Copyright 0 1992 by the Genetics Society of America
Perspectives Anecdotal, Historical and Critical Commentaries on Genetics Edited by James F. Crow and William F. Doue
Sixty Years Ago: The 1932 International Congressof Genetics James F. Crow Genetics Department, University of Wisconsin, Madison, Wisconsin 53706
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HE Sixth International Congress of Genetics, in Ithaca, New York in August, 1932 must have been an exciting experience in those dreary, depression, dust-bowl years. Five hundredand sixty-two people came to Cornel1 from 35 countries, often at considerable financial sacrifice, for in those days participants paid their own expenses (Figure 3 (foldout)). The Congress (JONES 1932)featureddemonstrations of living organisms, charts, photographs, and hundreds of microscopes for viewing specimens, especially cytological. Exhibits included 15 groups of invertebrates,includingaphids,echinoderms, mollusks and tunicates, many with living specimens. There were 15 vertebrate species and 35 genera of plants. In addition there were an equal number of vegetable crops, flowers and fruits. The description of the exhibits took about 250 pages in the volume of abstracts available at the start of the Congress. The sheer magnitude and variety must have been overwhelming. The living plant exhibits, extending over about a hectare, attracted the most attention. These involved carefulplanning, with many of the types grown experimentally the year before to determine planting time so that demonstrations would peak in August. This was done under thesupervision of MARCUS RHOADES, who did much of the field work himself, and that same year received his Ph.D. from Cornell. (RHOADES recently died and will be the subject of an obituary in a forthcoming issue.) T h e most popular exhibit was a “living chromosomemap” inwhich mutant maize plants were arranged in positions corresponding to the locations of the mutations on the linkage map (Figure 1). The Congress set aside ample time for seeing the exhibits, including all day Sunday for those who didn’t want to see Niagara Falls. Like the currently popular poster sessions, these offered the opportunity for individual discussions. According to the Congressdescription, “Even a most retiring person will easily find an opportunity to approach a Genetics 131: 761-768 (August, 1992)
person demonstrating his exhibits, in order to ask a question or to start a discussion.” My reason for featuring this particular Congress, however, is not the exhibits, but the number of outstanding addresses that have had a lasting influence. The year 1932 was near the end of the golden era of classical genetics, the period when the tools were breeding experiments and the microscope and when the riddlesof genetic transmission were largely solved. The advances of the previous 32 years had created the new science of genetics. Before discussing the Sixth Congress, I’ll briefly mention some other early ones. T h e First antedated the rediscovery of MENDEL’Slaws. T h e “International Conference on Hybridisation and on the Cross-Breeding of Varieties” was held in London on July 11-1 2, 1899atthe instigation of the Royal Horticultural Society. T h e Congress featured a talk by WILLIAM BATESON, soon to become one of MENDEL’Searliest and strongest supporters. T h e Second “International ConferenceonPlantBreeding and Hybridization” occurred in 1902, in New York City. Four years later the Royal Horticultural Society again convenedan “International Conference onHybridisation and Plant Breeding” in London in 1906,and again BATESON was featured, this time as President.In histalk he urgedthat his new verbal construct genetics be adopted. Accordingly, the printed volume was entitled Report of the ThirdInternational Conference on Genetics. For the first time, animal experiments were included. The Fourth Congressof Genetics took place in Paris in 191 1. The Fifth was scheduled for 1916, five years later, but World War I intervened, and it was not held until 1927, in Berlin. By then genetics and geneticists hadgrown; 903 membersfrom 35 countries attended, and the Proceedings include 148 papers. The Sixth Congress, the subject of this essay, was held on schedule five years later. T h e Seventh was
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planned for 1937 in Moscow, but was postponed a year by the National Academy of the USSR and was eventually held in Edinburgh in 1939. One month before the opening the Russian delegation withdrew and 50 papers were cancelled. O n the second day of the Congress, British citizens living in Germany were advised to return home and several non-British delegations left. The Congress continued, however, with most of the members remaining.O n September 1, the day after ad-journment, fighting began on the PolishGerman border. The SS Athenia with several geneticists aboard was torpedoed in the Atlantic. B. PRICE, C. W. COTTERMAN, W. LAWRENCE and W. R. SINGLETON survived but the F. W. TINNEYS of the University of Wisconsin perished. T o return to the 1932 Congress: the President was T. H. MORGAN(Figure 2). His attendance was greeted with great relief, for he had recently been in a severe automobile wreck and his friends feared that he would not recover i n time. MORGAN'Saddress dealt mainly with history, but he listedwhat he regarded as the five mostimportant problems. They were: ( 1 ) "growth and duplication" of genes; (2) physical interpretation of synapsis and crossing over: (3) relation of genes to characters; (4) nature of themutation process; ( 5 ) application to plant and animal breeding. This reflected the state of genetics at the time. Transmission genetics was essentially solved, and it was time to understand the geneand development in a morebasic way. MORGANhad no way of knowing that the study oftiny organisms andenormous molecules would converge in the decades ahead. One of the most exciting and influential addresses was delivered by H. J. MULLER. This was not long after his discovery of X-ray mutagenesis, and he exploited this tool to the fullest. He reviewed the induc-
