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C. H. Waddington

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960: 981: 972: 908:". Adam S. Wilkins wrote that " in his lifetime... was widely perceived primarily as a critic of Neo-Darwinian evolutionary theory. His criticisms ... were focused on what he saw as unrealistic, 'atomistic' models of both gene selection and trait evolution." In particular, according to Wilkins, Waddington felt that the Neo-Darwinians badly neglected the phenomenon of extensive gene interactions and that the "randomness" of mutational effects, posited in the theory, was false. Even though Waddington became critical of the neo-darwinian synthetic theory of evolution, he still described himself as a Darwinian, and called for an 830: 179: 591:. In 1910, at the age of four, he was sent to live with family in England including his aunt, uncle, and Quaker grandmother. His parents remained in India until 1928. During his childhood, he was particularly attached to a local druggist and distant relation, Dr. Doeg. Doeg, whom Waddington called "Grandpa", introduced Waddington to a wide range of sciences from chemistry to geology. During the year following the completion of his entrance exams to university, Waddington received an intense course in chemistry from 77: 36: 634:, earning a First in Part II in geology in 1926. In 1928, he was awarded an Arnold Gerstenberg Studentship in the University of Cambridge, whose purpose was to promote "the study of Moral Philosophy and Metaphysics among students of Natural Science, both men and women." He took up a Lecturership in Zoology and was a Fellow of 923:
did not describe himself as a Lamarckian, but by revealing mechanisms of inheritance of acquired characteristics, I think he should be regarded as such. The reason he did not do so is that Lamarck could not have conceived of the processes that Waddington revealed. Incidentally, it is also true to say
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to represent this cellular developmental process. The idea was based on experiment: Waddington found that one effect of mutation (which could modulate the epigenetic landscape) was to affect how cells differentiated. He also showed how mutation could affect the landscape, and used this metaphor in
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In the early 1930s, Waddington and many other embryologists looked for the molecules that would induce the amphibian neural tube. The search was beyond the technology of that time, and most embryologists moved away from such deep problems. Waddington, however, came to the view that the answers to
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A remarkable number of his contemporary colleagues in Edinburgh became Fellows of the Royal Society during his time there, or shortly thereafter. Waddington was an old-fashioned intellectual who lived in both the arts and science milieus of the 1950s and wrote widely. His 1969 book
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Ritterbush, P. C. (1970). "Behind Appearance. A Study of the Relations between Painting and the Natural Sciences in This Century. C. H. Waddington. M.I.T. Press, Cambridge, Mass., 1969. xii, 258 pp., illus. (71 color plates and 136 black-and-white illustrations)".
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wing. In a period of great creativity at the end of the 1930s, he also discovered mutations that affected cell phenotypes and wrote his first textbook of "developmental epigenetics", a term that then meant the external manifestation of genetic activity.
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that Lamarck did not invent the idea of the inheritance of acquired characteristics. But, whether historically correct or not, we are stuck today with the term 'Lamarckian' for inheritance of a characteristic acquired through an environmental influence.
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modulates development. Among other metaphors, Waddington asks us to imagine a number of marbles rolling down a hill. The marbles will sample the grooves on the slope, and come to rest at the lowest points. These points represent the eventual
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Waddington was very active in advancing biology as a discipline. He contributed to a book on the role of the sciences in times of war, and helped set up several professional bodies representing biology as a discipline.
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Waddington co-founded The Institute for Advanced Studies in the Humanities at the University of Edinburgh in 1969 with Professor John MacQueen, Professor of Scottish Literature and Oral Tradition.
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mechanism that allows certain acquired characteristic to become heritable. According to Navis, (2007) "Waddington focused his genetic assimilation work on the crossveinless trait of
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which would allow an animal's response to an environmental stress to become a fixed part of its developmental repertoire, and then went on to show that the mechanism would work.
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laboratory in California, even though this was a time when most embryologists felt that genes were unimportant and just played a role in minor phenomena such as eye colour.
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In the late 1930s, Waddington produced formal models about how gene regulatory products could generate developmental phenomena, showed how the mechanisms underpinning
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Huang, Sui (2012). "The molecular and mathematical basis of Waddington's epigenetic landscape: A framework for post-Darwinian biology?".
