reading5inversionsindrosophila

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Unformatted text preview: Proc. Natl. Acad. Sci. USA Vol. 88, pp. 10367—10371, November 1991 Population Biology Four decades of inversion polymorphism in Drosophila pseudoobscura (evolution / genetic change / chromosome inversions) WYATT W. ANDERSONa, JONATHAN ARNOLDa, DAVID G. BALDWINb, ANDREW T. BECKENBACHC, CELESTE J. BROWNa, STEPHEN H. BRYANTd, JERRY A. COYNEe, LAWRENCE G. HARSHMAN f 1 WILLIAM B. HEEDb,_DUANE E. JEFFERYg, LOUIS B. KLACZKOh, BETTY C. MOORE‘, JEAN M. PORTERa, JEFFREY R. POWELLJ, TIMOTHY PROUTf, STEPHEN W. SCHAEFFERk, J. CLAIBORNE STEPHENs', CHARLES E. TAYLOR‘“, MONTE E. TURNER“, GABRIEL O. WILLIAMSg, AND JOHN A. MOORE‘° aDepartment of Genetics, University of Georgia, Athens, GA 30602; bDepartment of Ecology and Evolution, University of Arizona, Tucson, AZ 85721; cDepartment of Biological Sciences, Simon Fraser University, Burnaby, B.C. VSA 1S6, Canada; dDepartment of Biology, California Polytechnic University, Pomona, CA 91768; eDepartment of Ecology and Evolution, The University of Chicago, Chicago, IL 60637; fDepartment of Genetics, University of California, Davis, CA 95616; I3Department of Zoology, Brigham Young University, Provo, UT 84602; lDepartment of Biology, University of California, Riverside, CA 92521; JDepartment of Biology, Yale University, New Haven CT 06511; mDepartment of Biology, University of California, Los Angeles, CA 90024; nDepartment of Biology, University of Akron, Akron, OH 44325; hDepartamento de Genetica, Universidade Estadual de Campinas, Campinas, Brazil; kDepartment of Biology, Pennsylvania State University, University Park, PA 16802; and 1Laboratory of Viral Carcinogenesis, National Cancer Institute, Frederick, MD 21702 Contributed by John A. Moore, August 14, 1991 ABSTRACT We report data that continue the studies of Dobzhansky and others on the frequencies of third-chromo- some inversions in natural populations of Drosophila pseudoob- scum in North America. The common gene arrangements continue to be present in frequencies similar to those described four decades ago, and the broad geographic patterns also remain unchanged. There is only one pronounced trend over time: the increase in frequency of the Tree Line inversion in Pacific coast populations. For more than 40 years Dobzhansky and his associates have monitored the frequencies of third-chromosome inversions in natural populations ofDrosophila pseudoobscura (1—5). This species, like many others in the genus, has an extraordinary amount Of chromosome polymorphism. D. pseudoobscura and its sibling species Drosophila persimilis are polymorphic for several dozen third-chromosome inversions (with only one in common) and are fixed for single-inversion differences on the X, second, and fourth chromosomes. Since polymor- phism must be a transient stage in the establishment of interspecific differences, long-term monitoring of gene ar— rangement frequencies may be useful in understanding this process. In addition, such surveys offer the possibility of documenting balancing selection as well as stasis or micro- evolution over long periods of human observation. The third-chromosome inversions of D. pseudoobscura are identified by differences in the banding patterns of