tion of mutations and chromosome breakage by radiation, and amongother things emphasized how the use of deletions enabled h i m to discern thedifference between gain and loss of gene function. He introduced the now-standard vocabulary of amorphic, hypomorphic, hvpermorphic. neomorphic and antimorphic mutations. MULLERalso introduced the idea of dosage compensation and its elucidation by hypomorphic mutations. Although his preferred mechanism turnedout to be wrong, the observation was remarkably astute and the generalizations from it to the meticulousness of evolution remarkably far-reaching. I have long erl-joyed MULLER'S reasoning from dosage compensation of mutant alleles to the conclusion, later abundantly confirmed, that normal alleles are not completely dominant. What a wealth of ingenuity of both concept and experimental trickerv he displayed! Those who claim MUILER as the idea man of early genetics can find ample supporting evidence here. A. H. STURTEVANT, with typical conciseness, discussed the possibility of using mosaics for elucidating developmental patterns. The newly discovered clarpt mutation in Drosophilasimulans, which produceda high rate of nondisjunction in early cleavage, greatly facilitated this analysis. I t was to be many years before the corresponding mutation was found in Drosophila melanogaster. The system was then exploited for fate mapping andneurogenetics, and SEYMOURBENZER coined the eponymous unit sturt. CURTSTERN presented his famous experiment using translocations to demonstrate the physical reality of crossing over. His diagram from this Congress has been reproduced in one textbook after another. Essentially the same experiment had been done in maize by HARRIETCREIGHTONand BARRARAMC-
Perspectives
CLINTOCK, but was published elsewhere. At this Congress they went furtherand used doubly heteromorphicchromosomes todemonstratefour-strand crossing over. This was a subject of interest at the time because of BELLING’S hypothesis of crossing over as a copying switch between newly replicated strands, which would not permit four-strand exchange. Mcname appears at many places in the proCLINTOCK’S ceedings. She gave a paper on nonhonlologous chromosome associations. She also did the cytology for another important paper, that of L. J. STADLER. STADLER, deep and thoughtful as always, argued cogently that radiation produces mainly chromosome rearrangements and few, if any, “real” genemutations. In contrast, TIMOF~EFF-RESSOVSKY discussed forward and backward mutations in great detail and believed that X-rays could indeed produce both. He ended his paper on a euphoric note: “We geneticists are in a very happy condition: our science is young, its ‘development curve’is rising rapidly and the future w i l l bring us the most interesting facts and views concerning the gene problem.”Alas, this was one year beforeHitlercame to power and TIMOF~EFF’S life changed for the worse as he was caught up i n both tragic dictatorships, Germany and USSR. R. A. EMERSONgave a thorough review of maize genetics. All 10 linkage groupshad by then been identified, some 100 mutations had been assigned to a group,andabout 50 had been reasonably well mapped (Figure 1). The assignment of linkage groups to chromosomes was done mainly by MCCLINTOCK,
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using trisomics. The trisomics, in turn, were obtained using GEORGEBEADLE’S asynaptic gene, which produced abundant triploids from which all the primary trisomics were easily derived. EMERSON also described the variegated pericarp genes, now known to be the result of a transposable element. Later R. A. BRINK made this mutant the focus of his study, and variegated pericarp became a part of the MCCLINTOCK legend. BEADIXwasjust deserting corn for Drosophila and w a s engaged i n showing, through the use of attachedX chromosomes, that crossing over was a four-strand phenomenonandthatchromatidinterference was negligible. The genetical and cytological analysis of crossing over was an important subject at thetime and several other papers were also devoted to it. A. F. BLAKESLEE described a mountain of work on the jimsonweed, Datura.Cleverly starting with a doubled haploid, he systematically found all the primary trisomics and most of the possible secondaries (extra isochromosomes) and tertiaries (fusions of arms from different chromosomes). Each had a characteristic phenotype which, because the strain was isogenic to begin with, was caused solely by gene dosage effects. (I recall the euphoric but short-lasting belief shared with m y late colleague KLAUS PATAU,after the discovery of the first human trisomic, that by analogy with Datura he would soon identify 22 more.) BLAKESLEE alsoused the trisomics to assign mutant genes to specific chromosome arms. 1\11 the Datura species had the same chromosome morphology and number, yet
Albrecht, 249 Alderman, 13 Anderson, 3 I 5 Armstrong, 375 Arnason, 358 Babcock, 91 Bangson, 159 Banta, 229 Baron, 349 Barrows, 8 Beers, 1 18 Belfield. Mrs., 8 3 Bennion. 78 Besley. 9 Bittner, 224 Blakeslee. 187 Bonnevie, 170 Bostian, 568 Bowers, 143 Bowstead, 247 Boyden, 326 Brandt. 138 Bregger.14 Brehme, 3 Briggs, 2 17 Brmk. 232 Brittingham. 346 Brown, 286 Brunson. 293 Bryall. 1 13 Buchhol;., 144 Burhoe. 313 Burnham. 363 Bussell. 26 Carothers, 2 19 Carter. 199 Castle. W. E., 318 Castle, Mrs. W. E., 264 Chapman. 162 Child, 350 Christian, 99 Chroboc7ek. 101 Chung, 106 Clark, E. B.. 381 Clark, F. H., 379 Clarke, 299 Clausen, 279 Cleland, 24 1 Coffnlan, 364 Cole, 180 Colin, 190 Collins, 2 18 Cook, 267 Cooper, 250 Comer. 367 Crawford, 72 Creighton. 43 Crew, 2 Crofts, 204 Currence, 386 Cutler, 46 Darlington, 68 Davenport, 21 3 Davis, Mrs. Barbara, 7 1 Davis, D. W.. 288 Davis, M. E., 287 Dawson, 164 Deakin, 300 Demerec, 291 Derick, 369 Dermen, 194 Diver, 298 Dobrovolskaia-Zavadskaia, 32
Dob~hansky, 35 Dodge, B. O., 243 Dodge, Mrs. B. O., 344 Duchemin. 3 17 Dunn, 36