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His family moved to India and until nearly three years of age, Waddington lived in India, where his father worked on a tea estate in the
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inheritance, the acquisition of inherited characteristics through the effects of the environment during an organism's lifetime.
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inheritance. They denied that genetic assimilation had taken place, and asserted that Waddington had simply observed the
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Induction and the Origin of Developmental Genetics - works by Salome Gluecksohn-Schoenhimer and Conrad Hal Waddington
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disagree, writing that "genetic assimilation, looks, but is not Lamarckian. It is a special case of the evolution of
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Waddington was married twice. His first marriage produced a son, C. Jake Waddington, professor of physics at the
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Behind Appearance : A Study in the Relationship Between Painting and the Natural Sciences in this Century
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development could be studied through a systematic analysis of mutations that affected the development of the
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Allen, Matthew. 2015. "Compelled by the Diagram: Thinking through C. H. Waddington’s Epigenetic Landscape."
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in Middletown, Connecticut. His personal papers are largely kept at the University of Edinburgh library.
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Yoxen, Edward. 1986. "Form and Strategy in Biology: Reflections on the Career of C. H. Waddington." In
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Behind Appearance; a Study of the Relations Between Painting and the Natural Sciences in This Century
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Wilkins, Adam S (2015). "Waddington's Unfinished Critique of Neo-Darwinian Genetics: Then and Now".
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Waddington, C. H. (1942). Canalization of development and the inheritance of acquired characters.
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before him) that evolution mainly occurred through mutations that affected developmental anatomy.
2249: 1932: 1798: 1755: 1538: 745: 603:." From these lessons in metaphysics, Waddington first gained an appreciation for interconnected 2510: 2494: 2290: 2285: 2194: 1976: 1940: 1891: 1842: 1790: 1715: 1690: 1686:
A Conceptual History of Modern Embryology: Volume 7: A Conceptual History of Modern Embryology
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of genetic variants that already existed in the study population. Other biologists such as
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Pages from a photograph album, given to Waddington by his colleagues on his 50th birthday.
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Robertson, Alan. 1977. "Conrad Hal Waddington. 8 November 1905–26 September 1975."
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Robertson, Alan. 1977. "Conrad Hal Waddington. 8 November 1905 – 26 September 1975."
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Robertson, Alan. 1977. "Conrad Hal Waddington. 8 November 1905 – 26 September 1975."
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despite variation in genotype or environment. He also identified a mechanism called
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Robertson, Alan. 1977. Conrad Hal Waddington. 8 November 1905 – 26 September 1975.
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This was the essence of the approach that won the 1995 Nobel prize in medicine for
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and the medal of the British Society for Developmental Biology is named after him.
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Biological Organisation Cellular and Subcellular : Proceedings of a Symposium
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Conrad Waddington, known as "Wad" to his friends and "Con" to family, was born in
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considered that Waddington was using genetic assimilation to support so-called
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The Life Organic: the Theoretical Biology Club and the Roots of Epigenetics
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systems. Waddington reflected that this early education prepared him for
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Supplement, Historical Register of the University of Cambridge, 1921-30
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both thought that Waddington was using genetic assimilation to support
813: 600: 577: 565: 298: 1936: 1202:––– (1952). Selection of the Genetic Basis for an Acquired Character. 2129: 1248:. Ed. John B. Cobb and David R. Griffin. University Press of America. 631: 588: 549: 359: 1244:––– (1977).(published posthumously). Whitehead and Modern Science. 881:
Waddington's theory of genetic assimilation was controversial. The
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In 1972, Waddington founded the Centre for Human Ecology in the
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to Hal and Mary Ellen (Warner) Waddington, on 8 November 1905.
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and development of living things. He also studied philosophy.
