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Arthur Cain

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260:) are dimorphic: the common morph ('white') is white in winter and brownish-grey dorsally in summer; the other morph ('blue') is light brown/blue in winter and dark brown in summer. The two morphs interbreed freely. Despite the obvious advantage of white in avoiding predation, blue is actually the most frequent morph in 269:
That very phenomenon which was to be used by Fisher & Ford in studies on natural selection is here shown by cogent argument and the facts of field natural history to be inexplicable by selection. But Elton knew that a far greater range of other characters have the same implications, namely, all,
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Also, Robson and Richards "showed a surprising reluctance to allow any example of natural selection; their cautious qualification that characters were non-adaptive as far as they could see became, too often, a certainty that they were non-adaptive; and their arguments were sometimes one-sided." Cain
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appreciated that, even if a character was in itself non-adaptive, the gene or genes determining it might affect other, adaptive, characters which were always under selective influence. Ford understood the significance of
348:'s demonstration that a neutral gene derived from a single mutation could only be in about the same number of individuals as there had been generations since its inception. Also, as Cain's own research showed, much 176:, one of the first studies to demonstrate natural selection by predators acting on a colour polymorphism, is now regarded as a classic. It generated a long series of further studies by Cain, including the formal 80:
After leaving the military in November 1945, Cain returned to Oxford to pursue research in the Department of Zoology. He became a Departmental Demonstrator in October 1946, finishing his
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What I wanted to know from all these great people was, how exactly did they know that a character was non-adaptive or neutral? They didn't know, and they couldn't know.
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David Lack was the only religious man I knew at that period who did not allow his religion to dictate his view of natural selection.
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in 1948. From January 1949 until 1964 Cain was employed as University Demonstrator in Animal Taxonomy. In addition he was appointed
1216: 1211: 74: 1236: 997: 1149: 332:, who were also believing Christians: Fisher from the start of his career was a leading proponent of natural selection. 901: 874: 697: 604: 228: 188:
material to follow changes in time as well as space. Later he turned to the study of variation in shell shape.
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laid the blame on their "vitalistic or perhaps theistic attitudes... Robson and Richards were far from alone.
93: 264:. Elton also gave a number of other examples which he claimed could not be explained by natural selection. 404:
Cain A.J. 1983. Ecology and ecogenetics of terrestrial molluscan populations. In Russell-Hunter W.D. (ed)
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Cain A.J., King J.M.B. and Sheppard P.M. 1960. New data on the genetics of polymorphism in the snail
360:(a full bibliography listing 148 items appears as supplementary material to Bryan Clarke's obituary) 66: 1165: 1022: 232: 100: 89: 54: 352:
is maintained by differential selection in the diversity of environments within a species' range.
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Cain A.J. and Provine W.B. 1991. Genes and ecology in history. In Berry R.J. & others (eds)
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of the variation, the discovery of area effects and the analysis of climatic influences. With
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Cook, L.M. (2024). "Arthur Cain and ecological genetics in the Oxford Zoology Department".
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Cain A.J. and Provine W.B. 1991. Genes and ecology in history. In Berry R.J. et al. (eds)
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in Zoology in 1941. Entering the British army in December 1941, Cain was commissioned
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Cain. This was the stimulus for Cain's research on evolution in natural communities.
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or nearly all, the differences (non-polymorphic) between closely related species
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Towards the end of his life Cain was persuaded to reminisce about the status of
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Cain A.J. 1971. Colour and banding morphs in subfossil samples of the snail
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in Warwickshire, England. In 1939 he was awarded a prestigious scholarship (
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Currey J.D. and Cain A.J. 1968. Climate and selection of banding morphs in
471:: the 33rd Symposium of the British Ecological Society. Blackwell, Oxford. 341: 282: 251: 1077: 1028: 762:
Cain & Provine W.B. 1991. Genes and ecology in history. In Berry R.J.
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Cain & Provine W.B. 1991. Genes and ecology in history. In Berry R.J.
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Cain & Provine W.B. 1991. Genes and ecology in history. In Berry R.J.