Dunning, 184 Durham. 195 Earl, 353 East, 216 Einsele, 1 I 5 Elliot, 385 Ellis, 52 Emerson. R. A,, 25 Emerson, S. H., 333 Erlanson. 330 Estabrook, 88 Eyster. 165 Federley. 169 Feldman, 322 Feng, C. F., 6 3 Fisher, 132 Fitzpatrick, 373 Forbes, 152 Fowlds. 65 Fraser, 23 French, 16 Frets, 6 1 Frost, 360 Gaines, E. F . , 56 Gaines. M., 57 Garber, 200 Gates, R. R., 188 Gates, W. H., 225 Gates, Mrs. W. H., 226 Gay, 29 Gerould,153 Ghigi, 96 Gibson, 84 Gini, 94 Glass, H. B., 341 Glass, L. D., 81 Goldschmidt. 97 Good. 258 Goodale, 295 Goodrich, 126 Gordon, C. D., 161 Gordon, M., 53 Gowen, 28 Green, 32I Gregory, 139 Grossman. 149 Gumbel, 176 Haines, 34 Haldane. 54 Hall, 120 Hamilton, 351 Hanna, 281 Hansen. 17 1 Harland, 230 Hardy, 352 Hart. 259 Hawryluk, 238 Hayden, 3.55 Hays, F. A , , 49 Hays, Mrs. F. A., 50 Hearne, 158 Heizer, 18 Herriott, 33 Hersh, 301 Hetzer, 146 Hill, Mrs. H. D.. I73 Hill, J. B., 174 Hill, L. H., 198 Hinman, 274 Holmes, 79 Hoover, 201 Horowitz, I 1 Houghtaling, 10 Howlett, 76 Hubert, 137 Hume, 236 Hunt. 136
Hurst, 27 Huskins, 80 Husred. 303 Hutt, 2 Ibsen, 310 Irnnler, 155 Ingersoll, 361 Irwin, 338 Jaap. 4 Jackson, 203 Jenkins, 305 Johnson-Little. 266 Jones, 290 Kaliss, 192 Kamenoff, 39 Kaufmann, B. P.. 270 Kaufmann, Mrs. B. P., 297 Kelly, 160 Kemp, 278 Kendall, 339 King, 260 Kirk, 282 Knowles, 335 Kopf, 316 Kornhauser, 15 I Krantz. 384 Krug, 277 Kulkarni, 98 Kwan, 129 Lambert, 362 Lampe, 89 Lancefield, 208 Lebedeff, 239 Lenderking, 334 Leonard, 110 Lesley. 257 L'Heritier, 336 Lindegren, C. C., 75 Lindegren, Mrs. G., 74 Lindsay, 25 1 Lindstt-om, 30 Little, 3 19 Livermore, 273 Lods, 107 Longley, 205 Low. 3 I 1 Luce, 304 Lush, 123 Lynch, 183 Ma, 102 MacArthur, 222 McClintock, 41 McConkey, 283 MacDowell, 215 McEwen, 214 McFadden. 64 McGibbon, 359 McGregor, 374 Macklin, 147 Macoun, 166 McPhee, 268 MacRae, 357 McRostie, 284 Mahoney, 172 Mangelsdorf, 314 Margolis. 348 Marshak, 312 Marshall, 227 Mavor, 127 Maw, A. J. G., 365 Maw, W. A,, 121 Meacharn, 366 Merrell, 90 Metz, 240 Miles, 100 Minns, 37 Mohr, 0. L., 69
Mohr, Mrs. 0. L., 70 Morgan, T. H., 24 Morgan, Mrs. T. H., 116 Morrison, 182 Moyer, 128 Murray, 323 Nabours, 223 Nachtsheim, 168 Neal, 253 Nebel, 197 Neely, 275 Newman, H. H., 86 Newman, L. H., 45 Nolla. 202 Painter, 196 Park, 324 Parker, 254 Parris, 130 Passrnore. 6 Patterson, 207 Perry, E. M.,342 Perry, M. M., 343 Phelps, 38 Pincus, 2 12 Plough, 327 Plunkett, 329 Popenoe, 263 Powers, 156 Prat, 255 Proulx,122 Quisenberry, I63 Raffel, 134 Reed, 380 Rheinheimer, J.. 59 Rheinheimer, Mrs. J., 58 Rhoades, M. M., 40 Rhoades, V. H., 42 Richards, A., 8 5 Richards, M. H., 67 Richey,131 Rieman, 388 Ritzman, 262 Robb, 383 Roberts, 269 Robertson, E. W., 108 Robertson, W. R. B., 48 Romanoff, 31 Roque, 276 Rosinski, 51 Rousseau, 179 Ru, 347 Saenko, 82 Salisbury, I7 Sanders, 175 Sando, 109 Saunders, 141 Sawin, 378 Sax, H.J., 345 Sax, K., 242 Schaffner, J. H., 103 Schaffner, Mrs. J. H., 104 Schmid, 185 Schneider, 27 1 Schultz, 21 Scott, A. C., 193 Scott. M. H.. 289 Senn, H. A., 331 Senn, P. H., 252 Shoemaker, 233 Showalter, 308 Shrigley, 372 Shull, A. F., 206 Shull, G. H., 60 Sirnpson, 7 Singleton, 306 Sinnott, 1 86 Slate, 12
Smith, L., 44 Smith, 0. D., 237 Smith, S. G., 294 Smith, S. N.. 20 Smith, T. L., 307 Smith, W. K., 377 Snell, 340 Snyder, L. H., 92 Snyder, Mrs. L. H., 93 Speiden, 62 Spencer, 77 Spier, 157 Sprague, 1 12 Stadler, 22 Stanton, 11 I Stark, I19 Stebbins, 140 Steele, 248 Stern, 66 Stockard, 87 Stout, 142 Strandskov. 189 Stringfield, 376 Strong, 256 Stuck, 114 Sturtevant. 328 Surnmerby, 167 Sutherland, 371 Swanson, 235 Taylor, 356 Thomas, 280 Thompson, 15 Timofkff-Ressovsky, H., 73 Timofieff-Ressovsky, N., 154 Tulloss, 265 T u i l y , 332 Turk, 19 Upp, 125 Vandel, A , , 95 Vandel, Mrs. A., 135 Vandendries, 177 Van Lone, Mrs. A , , 245 Van Lone, E. E., 246 Varrelman, 209 Vavilov, I48 Vicari, 1 17 Vilmorin, de, 221 Waldron. 55 Waller, 325 Warbritton, 211 Warfel, 296 Warren, D. C., 124 Warren, H. S., 210 Warren, P. A., 234 Warwick, 244 Waters, 320 Watson, 272 Weinstein, 133 Weismann, 191 Wenstrup, 5 Wentworth, 181 Whitaker, 302 White, E. G., 354 White, 0.E., 23 I Whiteside, 370 Whiting, 220 Whitney, D. D., 150 Whitney, L. F . , 261 Wiggins, 292 Wilcox, 387 Winge, 228 Wood, 285 Woolley, 145 Worzella, 47 Yang, 105 Yarnell, 309 Zeleny, 337 Zulueta, 178
FIGURE3 . - C h x q >
p h o t o g r a p h made o n A u g u s t 2.5, 1932. Of .562 m e m b e r s r e g i s t e r e d at Ith;lca, 389 are shown.
I F. A. E. Crew 27 C. C. Hurst J. W. Gowen B. 2 F. 28 Hutt 3 Katherine S . Brehme29 E. H.Gay 4 R. G .Jaap 30 E. W. Lindstrom 5 Edward J. Wenstrup31Anastasia J. Romanoff 6 Sara F. Passmore 32 N. Dobrovolskaia-Zavadskaia 7 J. L. S.Simpson 33 Mrs. F. W. Herriott 8 Florence L. Barrows 34GeorgeHaines 9 Helen Besley 35 Th. Dobzhansky 61 10 Helen Houghtaling 36 L. C. Dunn 62 1 I Solomon Horowita 37 Lua A. Minns I2 G . L. Slate 38 Lillian Phelps 13 W. Alderman H. 39 R. J. Kamenoff 14 John T. Bregger 40 Marcus M. Rhoades 1 BarbaraMcClintock 15 David H.Thompson4 16 A. P. French 42 Rhoades Virginia H. 17 Glen Salisbury Harriet 43 B. Creighton 69 18 E. E. Heizer Luther 44Smith I 9 Kenneth L. Turk 4 5 L. Newman H.7 20 Stuart N. Smith Crawford Mary 4672 Cutler G . H. 2 I Jack Schulta 4 7W. W. Worzella 22 L. J. Stadler 4 8 W. R. B. Robertson Fraser 23 A. C. 4 9 F. A. Hays 24 T. H. Morgan 50 Mrs. F. A. Hays 25 R. A. Emerson 51 B. Rosikki 26 F. P. Bussell 5 2 Zenas H. Ellis
53 Myron Gordon 79 54 J. B. S. Haldane 55 L. R. Waldron 56 E. F. Gaines 57 Margaret Gaines 58 Mrs. J. Rheinheimer 59 J. Rheinheimer 60 George H. Shull G. P. Frets N. R. Speiden 63 C. F. Feng 64 E.S. McFadden 6 5 Matthew Fowlds 6692 Stern Curt 67 MildredHoge Richards 68 C. D. Darlington 0.L. Mohr 70 Mrs. 0.L. Mohr 1 Mrs. Barbara Davis 7 3 H. Timofiiff-Ressovsky 74 Mrs. GertrudeLindegren 7 5 Carl C. Lindegren 76 F. S. Howlett 77 W. P. Spencer 103 John 78 Noel L. Bennion 104
S.J. Holmes 105 80 C. L. Huskins 81 L. C. Glass 82 S. M. Saenko 108 83 Mrs. S. Belfield 8 4 Roy E. Gibson 85 A. Richards 8 6 H. H. Newman 87 C. R. Stockard 88 Estabrook A. H. 8 9 Lois l ~ m p e 90 W. D. Merrell 91 E. B. Babcock Snyder L. H. 9 3 Mrs. L. H. Snyder Corrado 94 120 Gini 9 5 A. Vandel 96 A. Ghigi 97 R. Goldschmidt 9 8 L. G . Kulkarni 124 99 C. Stuart Christian 100 L. Gordon Miles 101 E. Chroboczek 127 102 P. C. Ma Schaffner H. Mrs. John Schaffner H.