polytene chromosomes in the larval salivary glands. Molecular data suggest that each inversion has arisen only once and that the polymorphism is at least 1 million years Old (6). Because the inversions are overlapping and show very little recombina- tion in heterokaryotypes, they form semi-isolated gene pools and, hence, have become genetically differentiated (6). The inversions also differ in fitness, as documented by laboratory experiments and observations in nature of seasonal cyling and differential mating success of males (1, 7—9). However, the ecological interactions that maintain the polymorphisms in nature are unknown. Almost all of the data on inversion frequencies in this species have come from Dobzhansky’s studies Of seasonal, annual, and geographic variation. Many localities (mainly in the western United States) were monitored at roughly 10-year The publication costs of this article were defrayed in part by page charge payment. This article must therefore be hereby marked “advertisement” in accordance with 18 U.S.C. §1734 solely to indicate this fact. 10367 intervals between 1936 and 1973, and a three-decade over- view was published in 1975 (4). We report here the results of the latest survey, conducted in the early 1980s and covering 48 localities in the northern part of the species range. Our methods were the same as those empldyed in previous studies: freshly collected females were allowed to oviposit on medium, and the salivary gland polytene chromosomes were analyzed in one to eight larvae from each female. Occasion— ally, wild—caught males were mated to tester females and chromosomes were analyzed in a sample of larvae (4). Table 1 gives the data from our survey, along with a summary of previous observations from these localities, which are de- scribed in previous surveys (1—4). Eight gene arrangements reach a frequency of at least 10% in some localities: ST, AR, CH, PP, TL, SC, 0L, and EP. Inversions OL, SC, and EP are rare in many localities but consistently appear in large samples. Eighteen rare arrangements have been described, and two more are undescribed, giving a total Of 28 inversions segregating in North America. Fig. 1 shows the diversity of gene arrangements within localities, given as a contour map, whereas Fig. 2 summarizes the common inversions in each population. The most important conclusion to be drawn from these data is that during half a century the common inversions have continued to segregate within populations at fairly similar frequencies. The general geographic pattern has also re- mained similar. ST is in highest frequency in Pacific Coast populations, especially west of the Sierra Nevada, where AR, CH, and TL are also common. AR becomes the predominant inversion between the Sierra Nevada and the Rocky Moun- tains and shares prominence with PP at the eastern edge of the species distribution. CH is predominant in southern California. Finally, western California has remained the region of highest diversity. Superimposed on this temporal stability are significant fluctuations in frequencies in some populations. These fluc— tuations, which have been seen in previous surveys (4), should be viewed with caution, for most populations were sampled only once in a given survey and seasonal variation has been seen in some localities (refs. 7 and 8; J.A.M., B.C.M., and C.E.T., unpublished data). This source Of error does not apply, however, to broad regional changes over time. Dobzhansky (ref. 3, p. 823) noted that the frequency of the PP Abbreviations: ST, standard; AR, arrowhead; CH, Chiricahua; PP, Pike’s Peak; TL, Tree Line; SC, Santa Cruz; 0L, Olympic; EP: Estes Park. “To whom reprint requests should be addressed. 