686:: the 33rd Symposium of the British Ecological Society. Blackwell, Oxford. 474:
Cain A.J. and Sheppard P.M. 1950. Selection in the polymorphic land snail
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Cain A.J. 1977. The efficacy of natural selection in wild populations. In
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Clarke, B.C. (2008). "Arthur James Cain. 25 July 1921 – 20 August 1999".
128: 1070: 289:" (p7)... In Cambridge matters were even worse" (p8, giving as examples 1044: 992: 982: 962: 240: 185: 132: 103:, and he later (1968) was appointed Derby Professor of Zoology at the 336: 151: 30: 286: 108: 50: 261: 236: 204: 85: 909: 401:. Special publication #12, 111–33. Academy of Natural Sciences. 215:
to our understanding of variation between taxonomic categories.
195:. He made important contributions to the theory and practice of 99:
In 1964, Cain left Oxford to become professor of zoology at the
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Collection of papers held by the American Philosophical Society
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Cain A.J. and Currey J.D. 1968. Ecogenetics of a population of
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Cain A.J. and Currey J.D. 1968. Ecogenetics of a population of
247: 244: 167: 147: 208: 162:, Cain studied the ecological genetics of colour and banding 231:. The general attitude was sceptical of natural selection. 227:
in pre-war Oxford and how it changed over the years of the
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Cain A.J., Sheppard P.M. and King J.M.B. 1968. Studies on
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Cain A.J. and Sheppard P.M. 1954. Natural selection in
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Clarke B.C. 1979. The evolution of genetic diversity.
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Clarke B.C. 1979. The evolution of genetic diversity.
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Biographical Memoirs of Fellows of the Royal Society
218: 429:on the Larkhill Artillery Ranges, Salisbury Plain. 191:In population genetics he clarified the concept of 92:in 1954, besides serving as Lecturer in Zoology at 1227:Royal Electrical and Mechanical Engineers officers 425:Cain A.J. and Currey J.D. 1963b. Area effects in 411:Cain A.J. and Currey J.D. 1963a. Area effects in 111:status at Liverpool upon his retirement in 1989. 1203: 814:Cain A.J. and Currey J.D. 1963. Area effects in 506:I. The genetics of some morphs and varieties of 73:(R.E.M.E.) on its formation. He was promoted to 207:and taxonomic importance, on the status of the 69:(engineering) and was later transferred to the 792:. All Souls Studies, Faber & Faber, London 243:polymorphism, which can be found in all three 150:, his main work lay in the field developed by 25:(25 July 1921 – 20 August 1999) was a British 895: 537:from the climate optimum to the present day. 562: 560: 558: 138:Though he initially conducted research with 902: 888: 45:Arthur James Cain was born and grew up in 555: 71:Royal Electrical and Mechanical Engineers 324:He might have added, had he known them, 114: 649: 619: 355: 1252:British Army personnel of World War II 1204: 605:"Arthur J. Cain Papers Mss.Ms.Coll.63" 566: 408:vol 6, p597-647. Academic Press, N.Y. 399:The changing scene in natural sciences 1257:Academics of the University of Oxford 883: 803:Genetical theory of natural selection 88:of the Zoological Collections at the 736:Robson G.C. and Richards O.W. 1936. 