Yun-Kuei Yang 106 H. C. Chung I07 E. A. Lods Robertson D. W. 109 W. J. Sando I10 W. H. Leonard I I 1 T. R. Stanton I I 2 G . F. Sprague 1 I3 A. A. Bvran Florence 1Stuck 14 I 15 W. C. Einsele I 16 Mrs. T. H. Morgan 117 F. M. Vicari 1 I8 Catherine V. Beers I I 9 Mary B. Stark G . 0.Hall 121 W. A. Maw 122 Maurice Proulx 123 J. L. Lush D. C. Warren 125 Charles W. Upp 126 H. B. Coodrich J. W. Mavor I 2 8 Raymond T. Mover 129 C. C. Kwan
130 C. K. Parris 131 F. D. Richey 132 R. A. Fisher 133 Alexander Weinstein 134 Daniel Raffel 135 Mrs. A. Vandel 136 H. R. Hunt I37 Kurt Hubert 138 A. E. Brandt 139 P. W. Gregory I40 G . L. Stebbins. Jr. 14 I A. P. Saunders 142 A. B. Stout 143 C. G . Bowers 144 J. T. Buchhols 145 G . W. Woolley 146 H. 0.Hetzer 147 M. T. Macklin 148 N. 1. Vavilov 149 E. F. Grossman 150 D. D. Whitney 151 S. 1. Kornhauser 152 W. T. M. Forbes 153J. H. Gerould 154 N. TimofPeff-Ressovsky I55 F. R. lmmer
156 L. R. Powers 157 Jane Spier 158 Marie Hearne 159 J. S . Bangson 160 J. P. Kellv 161 C. D. Gordon 162 Arthur B. Chapman 163 John H. Quisenberry 164 Walker M. Dawson 165 William H. Eyster 166 W. T. Macoun 167 R. Summerby 168 H. Nachtsheim 169 Harry Federley 170 Kristine Bonnevie 17 1 Mrs. Hansen I72 C. H. Mahoney 173 Mrs. Helen D. Hill 174 J. Ben Hill 175J. Sanders I76 E. J. Gumbel 177 RenP Vandendries 178 A. Zulueta 179 Jacques Rousseau I80 L. J. Cole 181 E. N. Wentwonh
182 F. B. Morrison 183 C. J. Lynch 184 Wilhelmina F. Dunning 185 A. Schmid 186 E. W. Sinnott 187 A. F. Blakeslee 188 R. Ruggles Gates 189 H. H. Strandskov I90 Edward C. Colin 191 M. N. Weismann 192 Nathan Kaliss 193 A. C. Scott 194 Haig Dermen 195 G . B. Durham 196 Reginald H. Painter I97 B. Nebel 198 Lillian Hollingshead Hill 199 GeorgeS . Carter 200 R. J. Garber 201 Max M. Hoover 202 J. A. B. Nolla 203 V. W. Jackson 204 JohnT. Corfts 205 W. H. Longley 206 A. Franklin Shull 207 J. T. Patterson
208 D. E. Lancefield 209 F. A. Varrelman 2 10 Herbert S . Warren 2 I I Virgene Warbritton 2 I 2 G . Pincus 213 C. B. Davenport 2 14 R. S. McEwen 215 E. C. MacDowell 216 E.M.FASI 2 17 Fred N. Briggs 218 J. L. Collins 2 I9 E. Eleanor Carothers 220 P. W. Whiting 22 I Roger d e Vilmorin 222 J. W . MacArthur 223 R. K. Nabours 224 JohnJ. Bittner 225 Wm. H. Gates 226 Mrs. Wm. H. Gates 227 Ruth Marshall 228 0. Winge 229 A. M. Banta 230 S. C. Harland 231 0. E. White 232 R. A. Brink 233 D. N. Shoemaker
234 Paul A. Warren 235 A. F. Swanson 236 E. P. Hume 237 0.D. Smith 238 T. Hawryluk 239 G . A. Lebedeff 240 C. W. Metz 24 1 Ralph E. Cleland 242 Karl Sax 243 B. 0. Dodge 244 B.L. Warwick 245 Mrs. Adeline Van Lone 246 E.E. Van Lone 247 J. E. Bowstead 248 D. G . Steele 249 H. R. Albrecht 250 D. C. Cooper 251 Ruth H. Lindsay 252 P. H. Senn 253 N. P. Neal 254 M. C. Parker 255 Henri Prat 256 L. C. Strong 257 J. W. Lesley 258 H. U. Good 259 E. H. Hart
60 H. D. King 61 L.F. Whitney 62 E. G . Ritzman 63 Pall1 Popenoe 64 Mrs. W. E. Castle 6 5 Mrs. F. S. Tulloss 66 Beatrice Johnson-Little 67 Robert Cook 68 Hugh C. McPhee 69 Elmer Roberts 70 B. P. Kaufmann 71 Burch H. Schneider 72 L. R. Watson 73 J. R. Livermore 74 R. R. Hinman 7 5 W. Neely 76 Arruro Roque 77 Carlos A. Krug 78 Tage Kemp 79 J. Clausen 8 0 L. C. Thomas 81 W. F. Hanna 8 2 L. E. Kirk 83 0.McConkey 84 G . P. McRostie 85 T. R. Wood
286 Mary J. Brown 287 Mary Eleanor Davis 288 Donald W. Davis 289 Martha H. Scott 290 Donald F. Jones 291 M. Demerec 292 R. G . Wiggans 293 A. M. Brunson 294 S.G . Smith 295 H. D. Goodale 296 H. E. Warfel 297 Mrs. B. P. Kaufmann 298 Ernest C. Diver 299 A. E. Clarke 300 Alan Deakin 301 A. H. Hersh 302 T. W. Whitaker 303 Ladley Husted 304 Wilbur M. Luce 305 Merle T. Jenkins 306 W. R. Singleton 307 T. L. Smith 308 H. M. Showalter 309 S. H. Yarnell 310 H. L. Ibsen 31 I Albert Lorz
3 12 Alfred Marshak 3 1 3 S. 0. Burhoe 3 14 P. C. Mangelsdorf 3 15 Edgar Anderson 3 16 Kenneth Kopf 3 17 W. J. Duchemin 318 W. E. Castle 319 C. C. Little 320 N. F. Waters 32 1 C. V. Green 322 Horace Feldman 323 J. M. Murray 324 J. B. Park 325 A. E. Waller 326 Alan Boyden 327 H. H. Plough 328 A. H. Sturtevant 329 C. R. Plunkett 330 E. W. Erlanson 33 I H. A. Senn 332 Eva M.Tully 333 S. H. Emerson 334 Ruth E. Lenderking 335 Emerson G . Knowles 336 Philippe L'Heritier 337 Charles Zeleny
338 M. R. Irwin 339 J. 1. Kendall 340 G . D. Snell 341 H. Bentley Glass 342 E.M. Perry 343 M.M. Perry 344 Mrs. B. 0.Dodge 345 Hally J. Sax 346 Wm. H. Brittingham 347 S. K. Ru 348 OttoS . Margolis 349 A. L. Baron 350 George P. Child 351 L. H. Hamilton 352 M. H. Harnly 353 R. 0.Earl 354 E. Grace White 355 M. A. Hayden 356 L. W. Taylor 357 N. A. MacRae 358 T. J. Arnason 359 W. H. McGibbon 360 Howard B. Frost 361 Floyd lngersoll 362 W. V. Lambert 363 C, R. Burnham
364 F. A. Coffman 365 A. J. G . Maw 366 F.B. Meacham 367 J. B. Cotner 368 C. H. Bostian 369 R. A. Derick 370 A. G . Whiteside 371 J. R. G . Sutherland 372 Edward W. Shrigley 373 H. J. Fitzpatrick 374 W. G . McGregor 375 J. M. Armstrong 376 G . H. Stringfield 377 W. K. Smith 378 P. B. Sawin 379 F. H. Clark 380 S. C. Reed 381 Everett B. Clark 383 R. Cumming Robb 384 F. A. Krantz 385 Elliot 386 T. M. Currence 387 A. N. Wilcox 388 G . H. Rieman 389 Herben P. Riley
Perspectives the hybrid meioses producedchromosomerings, showing that the chromosome arms had been extensively shuffled by translocations-a most convincing demonstration that translocations were an important part of the evolutionary process. A high point of the 1932 Congress was the paper by N. I. VAVILOV,in which hereported extensive geographical studiesof the wild relatives of cultivated plants. He described a series of polyploid potatoes in South America, wheat varieties in Abyssinia, and many others. In those premolecular days, he realized that one could compare noncrossable species by looking for homologous chromosome changesand genetic variants. He emphasized thatthefuture of plant breeding must rely on wild varieties as sources of useful genetic variability and established foundation stocks in widely different latitudes in the USSR. Alas, VAVILOV’Smethods promised only hard work, more geographical expeditions, and slow (but certain) improvement of cultivated crops. In contrast LYSENKO’S expansive promises based on his eccentric Lamarckian views caught Stalin’s eye. It is ironic that, inhis Congress paper, VAVILOV called attention to the “remarkable discovery” by LYSENKO of “simple physiological methods of shortening the period of growth, of transforming winter varieties into spring ones and late varieties into early ones by inducing processes of fermentation in the seeds before sowing them,” thereby building up the man who would later be his ruination.’VAvrLov was the first of four speakers in a session on evolution. The other three were R. A. FISHER, J. B. S. HALDANE and SEWALL WRIGHT.This was one of the few times, if not the only one, that this triumvirate who founded the genetical theory of evolution appeared on the same platform. T h e session was organized by E. M. EAST, who asked each of the speakersto give anonmathematicalpresentation. HALDANE asked, “Can evolution be explained in terms of known genetical facts?” He concluded that a great many facts can be explained qualitatively and quantitatively, and “while we cannot yet explain all evolutionary phenomena in terms of known genetical facts, thenumber of phenomena so explicable increases
