:
1072:
1062:
1057:
1052:
1050:Origin of life
1047:
1042:
1037:
1035:Microevolution
1032:
1030:Macroevolution
1027:
1022:
1017:
1016:
1015:
1005:
1000:
995:
990:
985:
980:
975:
970:
968:Common descent
965:
964:
963:
953:
948:
946:Baldwin effect
943:
942:
941:
936:
926:
921:
916:
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883:
878:
872:
869:
868:
863:
861:
860:
853:
846:
838:
831:
830:
803:(2): 130–141.
787:
766:(2): 421–435.
746:
725:(1): 430–436.
705:
666:(17): 6581–6.
643:
614:(2): 144–151.
594:
553:
526:Int J Dev Biol
512:
453:
434:(2): 139–149.
418:
367:
316:
269:
267:
264:
263:
262:
255:
252:
215:
212:
147:
144:
142:the patagium.
120:
117:
62:
59:
13:
10:
9:
6:
4:
3:
2:
2555:
2544:
2541:
2539:
2538:Animal flight
2536:
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2515:
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2505:
2503:
2500:
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2279:
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2269:
2267:
2264:
2263:
2261:
2259:
2255:
2250:
2240:
2237:
2235:
2234:Pectoral fins
2232:
2230:
2227:
2225:
2222:
2220:
2217:
2215:
2212:
2210:
2207:
2205:
2202:
2200:
2197:
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2173:
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2119:
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2097:
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2088:
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2081:
2078:
2076:
2073:
2069:
2066:
2065:
2064:
2063:Phylogenetics
2061:
2059:
2056:
2054:
2051:
2049:
2046:
2044:
2041:
2037:
2034:
2032:
2029:
2027:
2024:
2023:
2022:
2019:
2017:
2014:
2012:
2009:
2007:
2004:
2003:
2001:
1997:
1991:
1988:
1984:
1981:
1979:
1976:
1972:
1969:
1968:
1967:
1966:Structuralism
1964:
1962:
1959:
1957:
1954:
1952:
1949:
1947:
1944:
1942:
1941:Catastrophism
1939:
1938:
1937:
1934:
1932:
1929:
1928:
1926:
1922:
1916:
1913:
1911:
1908:
1906:
1903:
1901:
1900:Neo-Darwinism
1898:
1896:
1893:
1891:
1888:
1886:
1883:
1881:
1878:
1876:
1873:
1869:
1868:
1864:
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1859:
1855:
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1850:
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1843:
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1835:
1834:
1832:
1830:
1826:
1820:
1817:
1815:
1814:Reinforcement
1812:
1810:
1807:
1805:
1802:
1800:
1797:
1795:
1792:
1790:
1787:
1785:
1782:
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1777:
1775:
1772:
1770:
1767:
1765:
1762:
1761:
1759:
1757:
1753:
1747:
1746:Catastrophism
1743:
1740:
1738:
1737:Macromutation
1734:
1733:Micromutation
1731:
1729:
1725:
1721:
1718:
1717:
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1713:
1709:
1703:
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1696:
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1668:
1666:
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1653:
1651:
1648:
1646:
1643:
1641:
1640:Immune system
1638:
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1633:
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1623:
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1614:
1611:
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1530:
1526:
1523:
1521:
1518:
1516:
1513:
1511:
1508:
1506:
1503:
1501:
1500:symbiogenesis
1498:
1497:
1496:
1493:
1491:
1488:
1486:
1483:
1481:
1478:
1477:
1475:
1473:
1468:
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1408:
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1403:
1401:
1398:
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1325:
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1300:
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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:
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1123:
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1079:
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1038:
1036:
1033:
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1028:
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1023:
1021:
1018:
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1009:
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989:
986:
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949:
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927:
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922:
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917:
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845:
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839:
836:
826:
822:
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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:-
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