655: 625: 1150:Edward Shackleton, Baron Shackleton 13: 1262:Alumni of Magdalen College, Oxford 1232:Royal Army Ordnance Corps officers 738:The variation of animals in nature 597: 365:Animal species and their evolution 14: 1273: 868: 392:Ecological genetics and evolution 239:as a discipline, pointed out the 219:Cain reminisces on pre-war Oxford 838:subject to strong area effects. 607:. American Philosophical Society 443:subject to strong area effects. 431:J. Linnaean Soc London (Zoology) 1217:British evolutionary biologists 1212:20th-century British biologists 848: 828: 808: 795: 782: 769: 756: 743: 698:Evolution: the modern synthesis 775:Ford E.B. 1964, 4th edn 1975. 730: 717: 704: 689: 676: 170:. Cain and Sheppard's work on 1: 549: 229:modern evolutionary synthesis 119:Cain's main interests lay in 1237:Fellows of the Royal Society 628:"A.J. Cain F.R.S. 1921-1929" 335:In contrast to many others, 7: 1029:Andrew David McLachlan 658:Archives of Natural History 370:Cain A.J. 1968. Studies on 235:, who led the emergence of 35:Fellow of the Royal Society 10: 1278: 779:. Chapman and Hall, London 701:(2nd ed 1963, 3rd ed 1974) 211:, and on the relevance of 57:, where he graduated with 1158: 1142: 925: 67:Royal Army Ordnance Corps 40: 1166:Nicole Marthe Le Douarin 1023:Gilbert George Lonzarich 101:University of Manchester 90:Oxford University Museum 55:Magdalen College, Oxford 1084:John Philip Simons 1071:John Martin Rowell 184:he made elegant use of 105:University of Liverpool 1050:Robert Ladislav Parker 632:Malacological Bulletin 583:10.1098/rsbm.2007.0034 540:Phil. Trans. R. Soc. B 513:Phil. Trans. R. Soc. B 377:Phil. Trans. R. Soc. B 367:. Hutchinson, London. 27:evolutionary biologist 1078:Ian John Russell 394:. Blackwell, Oxford. 330:Theodosius Dobzhansky 115:Scientific activities 1191:Edward Mills Purcell 840:Phil Trans Roy Soc B 820:Phil Trans Roy Soc B 790:Genetic polymorphism 670:10.3366/anh.20240897 445:Phil Trans Roy Soc B 417:Phil Trans Roy Soc B 356:Notable publications 121:evolutionary biology 33:. He was elected a 1124:Thomas Summers West 777:Ecological genetics 740:. Longmans, London. 390:. In Creed R. (ed) 281:... was an earnest 156:ecological genetics 125:ecological genetics 1222:British ecologists 1186:Barbara McClintock 973:Peter Goddard 801:Fisher R.A. 1930. 205:phyletic weighting 199:, the problems of 94:St Peter's College 59:first class honors 1199: 1198: 1129:Andrew Wiles 1094:Marshall Stoneham 710:Elton C.S. 1927. 626:Cook, L. (2000). 225:natural selection 213:natural selection 63:second lieutenant 20:Arthur James Cain 16:British biologist 1269: 1134:Ian Robert Young 1119:Rob Webster 1089:Roger Slack 1035:Michael A. Moore 1018:John Lawton 938:Kenneth Bagshawe 904: 897: 890: 881: 880: 863: 852: 846: 836:Cepaea nemoralis 832: 826: 812: 806: 799: 793: 788:Ford E.B. 1965. 786: 780: 773: 767: 764:Genes in ecology 760: 754: 751:Genes in ecology 747: 741: 734: 728: 725:Genes in ecology 721: 715: 708: 702: 695:Huxley J. 1942. 