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VAVILOVwas named president of the1939 Congress in Edinburgh. Shortly before the opening, he sent a letter noting that the Congress had been postponed for a year by the Academy of Sciences of the USSR so as to make better arrangements, and added, “TheInternational Committee, however, postponed the Seventh International Congress of Genetics until 1939 and chose as its place of meeting not the USSR but another country. Under such circumstances Soviet geneticists and plant and animal breeders do not consider it possible to take part in the Congress.“ Nobody who knew VAVILOV thought this represented his true feelings. F. A. E. CREWwas then chosen President and, with his usual grace, said, “1 understand that in those places where films are made, everystar has his shadow (technically known, I think, as a ‘stand-in’)who is required to look more or less like his principal and to take his place in the more arduous parts of his role. I would suggest to you that at the moment this is exactly what I am-a stand-in for a star. You invite me to play a part that VAVlLOV would have so adorned. Around my unwilling shoulders you drape his robes, and if in them I seem to walk ungainly, you will not forget that this mantle was tailored for a bigger man” (PUNNETT 1939). Soon after, VAVILOVwas arrested and died in prison,
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every year, and there is no sign that the possibilities of explanation are reaching a limit.” FISHERartfully noted that his title might well have been, in antiparallelism to HALDANE’S, “Can genetical phenomena be explained in terms of known evolutionary causes?” and discussed the evolution of such genetic fundamentals as dominance and linkage. He accepted EAST’Sadvice to suppress the mathematics and said, “As I am a mathematician by trade, perhaps I should explain that I shall use no mathematics, partly because I recognize that the first duty of amathematician, rather like that of a lion tamer, is to keep his mathematics in their place.” WRIGHT’S paper has turned out to be the most influential of the three. This is partly because FISHERand HALDANE had both recentlycompleted books that developed their ideas more completely, whereas WRIGHThad only writtenapaper that hardly anyone understood. His paper at theCongress was his first attemptto explain verbally the importance of population structure, random drift, and differential migration-what he later called the “shifting balance” theory of evolution, as controversial today as it wasin 1932. WRIGHT spent much of the remainder ofhis long life restating the theory and arguing for it, but hardly changing it. My pleasure in writing this essaywas enhanced by working with WRIGHT’Swell-worn copy of the Proceedings and inferring from his annotations on the abstracts which talks interested him most. In addition tothe plenaryspeakers, there were about200papers. T. S. PAINTERand MULLER reporteda cytological map of Drosophila. This was madefrom metaphase chromosomes, salivary chromosomes having not yet been discovered, and showed a large variation in gene density in different chromosome regions. C. C. LITTLE,the founderofThe Jackson Laboratory,argued against a highly publicized view that cancer in mice was a single recessive. DOBZHANSKY and STURTEVANT discussed variegated position effects produced by translocations. GEORGE SNELL,laterto win a Nobel Prize for his work on histocompatibility, reported fertility reduction in irradiated mice, presumably the resultof translocations. H. H. NEWMANdescribed 10 sets of identical twins who hadbeen rearedapart. D. F. JONES reported using two mutant genesto create a heterosexual strain of maize. (Some years later I explained the fundamentals of genetics to him, not knowing who he was. He was a quick study.) LILLIAN V. MORGANdescribed the properties of a ring chromosome, including the predicted absenceof single exchanges. There were scores of papers using plants and animals other than Drosophila and maize. There was also a paper entitled “Genetical Engineering,” meaning the application of genetic principles to animal and plant breeding. It is fascinating to see what kinds of problems were attracting attention in those days and what kinds of methods
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were used. The variety of animal and plant species discussed was much greater than ata genetics meeting today. What about theday-to-day aspects of this Congress? Remember that 1932 was the worst of the depression. Almost everybody was poor, and therewere no grants to pay travel and living expenses. Nevertheless, 856 registered. The total expenses of the Congress were $17,583.58. For comparison, the 1988 TorontoCongress spent Can$1,396,701.16 (=US$1,135,000) and 3702 attended. The advance registration fee was $10 for full members and $6 for students. Those who couldn’t afford the whole fee at one time could pay $5 down and the balance on arrival. Rooms in the residence halls at Cornell were $1.75 perday and rates in private rooming houses in the campus area ranged from $1 .OO to $1.50.Those who traveled by car were told that “thereare several very attractivecamping places within thirty minutes’ drive of Ithaca.” Railroad fare from NewYork to Ithaca was $8.93 and attendees could get a round trip with various excursion privi-
leges for 50% more. But the hard times took their toll; of 856 registered, only 562 were able to attend. Despite great advances around the periphery, the central question of genetics-the nature of the gene, and how it replicates and mutates-was still elusive. In a review of the Congress, R. C. COOK(1932) said that “Oceans of words were spilled in formal and informal gatherings to discuss the vital question: ‘What is the gene?’ but that important entity is still elusive. Perhaps in 1937 the answer may be forthcoming.” He was too optimistic; it would be two decades before WATSON and CRICK turned the trick. LITERATURE CITED