10368 Table 1. Population Biology: Anderson et al. Proc. Natl. Acad. Sci. USA 88 (I991) Frequencies (in percent) of gene arrangements in D. pseudoobscura populations and n, the effective number of chromosomes on which they are based Locality 1. Lillooet, BC 3" Port Coquitlam, BC 3. Okanagan, BC 4. Victoria, BC 5 . Bellingham, WA 6. Methow, WA 7. Seattle, WA 9° Gaston, OR 9. Spray, OR 1981—1982 10. Kerby, OR 1981—1982 11. St. Helena, CA 1971—1972 1980—1981 12. Spieth Reserve, Davis, CA 13. Georgetown, CA 14. Placerville, Camino, and Apple Hill, CA 15. Berkeley, CA 1980—1982 1962—1964 1980—1981 16. Mather, CA 17. Tassajara Hot Springs, Santa Lucia Mountains, CA 1962—1963 18. Lone Pine Canyon, Inyo National Forest, CA 19. Death Valley National Monument, CA 19a. Wildrose Canyon, Panamint Mountains May 1980 July 1980 Year 1982 1981 1940 1964 1972 1981 1983 1980 1940 1964 1972 1981 1940 1981 1982 1940 1965 1972 1940 1965 1940 1957 1963 1980 1981 1981 1940 1957 1963 1979 1945 1957 1963 1971 1972 1974 1975 1976 1981 1940 1957 1972 1980 1938 1957 1963 1983 1937 1940 1957 1963 1968 1972 246 162 30 80 160 180 83 92 100 208 320 228 44 156 79 88 70 148 185 64 30 125 108 200 362 166 70 144 180 106 108 300 206 82 242 5221 202 308 3 16 446 390 576 746 204 222 62 104 200 640 296 212 94 78 38 95 224 360 224 132 142 96 134 124 ST 33.6 13.2 36.7 55.0 54.4 31.7 19.3 20.7 52.0 79.3 60.6 46.1 52.3 7.1 17.1 25.0 44.3 17.6 12.1 18.8 43.3 17.8 35.2 51.0 51.4 51.8 28.6 30.6 41.1 25.5 26.9 57.0 49.5 11.0 30.2 46.6 34.2 35.7 45.3 54.7 34.3 22.2 29.5 23.5 18.0 14.5 51.0 54.0 40.6 52.3 36.8 21.3 25.6 65.8 28.4 13.8 30.8 25.5 24.3 33.1 37.5 26.9 48.4 AR 20.8 13.8 46.7 30.0 31.9 36.7 10.0 19.6 47.0 17.8 34.4 34.2 11.4 12.2 27.3 56.8 27.1 29.1 46.5 53.1 30.0 44.5 20.4 19.5 14.9 12.1 17.1 13.9 10.0 33.0 26.9 15.0 24.8 9.8 19.4 9.1 6.4 35.7 33.2 22.0 33.1 37.5 32.6 30.4 38.3 25.8 20.2 23.0 18.0 11.1 9.4 56.4 51.3 13.2 54.7 67.4 44.4 58.9 40.9 45.1 38.5 39.6 30.6 CH 3.1 9.6 10.0 1.2 0.5 16.1 8.7 1.0 0.9 29.5 29.5 11.4 11.4 5.7 5.4 2.3 18.8 13.3 4.8 11.1 2.0 2.5 1.8 4.3 18.7 5.0 3.8 6.5 1.3 3.4 5.3 19.1 14.9 17.2 3.8 6.3 11.5 17.0 6.6 12.2 14.0 21.0 12.5 4.0 23.6 26.6 25.9 18.0 5.1 7.9 7.4 18.8 21.1 11.2 5.3 11.3 13.6 23.9 16.9 PP 19.7 1.8 3.3 8.8 0.6 1.7 1.2 9.8 0.9 1.3 0.6 1.1 4.5 8.6 10.8 6.1 3.3 6.2 5.0 6.1 0.6 4.3 2.8 1.7 3.8 12.0 4.9 14.6 9.9 2.8 1.5 9.8 6.7 2.8 6.3 6.4 8.3 4.5 12.9 9.0 5.2 0.3 9.0 5.3 3.2 0.9 12.9 6.3 3.1 3.7 0.8 Frequency, % TL sc OL EP 20.8 1.9 58.7 1.8 1.2 3.3 5.0 13.1 28.3 0.5 0.5 40.2 1.2 12.0 39.1 2.2 2.9 4.1 15.8 0.4 1.3 6.8 43.6 1.3 5.8 38.6 4.6 2.3 14.3 37.2 24.7 0.5 0.5 7.4 7.8 1.6 6.7 3.3 22.6 1.4 0.7 17.6 12.0 2.8 ‘ 13.5 7.5 1.5 22.9 1.1 0.3 0.8 32.5 1.2 37.1 2.9 4.3 1.4 30.6 2.1 1.4 33.3 1.1 4.4 3.3 24.5 9.4 13.9 18.5 7.4 4.3 9.0 1.3 16.5 1.0 37.8 12.2 14.6 22.7 0.8 7.8 4.8 14.0 1.4 0.7 6.2 31.2 1.0 2.5 8.4 10.4 0.7 0.3 6.3 1.6 9.9 0.4 17.2 0.3 