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Presidents of the International Union of Biological Sciences
1911:"Conrad Hal Waddington, 8 November 1905 - 26 September 1975" 1223:––– (1956). Genetic assimilation of the bithorax phenotype. 1155:
O.R. in World War 2: Operational Research Against the U-Boat
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and became scientific advisor to the Commander in Chief of
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based on his research. Reviewing the debate in 2015, the
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Evolution and Consciousness: Human Systems in Transition
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Mind in Nature: The Interface of Science and Philosophy
556:, as well as left-wing political leanings. In his book 1346:, Cambridge: Cambridge University Press, 1932, p. 63. 727:, with whom he had two daughters, the anthropologist 1915:
Biographical Memoirs of Fellows of the Royal Society
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Biographical Memoirs of Fellows of the Royal Society
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Biographical Memoirs of Fellows of the Royal Society
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Biographical Memoirs of Fellows of the Royal Society
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Biographical Memoirs of Fellows of the Royal Society
486:(8 November 1905 – 26 September 1975) was a British 2548: 2477: 2405: 2359: 2352: 2316: 2268: 2232: 2165: 2102: 454: 428: 410: 390: 375: 365: 343: 333: 310: 291: 257: 101:. Unsourced material may be challenged and removed. 1386:"Good Fellowship - The Repository - Royal Society" 1997:. British Society for Developmental Biology. 2015 1821:"Conrad Waddington and the origin of epigenetics" 817:his discussions on evolution—he emphasised (like 772:, the ability of an organism to produce the same 1635:Navis, Adam R. (2007). "Conrad Hal Waddington". 1131:, Edinburgh: Edinburgh University Press (1971-3 1112:. 4 vols. Edinburgh: Edinburgh University Press. 744:embryology lay in genetics, and in 1935 went to 719:, but ended in 1936. He then married architect 1733: 1731: 1237:––– (1974). A Catastrophe Theory of Evolution. 1218:Symposia of the Society of Experimental Biology 921: 560:(1941), he touched on political topics such as 1511:(18) (published 21 September 2004): R740–742. 1318:. Ithaca, NY: Cornell University Press. Pg. 2. 1038:. Cambridge : Cambridge University Press. 859:Waddington proposed an evolutionary process, " 2671:Commanders of the Order of the British Empire 2075: 1188:--- (1946). Human Ideals and Human Progress. 1096:Principles of Development and Differentiation 8: 548:Waddington had wide interests that included 1084:. London : George, Allen, & Unwin. 707:He died in Edinburgh on 26 September 1975. 611:'s philosophy in the 1920s and 30s and the 64:Learn how and when to remove these messages 2666:Alumni of Sidney Sussex College, Cambridge 2356: 2082: 2068: 2060: 1239:Annals of the New York Academy of Sciences 280: 254: 2636:Fellows of the Royal Society of Edinburgh 1926: 1885: 1836: 1712:Evolution : a developmental approach 1524: 1066:. London : George Allen & Unwin. 1052:. London : George Allen & Unwin. 1045:. London : George Allen & Unwin. 1024:. London : George Allen & Unwin. 