K. C., 1932 The GeneticsCongress. J . Hered. 23: 355960. JONES, I). F,,ED., 1932 Proceedings of the Sixth International Congress of Genetics. Vol. 1. Transactions and General Addresses. Vol 2. Condensed Articles and Descriptions ofExhibits. Brooklyn Botanic Garden, Brooklyn, N Y . I’UNNE’I-I‘, K. C . , ED. 1939 Proceedings of the Seventh International (hzetical Congress. Cambridge University Press, Cambridge. COOK,
Perspectives Anecdotal, Historical and Critical Commentaries on Genetics Edited by James F. Crow and William F. Doue
Sixty Years Ago: The 1932 International Congressof Genetics James F. Crow Genetics Department, University of Wisconsin, Madison, Wisconsin 53706
T
HE Sixth International Congress of Genetics, in Ithaca, New York in August, 1932 must have been an exciting experience in those dreary, depression, dust-bowl years. Five hundredand sixty-two people came to Cornel1 from 35 countries, often at considerable financial sacrifice, for in those days participants paid their own expenses (Figure 3 (foldout)). The Congress (JONES 1932)featureddemonstrations of living organisms, charts, photographs, and hundreds of microscopes for viewing specimens, especially cytological. Exhibits included 15 groups of invertebrates,includingaphids,echinoderms, mollusks and tunicates, many with living specimens. There were 15 vertebrate species and 35 genera of plants. In addition there were an equal number of vegetable crops, flowers and fruits. The description of the exhibits took about 250 pages in the volume of abstracts available at the start of the Congress. The sheer magnitude and variety must have been overwhelming. The living plant exhibits, extending over about a hectare, attracted the most attention. These involved carefulplanning, with many of the types grown experimentally the year before to determine planting time so that demonstrations would peak in August. This was done under thesupervision of MARCUS RHOADES, who did much of the field work himself, and that same year received his Ph.D. from Cornell. (RHOADES recently died and will be the subject of an obituary in a forthcoming issue.) T h e most popular exhibit was a “living chromosomemap” inwhich mutant maize plants were arranged in positions corresponding to the locations of the mutations on the linkage map (Figure 1). The Congress set aside ample time for seeing the exhibits, including all day Sunday for those who didn’t want to see Niagara Falls. Like the currently popular poster sessions, these offered the opportunity for individual discussions. According to the Congressdescription, “Even a most retiring person will easily find an opportunity to approach a Genetics 131: 761-768 (August, 1992)
person demonstrating his exhibits, in order to ask a question or to start a discussion.” My reason for featuring this particular Congress, however, is not the exhibits, but the number of outstanding addresses that have had a lasting influence. The year 1932 was near the end of the golden era of classical genetics, the period when the tools were breeding experiments and the microscope and when the riddlesof genetic transmission were largely solved. The advances of the previous 32 years had created the new science of genetics. Before discussing the Sixth Congress, I’ll briefly mention some other early ones. T h e First antedated the rediscovery of MENDEL’Slaws. T h e “International Conference on Hybridisation and on the Cross-Breeding of Varieties” was held in London on July 11-1 2, 1899atthe instigation of the Royal Horticultural Society. T h e Congress featured a talk by WILLIAM BATESON, soon to become one of MENDEL’Searliest and strongest supporters. T h e Second “International ConferenceonPlantBreeding and Hybridization” occurred in 1902, in New York City. Four years later the Royal Horticultural Society again convenedan “International Conference onHybridisation and Plant Breeding” in London in 1906,and again BATESON was featured, this time as President.In histalk he urgedthat his new verbal construct genetics be adopted. Accordingly, the printed volume was entitled Report of the ThirdInternational Conference on Genetics. For the first time, animal experiments were included. The Fourth Congressof Genetics took place in Paris in 191 1. The Fifth was scheduled for 1916, five years later, but World War I intervened, and it was not held until 1927, in Berlin. By then genetics and geneticists hadgrown; 903 membersfrom 35 countries attended, and the Proceedings include 148 papers. The Sixth Congress, the subject of this essay, was held on schedule five years later. T h e Seventh was
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( 1 9.72).
planned for 1937 in Moscow, but was postponed a year by the National Academy of the USSR and was eventually held in Edinburgh in 1939. One month before the opening the Russian delegation withdrew and 50 papers were cancelled. O n the second day of the Congress, British citizens living in Germany were advised to return home and several non-British delegations left. The Congress continued, however, with most of the members remaining.O n September 1, the day after ad-journment, fighting began on the PolishGerman border. The SS Athenia with several geneticists aboard was torpedoed in the Atlantic. B. PRICE, C. W. COTTERMAN, W. LAWRENCE and W. R. SINGLETON survived but the F. W. TINNEYS of the University of Wisconsin perished. T o return to the 1932 Congress: the President was T. H. MORGAN(Figure 2). His attendance was greeted with great relief, for he had recently been in a severe automobile wreck and his friends feared that he would not recover i n time. MORGAN'Saddress dealt mainly with history, but he listedwhat he regarded as the five mostimportant problems. They were: ( 1 ) "growth and duplication" of genes; (2) physical interpretation of synapsis and crossing over: (3) relation of genes to characters; (4) nature of themutation process; ( 5 ) application to plant and animal breeding. This reflected the state of genetics at the time. Transmission genetics was essentially solved, and it was time to understand the geneand development in a morebasic way. MORGANhad no way of knowing that the study oftiny organisms andenormous molecules would converge in the decades ahead. One of the most exciting and influential addresses was delivered by H. J. MULLER. This was not long after his discovery of X-ray mutagenesis, and he exploited this tool to the fullest. He reviewed the induc-