0.5 0.3 15.1 0.5 1.4 19.2 3.2 2.6 20.1 3.9 1.5 21.6 1.4 1.8 0.4 25.8 1.9 13.5 1.0 2.0 7.5 0.5 8.9 2.2 1.1 0.5 7.4 2.0 0.3 24.1 0.9 1.9 0.9 3.2 1.1 7.7 1.3 7.9 6.3 3.1 2.7 0.4 16.7 4.2 7.3 4.5 0.7 0.7 3.2 Others 2.1 (KB) 0.9 (new) 0.04 (EB, BE) 0.6 (MA) 0.4 (MA) Population Biology: Anderson et al. Proc. Natl. Acad. Sci. USA 88 (1991) 10369 Table 1. (Continued) Frequency, % Locality Year n ST AR CH PP TL SC OL EP Others 19b. Furnace Creek 1980 334 8.7 78.4 7.8 3.0 1.2 0.6 0.3 (FC) 20. China Ranch, Tecopa, CA 1981 234 15.8 67.5 7.7 3.4 4.7 0.4 0.4 21. Santa Barbara, CA 1940 438 47.5 20.8 16.2 7.3 7.8 0.5 1963 332 56.6 15.1 7.2 7.2 11.4 2.4 1973 204 52.9 10.8 16.7 0.5 18.1 1.0 1981 312 22.4 6.4 35.3 1.0 24.7 1.0 2.6 6.7 22. Santa Cruz Island, CA 1936 42 54.8 16.7 28.6 1940 72 43.1 18.1 6.9 31.9 1963 400 62.2 12.8 9.8 1.2 4.8 7.2 2.0 1970 204 41.2 14.2 15.2 1.0 10.8 13.2 1.0 3.4 1980—1981 612 38.4 5.9 14.5 0.8 6.6 20.2 0.3 13.2 23. San Gabriel Mountains, CA 1936—1937 101 34.7 27.7 26.7 10.9 1963—1964 134 68.7 16.4 7.5 0.8 6.7 1973 72 38.9 18.1 18.1 2.8 20.8 1.4 (SJ) 1980—1981 155 26.5 3.9 52.9 3.2 12.3 1.3 24. Riverside, CA 1963 124 68.5 10.5 8 1 2.4 5 6 4.0 0.8 1980—1981 386 32.9 11.6 37.8 2.1 14.8 0.2 0.2 0.1 25. San Jacinto Mountains, CA 25a. Pinyon Flats 1939—1942 3021 40.9 26.4 28.0 4.0 0.6 1952—1956 5702 47.7 21.5 15.4 8.7 5.3 0.7 0.6 0.1 1963 604 72.7 10.9 3.5 6.3 5.8 0.8 1970 1080 65.3 20.1 4.5 2.8 6.9 0.1 0.3 1978 201 58.2 18.4 10.5 2.0 9.5 1.5 1980 146 69.9 10.3 11.6 2.7 5.5 " 25b. Keen Camp 1939—1940 4368 29.9 26.1 40.2 3.6 0.2 1948—1949 571 44.5 19.8 35.7 1955—1956 1838 33.6 25.8 32.5 5.3 2.8 1966 438 48.2 24.2 14.2 3.2 10.3 25c. Indian Mountain 1974—1975 666 44.7 17.0 22.7 2.3 8.9 0.8 0.6 0.6 25d. James Reserve 1978—1979 414 44.9 13.5 30.7 1.9 8.5 0.2 0.2 25e. Andreas Canyon 1940—1942 782 58.7 25.8 12.5 2.9 1978—1979 86 55.8 16.3 11.6 16.3 26. Anza Borrego State Park, CA 1938 132 53.8 30.3 13.6 2.3 1941 42 59.5 35.7 4.8 1966 200 69.5 16.0 3.0 2.0 9.0 0.5 1973 450 75.3 16.4 2.7 0.4 5.1 1981 223 76.7 5.8 6.3 0.5 10.3 0.5 27. Lamoille Canyon, NV 1982 139 3.4 79.3 3.4 6.9 5.5 1.4 (TX, LA) 28. Lehman Caves National Monument, 1950 100 7.0 84.0 6.0 1.0 2.0 NV 1963 318 7.5 86.5 0.6 4.1 1.3 1973 264 1.9 96.2 1.1 0.4 0.4 ' 1980 241 3.5 94.2 0.6 1.2 0.3 0.3 (new) 29. Charleston Mountains, NV 1937 256 12.1 68.8 19.1 1955 126 24.6 55.6 9.5 6.3 4.0 1963 372 19.1 69.4 2.4 7.5 1.6 1972 200 23.0 64.5 10.5 1.0 1.0 1980 174 17.8 58.6 13.8 4.0 2.9 0.6 1.7 0.6 (CM) 30. Ferron, UT 1950 110 6.4 87.3 4.5 1.8 1965 54 5.6 81.3 1.9 9.3 1.9 1980 132 94.1 3.7 2.2 (EM, FE) 31. Bryce Canyon National Park, UT 1940 100 2.0 96.0 2.0 1950 84 4.8 92.9 2.4 1957 190 2.6 93.2 1.6 2.6 1965 200 2.5 92.0 4.0 1.5 1973 136 0.7 99.3 1978 880 0.6 97.6 1.6 0.1 0.1 (BR) 32. Betatakin, AZ 1957 200 3.0 96.5 0.5 1980 191 2.0 91.3 4.1 0.5 2.0 33. Grand Canyon National Park, AZ 1940 100 1.0 98.0 1.0 1957 200 5.5 91.0 2.5 1.0 1965 200 2.5 96.5 0.5 0.5 1973 244 2.5 94.3 2.5 0.8 1980 245 0.8 98.4 0.4 0.4 34. Flagstaff, AZ 1940 100 1.0 97.0 1.0 1.0 1957 200 6.5 89.0 2.0 2.5 10370 Population Biology: Anderson et al. Proc. Natl. Acad. Sci. USA 88 (1991) Table 1. (Continued) Frequency, % Locality Year n ST AR CH PP TL SC OL EP Others 1965 200 0.5 96.0 1.5 1.5 0.5 (new) 1973 206 95.1 4.9 1980 132 