1003:. London : George Alien & Unwin. 241:Learn how and when to remove this message 223:Learn how and when to remove this message 161:Learn how and when to remove this message 2606:Academics of the University of Edinburgh 2045:NAHSTE Project Record of C.H. Waddington 1089:New Patterns in Genetics and Development 1010:. Cambridge: Cambridge University Press. 186:This article includes a list of general 1442:Oxford Dictionary of National Biography 1284: 1258: 1216:––– (1953). Epigenetics and evolution. 955: 696:as Professor of Animal Genetics at the 568:as a "profound scientific philosophy". 2626:Fellows of Christ's College, Cambridge 2572:Index of evolutionary biology articles 1091:. New York: Columbia University Press. 1603:Goldberg, A. D., Allis, C. D., & 1450:10.1093/odnb/9780198614128.013.112261 1167:––– (1977) (published posthumously). 768:Waddington introduced the concept of 7: 692:After the war, in 1947, he replaced 99:adding citations to reliable sources 2646:People educated at Clifton College 2382:Evolutionary developmental biology 1230:––– (1961). Genetic assimilation. 1075:Darwinism and the Study of Society 1001:An Introduction to Modern Genetics 812:types. Waddington coined the term 516:evolutionary developmental biology 192:it lacks sufficient corresponding 25: 2023:from the original on 4 March 2020 638:until 1942. His friends included 45:This article has multiple issues. 2339:Evolution of sexual reproduction 1316:The Evolution of an Evolutionist 979: 970: 958: 628:Sidney Sussex College, Cambridge 177: 75: 34: 2621:Extended evolutionary synthesis 2616:20th-century British zoologists 1866:Journal of Experimental Zoology 1825:Journal of Experimental Biology 910:extended evolutionary synthesis 877:Neo-Darwinism versus Lamarckism 630:. He took the Natural Sciences 86:needs additional citations for 53:or discuss these issues on the 2651:British operations researchers 2110:Genotype–phenotype distinction 1413:. Wesleyan.edu. Archived from 1098:. New York: Macmillan Company. 1: 2367:Regulation of gene expression 1110:Towards a Theoretical Biology 1071:The Human Evolutionary System 506:who laid the foundations for 286:Conrad Hal Waddington in 1934 2641:Fellows of the Royal Society 2537:Endless Forms Most Beautiful 2317:Evolution of genetic systems 2125:Gene–environment correlation 2120:Gene–environment interaction 1973:10.1126/science.169.3947.751 1073:. In: Michael Banton (Ed.), 721:Margaret Justin Blanco White 2631:Wesleyan University faculty 2516:Christiane Nüsslein-Volhard 2019:. University of Edinburgh. 1637:Embryo Project Encyclopedia 1498:"Edward B. Lewis 1918-2004" 1436:Darling, Elizabeth (2019). 1267:Christiane Nüsslein-Volhard 1017:, George Allen & Unwin. 662:. His interests began with 2697: 2392:Hedgehog signaling pathway 2269:Developmental architecture 1683:Gilbert, Scott F. (2013). 1658:Peterson, Erik L. (2016). 1585:. Centre for Human Ecology 1559:. iBiology. Archived from 999:Waddington, C. H. (1939). 852: 731:(1943–) and mathematician 2569: 2219:Transgressive segregation 1752:10.1162/biot.2008.3.3.224 1689:. Springer. p. 205. 1526:10.1016/j.cub.2004.09.007 1496:Winchester, Guil (2004). 1059:. London: Pergamon Press. 1050:The Strategy of the Genes 723:, daughter of the writer 619:and others in the 1940s. 464: 403: 279: 2611:British palaeontologists 1909:Robertson, Alan (1977). 