tion of mutations and chromosome breakage by radiation, and amongother things emphasized how the use of deletions enabled h i m to discern thedifference between gain and loss of gene function. He introduced the now-standard vocabulary of amorphic, hypomorphic, hvpermorphic. neomorphic and antimorphic mutations. MULLERalso introduced the idea of dosage compensation and its elucidation by hypomorphic mutations. Although his preferred mechanism turnedout to be wrong, the observation was remarkably astute and the generalizations from it to the meticulousness of evolution remarkably far-reaching. I have long erl-joyed MULLER'S reasoning from dosage compensation of mutant alleles to the conclusion, later abundantly confirmed, that normal alleles are not completely dominant. What a wealth of ingenuity of both concept and experimental trickerv he displayed! Those who claim MUILER as the idea man of early genetics can find ample supporting evidence here. A. H. STURTEVANT, with typical conciseness, discussed the possibility of using mosaics for elucidating developmental patterns. The newly discovered clarpt mutation in Drosophilasimulans, which produceda high rate of nondisjunction in early cleavage, greatly facilitated this analysis. I t was to be many years before the corresponding mutation was found in Drosophila melanogaster. The system was then exploited for fate mapping andneurogenetics, and SEYMOURBENZER coined the eponymous unit sturt. CURTSTERN presented his famous experiment using translocations to demonstrate the physical reality of crossing over. His diagram from this Congress has been reproduced in one textbook after another. Essentially the same experiment had been done in maize by HARRIETCREIGHTONand BARRARAMC-
Perspectives
CLINTOCK, but was published elsewhere. At this Congress they went furtherand used doubly heteromorphicchromosomes todemonstratefour-strand crossing over. This was a subject of interest at the time because of BELLING’S hypothesis of crossing over as a copying switch between newly replicated strands, which would not permit four-strand exchange. Mcname appears at many places in the proCLINTOCK’S ceedings. She gave a paper on nonhonlologous chromosome associations. She also did the cytology for another important paper, that of L. J. STADLER. STADLER, deep and thoughtful as always, argued cogently that radiation produces mainly chromosome rearrangements and few, if any, “real” genemutations. In contrast, TIMOF~EFF-RESSOVSKY discussed forward and backward mutations in great detail and believed that X-rays could indeed produce both. He ended his paper on a euphoric note: “We geneticists are in a very happy condition: our science is young, its ‘development curve’is rising rapidly and the future w i l l bring us the most interesting facts and views concerning the gene problem.”Alas, this was one year beforeHitlercame to power and TIMOF~EFF’S life changed for the worse as he was caught up i n both tragic dictatorships, Germany and USSR. R. A. EMERSONgave a thorough review of maize genetics. All 10 linkage groupshad by then been identified, some 100 mutations had been assigned to a group,andabout 50 had been reasonably well mapped (Figure 1). The assignment of linkage groups to chromosomes was done mainly by MCCLINTOCK,
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using trisomics. The trisomics, in turn, were obtained using GEORGEBEADLE’S asynaptic gene, which produced abundant triploids from which all the primary trisomics were easily derived. EMERSON also described the variegated pericarp genes, now known to be the result of a transposable element. Later R. A. BRINK made this mutant the focus of his study, and variegated pericarp became a part of the MCCLINTOCK legend. BEADIXwasjust deserting corn for Drosophila and w a s engaged i n showing, through the use of attachedX chromosomes, that crossing over was a four-strand phenomenonandthatchromatidinterference was negligible. The genetical and cytological analysis of crossing over was an important subject at thetime and several other papers were also devoted to it. A. F. BLAKESLEE described a mountain of work on the jimsonweed, Datura.Cleverly starting with a doubled haploid, he systematically found all the primary trisomics and most of the possible secondaries (extra isochromosomes) and tertiaries (fusions of arms from different chromosomes). Each had a characteristic phenotype which, because the strain was isogenic to begin with, was caused solely by gene dosage effects. (I recall the euphoric but short-lasting belief shared with m y late colleague KLAUS PATAU,after the discovery of the first human trisomic, that by analogy with Datura he would soon identify 22 more.) BLAKESLEE alsoused the trisomics to assign mutant genes to specific chromosome arms. 1\11 the Datura species had the same chromosome morphology and number, yet
Albrecht, 249 Alderman, 13 Anderson, 3 I 5 Armstrong, 375 Arnason, 358 Babcock, 91 Bangson, 159 Banta, 229 Baron, 349 Barrows, 8 Beers, 1 18 Belfield. Mrs., 8 3 Bennion. 78 Besley. 9 Bittner, 224 Blakeslee. 187 Bonnevie, 170 Bostian, 568 Bowers, 143 Bowstead, 247 Boyden, 326 Brandt. 138 Bregger.14 Brehme, 3 Briggs, 2 17 Brmk. 232 Brittingham. 346 Brown, 286 Brunson. 293 Bryall. 1 13 Buchhol;., 144 Burhoe. 313 Burnham. 363 Bussell. 26 Carothers, 2 19 Carter. 199 Castle. W. E., 318 Castle, Mrs. W. E., 264 Chapman. 162 Child, 350 Christian, 99 Chroboc7ek. 101 Chung, 106 Clark, E. B.. 381 Clark, F. H., 379 Clarke, 299 Clausen, 279 Cleland, 24 1 Coffnlan, 364 Cole, 180 Colin, 190 Collins, 2 18 Cook, 267 Cooper, 250 Comer. 367 Crawford, 72 Creighton. 43 Crew, 2 Crofts, 204 Currence, 386 Cutler, 46 Darlington, 68 Davenport, 21 3 Davis, Mrs. Barbara, 7 1 Davis, D. W.. 288 Davis, M. E., 287 Dawson, 164 Deakin, 300 Demerec, 291 Derick, 369 Dermen, 194 Diver, 298 Dobrovolskaia-Zavadskaia, 32
Dob~hansky, 35 Dodge, B. O., 243 Dodge, Mrs. B. O., 344 Duchemin. 3 17 Dunn, 36
Dunning, 184 Durham. 195 Earl, 353 East, 216 Einsele, 1 I 5 Elliot, 385 Ellis, 52 Emerson. R. A,, 25 Emerson, S. H., 333 Erlanson. 330 Estabrook, 88 Eyster. 165 Federley. 169 Feldman, 322 Feng, C. F., 6 3 Fisher, 132 Fitzpatrick, 373 Forbes, 152 Fowlds. 65 Fraser, 23 French, 16 Frets, 6 1 Frost, 360 Gaines, E. F . , 56 Gaines. M., 57 Garber, 200 Gates, R. R., 188 Gates, W. H., 225 Gates, Mrs. W. H., 226 Gay, 29 Gerould,153 Ghigi, 96 Gibson, 84 Gini, 94 Glass, H. B., 341 Glass, L. D., 81 Goldschmidt. 97 Good. 258 Goodale, 295 Goodrich, 126 Gordon, C. D., 161 Gordon, M., 53 Gowen, 28 Green, 32I Gregory, 139 Grossman. 149 Gumbel, 176 Haines, 34 Haldane. 54 Hall, 120 Hamilton, 351 Hanna, 281 Hansen. 17 1 Harland, 230 Hardy, 352 Hart. 259 Hawryluk, 238 Hayden, 3.55 Hays, F. A , , 49 Hays, Mrs. F. A., 50 Hearne, 158 Heizer, 18 Herriott, 33 Hersh, 301 Hetzer, 146 Hill, Mrs. H. D.. I73 Hill, J. B., 174 Hill, L. H., 198 Hinman, 274 Holmes, 79 Hoover, 201 Horowitz, I 1 Houghtaling, 10 Howlett, 76 Hubert, 137 Hume, 236 Hunt. 136