95.3 4.0 0.7 35. Prescott, AZ 1940 100 11.0 79.0 9.0 1.0 1957 200 23.0 71.5 3.5 2.0 1963 412 10.4 82.3 5.1 2.2 1973 238 5.0 86.6 8.0 0.4 1980 178 4.5 91.6 3.9 36. Tempe, AZ 1981 203 0.5 90.7 8.8 37. Organ Pipe National Monument, AZ 1971 42 7.1 90.5 2.4 38. Sonoita, AZ 1941 42 7.1 59.5 33.3 1957 200 2.0 80.0 15.0 3.0 1965 200 6.5 77.5 11.5 4.0 0.5 1973 122 0.8 86.1 9.0 3.3 1980 159 4.8 78.6 12.5 4.2 0.8 (SO) 39. Chiricahua Mountains, AZ 1940 192 0.5 88.5 6.3 4.2 0.5 1957 400 0.2 85.0 11.5 2.5 0.2 0.5 (CC) 1959 200 84.0 11.5 4.0 0.5 (CC) 1964 198 1.5 88.9 6.1 3.0 0.5 1973 262 0.4 92.4 4.9 2.3 1980 275 85.1 11.4 2.8 0.5 0.3 (EM) 40. Muggins Gulch and Rist Canyon, CO 1941 64 4.7 17.2 57.8 7.8 12.5 1965 449 4.7 49.9 28.5 10.9 5.1 0.9 (CH, 0L, SC) R 1968—1970 450 6.4 34.9 29.3 14.9 9.6 4.9 (CH, OL, SC) M 1969—1970 410 3.4 34.6 37.3 15.4 6.3 2.9 (CH, OL, SC) 1980 41 4.7 46.5 16.3 25.6 2.3 4.7 41. Black Canyon of the Gunnison 1950 152 3.3 81.6 8.6 1.3 5.3 National Monument, CO 1964 182 5 .0 94.4 0.6 1981—1982 399 1.2 72.5 0.5 11.4 5.9 0.2 8.3 42. Rocky Mountain Biological Lab., CO 1970 48 4.2 83.3 6.2 4.2 2.1 1980 128 94.5 1.6 2.3 .0.8 0.8 1981 74 2.7 73.0 12.2 9.5 2.7 43. Hayden Creek, CO 1950 24 41.7 20.8 29.2 4.2 4.2 1964 180 17.2 47.2 2.2 25.0 1.7 6.7 1982 122 0.7 67.1 2.1 26.7 2.1 1.4 (HC) 44. Mesa Verde National Park, CO 1940 100 100.0 1957 200 96.5 1.5 0.5 1.5 (CC) 1964 206 1.9 97.6 0.5 1980—1981 147 98.5 1.5 45. Raton, NM 1940 100 78.0 1.0 20.0 1.0 1964 200 0.5 78.0 1.5 19.0 1.0 1980 184 72.6 3.7 22.0 0.6 1.2 (CU) 46. Capitan, Hondo, Ruidoso, and 1941 142 56.3 7.0 35.2 1.4 Lincoln, NM 1964 82 2.4 69.5 1.2 25.6 1.2 1965 200 2.5 42.0 1.0 47.5 2.0 5.0 1969 376 0.3 57.7 2.9 35.9 1.1 1.9 0.3 1970 310 1.6 68.7 3.2 25.5 0.6 0.3 1980 247 0.7 81.1 4.2 11.7 0.7 0.4 0.4 0.8 (SO, RU) 47. Marfa and Davis Mountains, TX 1939—1941 148 1.4 33.8 3.4 56.1 4.7 0.7 1964 200 15.0 1.5 82.5 1.0 1973 224 0.5 32.6 2.2 61.6 2.7 0.4 1982 288 33.0 4.6 57.8 3.0 0.7 0.3 0.7 48. Austin, TX 1939—1941 1279 20.6 71.0 6.3 1.5 0.7 1953 200 2.0 38.5 54.5 4.0 0.5 0.5 (TX) 1964 300 2.6 16.4 72.0 3.5 4.3 1.3 1982 176 21.0 0.6 66.3 5.5 0.6 5.5 0.6 (HI) See refs. 10 and 11 for discussion of n. In addition to the eight principal gene arrangements [Standard (ST), Arrowhead (AR), Chiricahua (CH), Pike’s Peak (PP), Tree Line (TL), Santa Cruz (SC), Olympic (0L), and Estes Park (EP)], others in low frequency are listed under “Others.” These include some previously described: Berkeley (BE), Cochise (CO), Cuemavaca (CU), East Bay (EB), Hidalgo (HI), Mammoth (MA), San Jacinto (SJ), Sonoita (SO), and Texas (TX). Many new arrangements were encountered and some of these have been given working names: Bryce (BR), Charleston Mountains (CM), Emory (EM), Fort Collins (FC), Ferron (FE), Hayden Creek (HC), Lamoille (LA), Ruidoso (RU). Other undiagnosed new arrangements were from St. Helena (locality 11), Lehman Caves (locality 28), and Flagstaff (locality 34). ° inversion increased in many California localities in earlier frequency of TL along the entire Pacific coast, a pattern noted surveys, but its frequency has since declined (4). The one clear in 1975 (4). As Fig. 3 indicates, this trend has continued in 13 directional trend in our data is the continuing increase in the of our 16 west coast