1710:Arthur, Wallace (2011). 1314:Waddington, C. H. 1975. 1171:. London: Jonathan Cape. 1043:Principles of Embryology 1036:The Epigenetics of birds 694:Francis Albert Eley Crew 449:Centre for Human Ecology 2397:Notch signaling pathway 2372:Gene regulatory network 2255:Dual inheritance theory 1373:A History of Embryology 1362:, 575-622. Pp. 579-580. 1298:, 575-622. pp. 575-76. 1157:. London: Elek Science. 1144:The Development of Mind 1029:The Scientific Attitude 1013:––– and others (1942). 883:evolutionary biologists 785:University of Edinburgh 717:University of Minnesota 698:University of Edinburgh 558:The Scientific Attitude 531:evolutionary biologists 521:Although his theory of 488:developmental biologist 437:University of Cambridge 433:University of Edinburgh 338:University of Cambridge 207:more precise citations. 2661:Theoretical biologists 2445:cis-regulatory element 2353:Control of development 2233:Non-genetic influences 2199:evolutionary landscape 1995:"The Waddington Medal" 1928:10.1098/rsbm.1977.0022 1860:Hall, Brian K (2004). 1787:10.1002/bies.201100031 1390:blogs.royalsociety.org 1304:10.1098/rsbm.1977.0022 1008:Organisers & Genes 926: 850: 799:is a metaphor for how 609:Alfred North Whitehead 2556:Nature versus nurture 2460:Cell surface receptor 2377:Evo-devo gene toolkit 2276:Developmental biology 2214:Polygenic inheritance 2140:Quantitative genetics 1121:Longuet-Higgins, H.C. 944:developmental biology 906:phenotypic plasticity 886:Theodosius Dobzhansky 832: 535:Theodosius Dobzhansky 529:explanation, leading 471:Conrad Hal Waddington 415:Developmental biology 259:Conrad Hal Waddington 2465:Transcription factor 2180:Genetic assimilation 2167:Genetic architecture 1232:Advances in Genetics 1108:–––, ed. (1968–72). 1077:. London: Tavistock. 919:writes however that 861:genetic assimilation 855:Genetic assimilation 835:genetic assimilation 825:Genetic assimilation 797:epigenetic landscape 791:Epigenetic landscape 778:genetic assimilation 746:Thomas Hunt Morgan's 679:operational research 666:but moved on to the 523:genetic assimilation 356:genetic assimilation 348:Epigenetic landscape 95:improve this article 2596:People from Evesham 2561:Morphogenetic field 2478:Influential figures 1887:10.1002/jez.b.20002 1878:2004JEZ...302....5H 1583:"Conrad Waddington" 1517:2004CBio...14.R740W 1185:150 (3811):563–565. 1022:How Animals Develop 843:Darwinian evolution 702:Wesleyan University 689:from 1944 to 1945. 677:he was involved in 660:John Desmond Bernal 445:Wesleyan University 370:Justin Blanco White 2656:Systems biologists 2250:Genomic imprinting 1838:10.1242/jeb.120071 1563:on 13 October 2016 1334:, 575-622. Pg 577. 1129:The Nature of Mind 1082:The Nature of Life 1064:The Ethical Animal 1015:Science and Ethics 851: 599:" and the "Arabic 110:"C. H. Waddington" 2578: 2577: 2511:Eric F. Wieschaus 2473: 2472: 2291:Pattern formation 2195:Fitness landscape 1967:(3947): 751–752. 1740:Biological Theory 1696:978-1-4615-6823-0 1486:, 575-622. P. 578 1459:978-0-19-861412-8 1271:Eric F. Wieschaus 1199:169 (4288):27-28. 1169:Tools for Thought 1164:. Addison-Wesley. 1101:––– (1970). 72). 