693: 687: 684:Genes in ecology 680: 674: 673: 653: 647: 646: 644: 642: 623: 617: 616: 614: 612: 601: 595: 594: 564: 508:Cepaea nemoralis 476:Cepaea nemoralis 469:Genes in ecology 455:Cepaea nemoralis 441:Cepaea nemoralis 363:Cain A.J. 1954. 285:and certainly a 254:. Arctic foxes ( 250:of the northern 178:genetic analysis 173:Cepaea nemoralis 1277: 1276: 1272: 1271: 1270: 1268: 1267: 1266: 1202: 1201: 1200: 1195: 1181:Edward B. Lewis 1154: 1138: 1099:Roger C. Thomas 988:Geoffrey Hewitt 953:Roger Blandford 921: 908: 871: 866: 853: 849: 833: 829: 813: 809: 800: 796: 787: 783: 774: 770: 761: 757: 748: 744: 735: 731: 722: 718: 709: 705: 694: 690: 681: 677: 654: 650: 640: 638: 624: 620: 610: 608: 603: 602: 598: 565: 556: 552: 358: 221: 129:animal taxonomy 117: 43: 17: 12: 11: 5: 1275: 1265: 1264: 1259: 1254: 1249: 1244: 1239: 1234: 1229: 1224: 1219: 1214: 1197: 1196: 1194: 1193: 1188: 1183: 1178: 1173: 1168: 1162: 1160: 1156: 1155: 1153: 1152: 1146: 1144: 1140: 1139: 1137: 1136: 1131: 1126: 1121: 1116: 1111: 1106: 1101: 1096: 1091: 1086: 1081: 1074: 1067: 1062: 1057: 1052: 1047: 1042: 1037: 1032: 1025: 1020: 1015: 1010: 1005: 1000: 995: 990: 985: 980: 975: 970: 965: 960: 955: 950: 945: 940: 935: 933:Brian Anderson 929: 927: 923: 922: 907: 906: 899: 892: 884: 878: 877: 870: 869:External links 867: 865: 864: 856:Proc Roy Soc B 847: 827: 807: 794: 781: 768: 755: 742: 729: 716: 712:Animal ecology 703: 688: 675: 648: 618: 596: 553: 551: 548: 524:Proc Roy Soc B 357: 354: 344:, and knew of 322: 321: 314: 313: 303:J.W.S. Pringle 274: 273: 257:Alopex lagopus 220: 217: 193:adaptive value 144:histochemistry 116: 113: 107:. He received 42: 39: 15: 9: 6: 4: 3: 2: 1274: 1263: 1260: 1258: 1255: 1253: 1250: 1248: 1245: 1243: 1240: 1238: 1235: 1233: 1230: 1228: 1225: 1223: 1220: 1218: 1215: 1213: 1210: 1209: 1207: 1192: 1189: 1187: 1184: 1182: 1179: 1177: 1176:Kenichi Fukui 1174: 1172: 1169: 1167: 1164: 1163: 1161: 1157: 1151: 1148: 1147: 1145: 1141: 1135: 1132: 1130: 1127: 1125: 1122: 1120: 1117: 1115: 1112: 1110: 1107: 1105: 1104:Anne Treisman 1102: 1100: 1097: 1095: 1092: 1090: 1087: 1085: 1082: 1080: 1079: 1075: 1073: 1072: 1068: 1066: 1063: 1061: 1058: 1056: 1053: 1051: 1048: 1046: 1043: 1041: 1038: 1036: 1033: 1031: 1030: 1026: 1024: 1021: 1019: 1016: 1014: 1013:Francis Kelly 1011: 1009: 1008:Michael James 1006: 1004: 1003:Richard Hynes 1001: 999: 996: 994: 991: 989: 986: 984: 981: 979: 978:Michael Green 976: 974: 971: 969: 966: 964: 961: 959: 956: 954: 951: 949: 946: 944: 941: 939: 936: 934: 931: 930: 928: 924: 920: 916: 915:Royal Society 912: 905: 900: 898: 893: 891: 886: 885: 882: 876: 873: 872: 861: 857: 851: 844: 841: 837: 831: 824: 821: 817: 811: 804: 798: 791: 785: 778: 772: 765: 759: 752: 746: 739: 733: 726: 720: 713: 707: 700: 699: 692: 685: 679: 671: 667: 663: 659: 652: 637: 633: 629: 622: 606: 600: 592: 588: 584: 580: 576: 572: 571: 563: 561: 559: 554: 547: 545: 542: 541: 536: 531: 529: 525: 520: 518: 515: 514: 509: 505: 500: 498: 495: 491: 486: 484: 481: 477: 472: 470: 465: 463: 460: 456: 451: 449: 446: 442: 437: 435: 432: 428: 423: 421: 418: 414: 409: 407: 402: 400: 395: 393: 389: 384: 382: 379: 378: 373: 368: 366: 361: 353: 351: 347: 343: 338: 333: 331: 327: 326:Ronald Fisher 319: 316: 315: 311: 308: 307: 306: 304: 300: 296: 295:Charles Raven 292: 288: 284: 280: 279:Alister Hardy 271: 267: 266: 265: 263: 259: 258: 253: 249: 246: 242: 238: 234: 233:Charles Elton 230: 226: 216: 214: 210: 206: 202: 198: 194: 189: 187: 183: 179: 175: 174: 169: 165: 164:polymorphisms 161: 160:P.M. Sheppard 157: 153: 149: 145: 141: 136: 134: 130: 126: 122: 112: 110: 106: 102: 97: 96:(1958–1961). 