populations. With the exception of PP, Population Biology: Anderson et al. FIG. 1. Inversion diversity measured as H = 1 — Epiz, where p, is the frequency of the ith inversion. The Mexican sites indicated are from earlier surveys and are not in Table 1. none of the other common gene arrangements has shown such a trend over the four decades of sampling. Apart from the recent change in the frequency of TL, inversions in North American populations of D. pseudoob- scura constitute many locally stable polymorphisms with pronounced differentiation across the species range. The field and laboratory evidence that the inversions differ in fitness implies that the local stability probably results from \7: 0 SA PT \ \ 1 SAT \ I . SACTE o ‘smfif i | l oSAT 7"r--4 ACT I 1 CAP FIG. 2. Inversions >10% in each site. Inversion identity is given by the first letter of the two—letter acronym except for SC, which is denoted by Sc. For Mexican sites, see Fig. 1 legend. Proc. Natl. Acad. Sci. USA 88 (1991) 10371 40 3 Near Vancouver, BC 40 16 Yosemite 20 20 I 1940 1 980 1 940 40 1 7 20 to m o 40 20 l 40 4o 21 20 20 I 40 40 20 20 W 40 1 0 40 20 20 I ll 40 11 40 24 20 20 K I 40 14 4o 25 20 20 K l 40 15 Berkeley 40 26 NearSan Diego 20 R 1940 1980 1940 1980 FIG. 3. Frequency of TL on the west coast. On each graph the ordinate is % TL and the abscissa is years with 1940 and 1980 as indicated. Site numbers from Table 1 are given. balancing selection. The relative stability is also indicated by the 1- to 2-million-year age of the system estimated from molecular analysis (6). The causes of the differentiation among populations remain a mystery. The increase in frequency of TL may be due to an envi- ronmental change favoring carriers of TL or to the occur— rence of a new favorable mutation linked to the arrangement. Further monitoring of natural populations combined with laboratory experiments may reveal the bases for stable polymorphism, geographic variation, and temporal changes in the frequency of these inversions. This work was supported by National Science Foundation Grants DEB-7968493 and BSR-8516188 to W.W.A. by a grant from the Natural Sciences and Engineering Research Council of Canada to A.T.B., and by Intramural Research Funds of the University of California, Riverside, to J.A.M. and C.E.T. Preparation of the manuscript was supported by National Institutes of Health Grant GM38462 to J.A.C. 1. Dobzhansky, T. & Sturtevant, A. (1938) Genetics 23, 28—64. 2. Dobzhansky, T. & Epling, C. (1944) Carnegie Inst. Washington Pub]. 554. 3. Lewontin, L. C., Moore, J. A., Provine, W. B. & Wallace, B. eds. (1981) Dobzhansky’s Genetics of Natural Populations I—XLIII (Columbia Univ. Press, New York). 4. Anderson, W., Dobzhansky, T., Pavlovsky, 0., Powell, J. R. & Yardley, D. (1975) Evolution 29, 24—36. Anderson, W. W. (1989) Genome 31, 239—245. Aquadro, C. F., Weaver, A. L., Schaeffer, S. W. &Anderson, W. W. (1991) Proc. Natl. Acad. Sci. USA 88, 305-309. Dobzhansky, T. (1943) Genetics 28, 162-186. Epling, C. (1953) Evolution 7, 342—365. Salceda, V. M. & Anderson, W. W. (1988) Proc. Natl. Acad. Sci. USA 85, 9870—9874. 10. Arnold, J. (1981) Biometrics 37, 495-504. 11. Arnold, J. & Norrison, M. L. (1985) Genetics 109, 785—798. 9‘5" 339°.“ ...
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