948:pre-molecular age 914:systems biologist 898:natural selection 729:Caroline Humphrey 468: 467: 455:Doctoral students 405:Scientific career 381:Caroline Humphrey 314:26 September 1975 251: 250: 243: 233: 232: 225: 171: 170: 163: 145: 68: 27:British biologist 16:(Redirected from 2688: 2521:William McGinnis 2490:Richard Lewontin 2485:C. H. 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Waddington 15: 12: 11: 5: 2694: 2692: 2684: 2683: 2678: 2673: 2668: 2663: 2658: 2653: 2648: 2643: 2638: 2633: 2628: 2623: 2618: 2613: 2608: 2603: 2598: 2593: 2583: 2582: 2576: 2575: 2570: 2567: 2566: 2564: 2563: 2558: 2552: 2550: 2546: 2545: 2543: 2542: 2541: 2540: 2528: 2523: 2518: 2513: 2508: 2507: 2506: 2495:François Jacob 2492: 2487: 2481: 2479: 2475: 2474: 2471: 2470: 2468: 2467: 2462: 2457: 2452: 2447: 2442: 2437: 2432: 2431: 2430: 2420: 2415: 2409: 2407: 2403: 2402: 2400: 2399: 2394: 2389: 2384: 2379: 2374: 2369: 2363: 2361: 2354: 2350: 2349: 2347: 2346: 2341: 2336: 2331: 2326: 2320: 2318: 2314: 2313: 2311: 2310: 2305: 2300: 2295: 2294: 2293: 2288: 2278: 2272: 2270: 2266: 2265: 2263: 2262: 2257: 2252: 2247: 2242: 2236: 2234: 2230: 2229: 2227: 2226: 2224:Sequence space 2221: 2216: 2211: 2206: 2201: 2192: 2187: 2182: 2177: 2171: 2169: 2163: 2162: 2160: 2159: 2154: 2153: 2152: 2142: 2137: 2132: 2127: 2122: 2117: 2112: 2106: 2104: 2100: 2099: 2089: 2087: 2086: 2079: 2072: 2064: 2058: 2057: 2052: 2047: 2040: 2039:External links 2037: 2035: 2034: 2008: 1986: 1950: 1901: 1852: 1831:(6): 816–818. 1808: 1781:(2): 149–157. 1765: 1746:(3): 224–232. 1727: 1721:978-1405186582 1720: 1702: 1695: 1675: 1668: 1650: 1627: 1609: 1596: 1574: 1548: 1488: 1472: 1458: 1428: 1402: 1377: 1364: 1348: 1336: 1320: 1307: 1283: 1281: 1278: 1275: 1274: 1257: 1256: 1254: 1251: 1250: 1249: 1242: 1235: 1228: 1221: 1214: 1207: 1200: 1193: 1186: 1177: 1174: 1173: 1172: 1165: 1158: 1151: 1140: 1135:in Edinburgh, 1113: 1106: 1099: 1092: 1085: 1078: 1067: 1060: 1053: 1046: 1039: 1032: 1025: 1018: 1011: 1004: 995: 992: 990: 989:Selected works 987: 986: 985: 978: 976: 969: 967: 964: 957: 930: 927: 902:Wallace Arthur 878: 875: 853:Main article: 847:Baldwin effect 826: 823: 792: 789: 740: 737: 712: 709: 656:Joseph Needham 644:Walter Gropius 617:Norbert Wiener 593:E. 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Holmyard 573: 570: 564:, and praised 492:paleontologist 466: 465: 462: 461: 459:Robert Edwards 456: 452: 451: 430: 426: 425: 412: 408: 407: 401: 400: 392: 388: 387: 377: 373: 372: 367: 363: 362: 345: 344:Known for 341: 340: 335: 331: 330: 324: 322:(aged 69) 312: 308: 307: 303:Worcestershire 297: 293: 289: 288: 285: 277: 276: 261: 258: 249: 248: 231: 230: 185: 183: 176: 169: 168: 83: 81: 74: 69: 43: 42: 40: 33: 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 2693: 2682: 2681:Reeves family 2679: 2677: 2674: 2672: 2669: 2667: 2664: 2662: 2659: 2657: 2654: 2652: 2649: 2647: 2644: 2642: 2639: 2637: 2634: 2632: 2629: 2627: 2624: 2622: 2619: 2617: 2614: 2612: 2609: 2607: 2604: 2602: 2599: 2597: 2594: 2592: 2589: 2588: 2586: 2573: 2568: 2562: 2559: 2557: 2554: 2553: 2551: 2547: 2539: 2538: 2534: 2533: 2532: 2529: 2527: 2524: 2522: 2519: 2517: 2514: 2512: 2509: 2505: 2502: 2501: 2500: 2499:Jacques Monod 2496: 2493: 2491: 2488: 2486: 2483: 2482: 2480: 2476: 2466: 2463: 2461: 2458: 2456: 2453: 2451: 2448: 2446: 2443: 2441: 2438: 2436: 2433: 2429: 2426: 2425: 2424: 2421: 2419: 2416: 2414: 2413:Homeotic gene 2411: 2410: 2408: 