95: 91: 87: 83: 78: 76: 72: 68: 64: 60: 56: 52: 48: 38: 36: 32: 28: 24: 21: 1109:Kenneth Wade 1076: 1069: 1065:Warren Roper 1060:Richard Peto 1055:Geoff Parker 1027: 968:Alwyn Davies 957: 943:John M. Ball 859: 855: 850: 842: 839: 835: 830: 822: 819: 815: 810: 802: 797: 789: 784: 776: 771: 763: 758: 750: 745: 737: 732: 724: 719: 711: 706: 696: 691: 683: 678: 664:(1): 73–85. 661: 657: 651: 639:. Retrieved 635: 631: 621: 609:. Retrieved 599: 574: 568: 543: 538: 534: 532: 530:, 453–474. 527: 523: 521: 516: 511: 507: 503: 501: 496: 493: 489: 487: 482: 479: 475: 473: 468: 466: 461: 458: 454: 452: 447: 444: 440: 438: 433: 430: 426: 424: 419: 416: 412: 410: 406:The Mollusca 405: 403: 398: 396: 391: 387: 385: 380: 375: 371: 369: 364: 362: 359: 350:polymorphism 342:pleiotropism 334: 323: 317: 309: 275: 268: 255: 252:palaeoarctic 222: 190: 171: 137: 118: 98: 79: 44: 19: 18: 1247:1999 deaths 1242:1921 births 1040:Kim Nasmyth 998:Julian Hunt 958:Arthur Cain 948:Adrian Bird 519:, 383–396. 464:, 393–411. 450:, 447–482. 422:, 269–299. 383:, 499–517. 291:W.H. Thorpe 182:John Currey 1206:Categories 1171:Paul ErdΕ‘s 1143:Statute 12 1045:Paul Nurse 993:Derek Hull 983:Brian Heap 963:Mark Child 862:, 453–474. 845:, 447–482. 825:, 269–299. 550:References 546:, 483–98. 499:, 89–116. 485:, 275–94. 299:James Gray 241:Arctic fox 186:sub-fossil 154:, namely, 140:John Baker 133:speciation 577:: 47–57. 337:E.B. Ford 283:Unitarian 152:E.B. Ford 77:in 1942. 37:in 1989. 31:ecologist 1114:Pat Wall 917:elected 591:73131707 494:Genetics 480:Heredity 459:Genetics 436:, 1–15. 287:vitalist 201:homology 197:taxonomy 109:emeritus 82:D. Phil. 51:Demyship 1159:Foreign 926:Fellows 919:in 1989 913:of the 911:Fellows 272:. Cain. 262:Iceland 237:ecology 158:. With 142:on the 86:Curator 75:captain 65:in the 816:Cepaea 641:2 July 611:2 July 589:  535:Cepaea 510:(L.). 504:Cepaea 490:Cepaea 478:(L.). 427:Cepaea 413:Cepaea 388:Cepaea 372:Cepaea 346:Fisher 297:, Sir 248:biomes 245:tundra 168:snails 148:lipids 41:Career 587:S2CID 320:Cain. 209:genus 53:) to 47:Rugby 805:p80. 643:2024 613:2024 328:and 301:and 131:and 29:and 860:205 843:253 823:246 666:doi 579:doi 544:253 528:205 517:253 448:253 420:246 381:253 374:V. 305:). 166:in 146:of 23:FRS 1208:: 858:. 818:. 766:p9 753:p8 727:p5 662:51 660:. 636:34 634:. 630:. 585:. 575:54 573:. 557:^ 526:. 497:39 492:. 462:45 457:. 434:45 415:. 293:, 203:, 135:. 127:, 123:, 903:e 896:t 889:v 714:. 672:. 668:: 645:. 615:. 593:. 581:: 483:4

Index

FRS
evolutionary biologist
ecologist
Fellow of the Royal Society
Rugby
Demyship
Magdalen College, Oxford
first class honors
second lieutenant
Royal Army Ordnance Corps
Royal Electrical and Mechanical Engineers
captain
D. Phil.
Curator
Oxford University Museum
St Peter's College
University of Manchester
University of Liverpool
emeritus
evolutionary biology
ecological genetics
animal taxonomy
speciation
John Baker
histochemistry
lipids
E.B. Ford
ecological genetics
P.M. Sheppard
polymorphisms

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