2404: 2398: 2395: 2393: 2390: 2388: 2385: 2383: 2380: 2378: 2375: 2373: 2370: 2368: 2365: 2364: 2362: 2358: 2355: 2351: 2345: 2342: 2340: 2337: 2335: 2332: 2330: 2327: 2325: 2322: 2321: 2319: 2315: 2309: 2306: 2304: 2301: 2299: 2296: 2292: 2289: 2287: 2284: 2283: 2282: 2281:Morphogenesis 2279: 2277: 2274: 2273: 2271: 2267: 2261: 2258: 2256: 2253: 2251: 2248: 2246: 2243: 2241: 2238: 2237: 2235: 2231: 2225: 2222: 2220: 2217: 2215: 2212: 2210: 2207: 2205: 2202: 2200: 2196: 2193: 2191: 2188: 2186: 2183: 2181: 2178: 2176: 2173: 2172: 2170: 2168: 2164: 2158: 2155: 2151: 2148: 2147: 2146: 2143: 2141: 2138: 2136: 2133: 2131: 2128: 2126: 2123: 2121: 2118: 2116: 2115:Reaction norm 2113: 2111: 2108: 2107: 2105: 2101: 2097: 2093: 2085: 2080: 2078: 2073: 2071: 2066: 2065: 2062: 2056: 2053: 2051: 2048: 2046: 2043: 2042: 2038: 2022: 2018: 2012: 2009: 1996: 1990: 1987: 1982: 1978: 1974: 1970: 1966: 1962: 1954: 1951: 1946: 1942: 1938: 1934: 1929: 1924: 1920: 1916: 1912: 1905: 1902: 1897: 1893: 1888: 1883: 1879: 1875: 1871: 1867: 1863: 1856: 1853: 1848: 1844: 1839: 1834: 1830: 1826: 1822: 1818: 1812: 1809: 1804: 1800: 1796: 1792: 1788: 1784: 1780: 1776: 1769: 1766: 1761: 1757: 1753: 1749: 1745: 1741: 1734: 1732: 1728: 1723: 1717: 1713: 1706: 1703: 1698: 1692: 1688: 1687: 1679: 1676: 1671: 1669:9780822944669 1665: 1661: 1654: 1651: 1646: 1642: 1638: 1631: 1628: 1625: 1623: 1620: 1613: 1610: 1606: 1605:Bernstein, E. 1600: 1597: 1584: 1578: 1575: 1562: 1558: 1552: 1549: 1544: 1540: 1536: 1532: 1527: 1522: 1518: 1514: 1510: 1506: 1499: 1492: 1489: 1485: 1482: 1476: 1473: 1461: 1455: 1451: 1447: 1443: 1439: 1432: 1429: 1416: 1412: 1406: 1403: 1391: 1387: 1381: 1378: 1374: 1368: 1365: 1361: 1358: 1352: 1349: 1345: 1340: 1337: 1333: 1330: 1324: 1321: 1317: 1311: 1308: 1305: 1301: 1297: 1294: 1288: 1285: 1279: 1272: 1268: 1262: 1259: 1252: 1247: 1243: 1240: 1236: 1233: 1229: 1226: 1222: 1219: 1215: 1212: 1208: 1205: 1201: 1198: 1194: 1192:August:29-36. 1191: 1187: 1184: 1180: 1179: 1175: 1170: 1166: 1163: 1159: 1156: 1152: 1149: 1145: 1141: 1138: 1134: 1130: 1126: 1122: 1118: 1114: 1111: 1107: 1104: 1100: 1097: 1093: 1090: 1086: 1083: 1079: 1076: 1072: 1068: 1065: 1061: 1058: 1054: 1051: 1047: 1044: 1040: 1037: 1033: 1030: 1026: 1023: 1019: 1016: 1012: 1009: 1005: 1002: 998: 997: 993: 988: 982: 977: 973: 968: 961: 956: 954: 951: 949: 945: 941: 935: 928: 925: 920: 918: 915: 911: 907: 903: 899: 895: 891: 887: 884: 876: 874: 872: 871: 866: 862: 856: 848: 844: 840: 836: 833:Waddington's 831: 824: 822: 820: 819:Ernst Haeckel 815: 811: 807: 802: 798: 795:Waddington's 790: 788: 786: 781: 779: 775: 771: 766: 763: 762: 757: 752: 750: 747: 738: 736: 734: 730: 726: 722: 718: 710: 708: 705: 703: 699: 695: 690: 688: 684: 680: 676: 671: 669: 665: 664:palaeontology 661: 657: 653: 649: 645: 641: 637: 633: 629: 625: 620: 618: 614: 610: 606: 602: 598: 594: 590: 586: 581: 579: 571: 569: 567: 563: 559: 555: 551: 546: 544: 540: 536: 532: 528: 524: 519: 517: 513: 509: 505: 501: 497: 493: 489: 484: 480: 476: 472: 463: 460: 457: 453: 450: 446: 442: 438: 434: 431: 427: 424: 420: 416: 413: 409: 406: 402: 396: 393: 389: 386: 382: 379:3, including 378: 374: 371: 368: 364: 361: 357: 353: 349: 346: 342: 339: 336: 332: 327: 313: 309: 304: 300: 294: 290: 283: 278: 273: 269: 265: 256: 253: 245: 242: 227: 224: 216: 213:December 2022 206: 202: 196: 195: 189: 184: 175: 174: 165: 162: 154: 151:December 2022 143: 140: 136: 133: 129: 126: 122: 119: 115: 112: –  111: 107: 106:Find sources: 100: 96: 90: 89: 84:This article 82: 78: 73: 72: 67: 65: 58: 57: 52: 51: 46: 41: 32: 31: 19: 2535: 2484: 2428:eyeless gene 2324:Evolvability 2298:Segmentation 2175:Canalisation 2145:Heterochrony 2135:Heritability 2103:Key concepts 2025:. 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Snow 601:Alchemists 543:Lamarckian 539:Ernst Mayr 533:including 496:geneticist 328:, Scotland 318:1975-09-27 188:references 121:newspapers 50:improve it 2455:Morphogen 2440:Engrailed 2423:Pax genes 2344:Tinkering 2190:Epistasis 2185:Dominance 2096:phenotype 2001:7 October 1981:0036-8075 1775:BioEssays 1645:1940-5030 1225:Evolution 1211:Evolution 1117:Kenny, A. 865:Darwinian 774:phenotype 739:Evolution 735:(1945–). 681:with the 527:Darwinian 326:Edinburgh 305:, England 56:talk page 2418:Hox gene 2406:Elements 2387:Homeobox 2021:Archived 1945:11615737 1896:14760651 1847:25788723 1819:(2015). 1803:19632484 1795:22102361 1760:84217300 1543:32648995 1535:15380080 1395:23 April 1227:10:1–13. 1127:(1972). 863:", as a 668:heredity 605:holistic 597:Gnostics 554:painting 419:genetics 376:Children 2549:Debates 2360:Systems 2286:Eyespot 2150:Neoteny 2027:24 July 1961:Science 1874:Bibcode 1513:Bibcode 1421:4 April 946:of the 814:chreode 673:During 578:Evesham 566:Marxism 316: ( 299:Evesham 201:improve 135:scholar 2450:Ligand 2130:Operon 1979:  1943:  1937:769627 1935:  1894:  1845:  1801:  1793:  1758:  1718:  1693:  1666:  1643:  1541:  1533:  1456:  1204:Nature 1197:Nature 1183:Nature 1176:Papers 1148:online 1137:online 810:tissue 711:Family 658:, and 632:Tripos 589:Kerala 550:poetry 525:had a 514:, and 411:Fields 398:(1960) 391:Awards 366:Spouse 360:creode 190:, but 137:  130:  123:  116:  108:  1933:JSTOR 1799:S2CID 1756:S2CID 1539:S2CID 1501:(PDF) 1253:Notes 1115:–––, 994:Books 481: 477: 270: 266: 142:JSTOR 128:books 2090:The 2029:2020 2003:2019 1977:ISSN 1941:PMID 1892:PMID 1843:PMID 1791:PMID 1716:ISBN 1691:ISBN 1664:ISBN 1641:ISSN 1591:2016 1569:2016 1531:PMID 1467:2024 1454:ISBN 1423:2010 1397:2018 1269:and 888:and 806:cell 626:and 572:Life 552:and 537:and 502:and 483:FRSE 383:and 311:Died 292:Born 272:FRSE 114:news 2094:of 1969:doi 1965:169 1923:doi 1882:doi 1870:302 1833:doi 1829:218 1783:doi 1748:doi 1521:doi 1446:doi 1300:doi 615:of 587:of 479:FRS 475:CBE 268:FRS 264:CBE 97:by 2587:: 2497:+ 1975:. 1963:. 1939:. 1931:. 1919:23 1917:. 1913:. 1890:. 1880:. 1868:. 1864:. 1841:. 1827:. 1823:. 1797:. 1789:. 1779:34 1777:. 1754:. 1742:. 1730:^ 1639:. 1537:. 1529:. 1519:. 1509:14 1507:. 1503:. 1484:23 1452:. 1444:. 1440:. 1388:. 1360:23 1332:23 1296:23 1123:, 1119:, 841:, 787:. 654:, 650:, 646:, 642:, 518:. 510:, 498:, 494:, 490:, 439:, 421:, 417:, 358:, 354:, 350:, 301:, 59:. 2197:/ 2083:e 2076:t 2069:v 2031:. 2005:. 1983:. 1971:: 1947:. 1925:: 1898:. 1884:: 1876:: 1849:. 1835:: 1805:. 1785:: 1762:. 1750:: 1744:3 1724:. 1699:. 1672:. 1647:. 1624:. 1622:4 1593:. 1571:. 1545:. 1523:: 1515:: 1469:. 1448:: 1425:. 1399:. 1302:: 1150:) 1139:) 320:) 244:) 238:( 226:) 220:( 215:) 211:( 197:. 164:) 158:( 153:) 149:( 139:· 132:· 125:· 118:· 91:. 66:) 62:( 20:)

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