Hurst, 27 Huskins, 80 Husred. 303 Hutt, 2 Ibsen, 310 Irnnler, 155 Ingersoll, 361 Irwin, 338 Jaap. 4 Jackson, 203 Jenkins, 305 Johnson-Little. 266 Jones, 290 Kaliss, 192 Kamenoff, 39 Kaufmann, B. P.. 270 Kaufmann, Mrs. B. P., 297 Kelly, 160 Kemp, 278 Kendall, 339 King, 260 Kirk, 282 Knowles, 335 Kopf, 316 Kornhauser, 15 I Krantz. 384 Krug, 277 Kulkarni, 98 Kwan, 129 Lambert, 362 Lampe, 89 Lancefield, 208 Lebedeff, 239 Lenderking, 334 Leonard, 110 Lesley. 257 L'Heritier, 336 Lindegren, C. C., 75 Lindegren, Mrs. G., 74 Lindsay, 25 1 Lindstt-om, 30 Little, 3 19 Livermore, 273 Lods, 107 Longley, 205 Low. 3 I 1 Luce, 304 Lush, 123 Lynch, 183 Ma, 102 MacArthur, 222 McClintock, 41 McConkey, 283 MacDowell, 215 McEwen, 214 McFadden. 64 McGibbon, 359 McGregor, 374 Macklin, 147 Macoun, 166 McPhee, 268 MacRae, 357 McRostie, 284 Mahoney, 172 Mangelsdorf, 314 Margolis. 348 Marshak, 312 Marshall, 227 Mavor, 127 Maw, A. J. G., 365 Maw, W. A,, 121 Meacharn, 366 Merrell, 90 Metz, 240 Miles, 100 Minns, 37 Mohr, 0. L., 69
Mohr, Mrs. 0. L., 70 Morgan, T. H., 24 Morgan, Mrs. T. H., 116 Morrison, 182 Moyer, 128 Murray, 323 Nabours, 223 Nachtsheim, 168 Neal, 253 Nebel, 197 Neely, 275 Newman, H. H., 86 Newman, L. H., 45 Nolla. 202 Painter, 196 Park, 324 Parker, 254 Parris, 130 Passrnore. 6 Patterson, 207 Perry, E. M.,342 Perry, M. M., 343 Phelps, 38 Pincus, 2 12 Plough, 327 Plunkett, 329 Popenoe, 263 Powers, 156 Prat, 255 Proulx,122 Quisenberry, I63 Raffel, 134 Reed, 380 Rheinheimer, J.. 59 Rheinheimer, Mrs. J., 58 Rhoades, M. M., 40 Rhoades, V. H., 42 Richards, A., 8 5 Richards, M. H., 67 Richey,131 Rieman, 388 Ritzman, 262 Robb, 383 Roberts, 269 Robertson, E. W., 108 Robertson, W. R. B., 48 Romanoff, 31 Roque, 276 Rosinski, 51 Rousseau, 179 Ru, 347 Saenko, 82 Salisbury, I7 Sanders, 175 Sando, 109 Saunders, 141 Sawin, 378 Sax, H.J., 345 Sax, K., 242 Schaffner, J. H., 103 Schaffner, Mrs. J. H., 104 Schmid, 185 Schneider, 27 1 Schultz, 21 Scott, A. C., 193 Scott. M. H.. 289 Senn, H. A., 331 Senn, P. H., 252 Shoemaker, 233 Showalter, 308 Shrigley, 372 Shull, A. F., 206 Shull, G. H., 60 Sirnpson, 7 Singleton, 306 Sinnott, 1 86 Slate, 12
Smith, L., 44 Smith, 0. D., 237 Smith, S. G., 294 Smith, S. N.. 20 Smith, T. L., 307 Smith, W. K., 377 Snell, 340 Snyder, L. H., 92 Snyder, Mrs. L. H., 93 Speiden, 62 Spencer, 77 Spier, 157 Sprague, 1 12 Stadler, 22 Stanton, 11 I Stark, I19 Stebbins, 140 Steele, 248 Stern, 66 Stockard, 87 Stout, 142 Strandskov. 189 Stringfield, 376 Strong, 256 Stuck, 114 Sturtevant. 328 Surnmerby, 167 Sutherland, 371 Swanson, 235 Taylor, 356 Thomas, 280 Thompson, 15 Timofkff-Ressovsky, H., 73 Timofieff-Ressovsky, N., 154 Tulloss, 265 T u i l y , 332 Turk, 19 Upp, 125 Vandel, A , , 95 Vandel, Mrs. A., 135 Vandendries, 177 Van Lone, Mrs. A , , 245 Van Lone, E. E., 246 Varrelman, 209 Vavilov, I48 Vicari, 1 17 Vilmorin, de, 221 Waldron. 55 Waller, 325 Warbritton, 211 Warfel, 296 Warren, D. C., 124 Warren, H. S., 210 Warren, P. A., 234 Warwick, 244 Waters, 320 Watson, 272 Weinstein, 133 Weismann, 191 Wenstrup, 5 Wentworth, 181 Whitaker, 302 White, E. G., 354 White, 0.E., 23 I Whiteside, 370 Whiting, 220 Whitney, D. D., 150 Whitney, L. F . , 261 Wiggins, 292 Wilcox, 387 Winge, 228 Wood, 285 Woolley, 145 Worzella, 47 Yang, 105 Yarnell, 309 Zeleny, 337 Zulueta, 178
FIGURE3 . - C h x q >
p h o t o g r a p h made o n A u g u s t 2.5, 1932. Of .562 m e m b e r s r e g i s t e r e d at Ith;lca, 389 are shown.
I F. A. E. Crew 27 C. C. Hurst J. W. Gowen B. 2 F. 28 Hutt 3 Katherine S . Brehme29 E. H.Gay 4 R. G .Jaap 30 E. W. Lindstrom 5 Edward J. Wenstrup31Anastasia J. Romanoff 6 Sara F. Passmore 32 N. Dobrovolskaia-Zavadskaia 7 J. L. S.Simpson 33 Mrs. F. W. Herriott 8 Florence L. Barrows 34GeorgeHaines 9 Helen Besley 35 Th. Dobzhansky 61 10 Helen Houghtaling 36 L. C. Dunn 62 1 I Solomon Horowita 37 Lua A. Minns I2 G . L. Slate 38 Lillian Phelps 13 W. Alderman H. 39 R. J. Kamenoff 14 John T. Bregger 40 Marcus M. Rhoades 1 BarbaraMcClintock 15 David H.Thompson4 16 A. P. French 42 Rhoades Virginia H. 17 Glen Salisbury Harriet 43 B. Creighton 69 18 E. E. Heizer Luther 44Smith I 9 Kenneth L. Turk 4 5 L. Newman H.7 20 Stuart N. Smith Crawford Mary 4672 Cutler G . H. 2 I Jack Schulta 4 7W. W. Worzella 22 L. J. Stadler 4 8 W. R. B. Robertson Fraser 23 A. C. 4 9 F. A. Hays 24 T. H. Morgan 50 Mrs. F. A. Hays 25 R. A. Emerson 51 B. Rosikki 26 F. P. Bussell 5 2 Zenas H. Ellis
53 Myron Gordon 79 54 J. B. S. Haldane 55 L. R. Waldron 56 E. F. Gaines 57 Margaret Gaines 58 Mrs. J. Rheinheimer 59 J. Rheinheimer 60 George H. Shull G. P. Frets N. R. Speiden 63 C. F. Feng 64 E.S. McFadden 6 5 Matthew Fowlds 6692 Stern Curt 67 MildredHoge Richards 68 C. D. Darlington 0.L. Mohr 70 Mrs. 0.L. Mohr 1 Mrs. Barbara Davis 7 3 H. Timofiiff-Ressovsky 74 Mrs. GertrudeLindegren 7 5 Carl C. Lindegren 76 F. S. Howlett 77 W. P. Spencer 103 John 78 Noel L. Bennion 104
S.J. Holmes 105 80 C. L. Huskins 81 L. C. Glass 82 S. M. Saenko 108 83 Mrs. S. Belfield 8 4 Roy E. Gibson 85 A. Richards 8 6 H. H. Newman 87 C. R. Stockard 88 Estabrook A. H. 8 9 Lois l ~ m p e 90 W. D. Merrell 91 E. B. Babcock Snyder L. H. 9 3 Mrs. L. H. Snyder Corrado 94 120 Gini 9 5 A. Vandel 96 A. Ghigi 97 R. Goldschmidt 9 8 L. G . Kulkarni 124 99 C. Stuart Christian 100 L. Gordon Miles 101 E. Chroboczek 127 102 P. C. Ma Schaffner H. Mrs. John Schaffner H.
Yun-Kuei Yang 106 H. C. Chung I07 E. A. Lods Robertson D. W. 109 W. J. Sando I10 W. H. Leonard I I 1 T. R. Stanton I I 2 G . F. Sprague 1 I3 A. A. Bvran Florence 1Stuck 14 I 15 W. C. Einsele I 16 Mrs. T. H. Morgan 117 F. M. Vicari 1 I8 Catherine V. Beers I I 9 Mary B. Stark G . 0.Hall 121 W. A. Maw 122 Maurice Proulx 123 J. L. Lush D. C. Warren 125 Charles W. Upp 126 H. B. Coodrich J. W. Mavor I 2 8 Raymond T. Mover 129 C. C. Kwan
130 C. K. Parris 131 F. D. Richey 132 R. A. Fisher 133 Alexander Weinstein 134 Daniel Raffel 135 Mrs. A. Vandel 136 H. R. Hunt I37 Kurt Hubert 138 A. E. Brandt 139 P. W. Gregory I40 G . L. Stebbins. Jr. 14 I A. P. Saunders 142 A. B. Stout 143 C. G . Bowers 144 J. T. Buchhols 145 G . W. Woolley 146 H. 0.Hetzer 147 M. T. Macklin 148 N. 1. Vavilov 149 E. F. Grossman 150 D. D. Whitney 151 S. 1. Kornhauser 152 W. T. M. Forbes 153J. H. Gerould 154 N. TimofPeff-Ressovsky I55 F. R. lmmer
156 L. R. Powers 157 Jane Spier 158 Marie Hearne 159 J. S . Bangson 160 J. P. Kellv 161 C. D. Gordon 162 Arthur B. Chapman 163 John H. Quisenberry 164 Walker M. Dawson 165 William H. Eyster 166 W. T. Macoun 167 R. Summerby 168 H. Nachtsheim 169 Harry Federley 170 Kristine Bonnevie 17 1 Mrs. Hansen I72 C. H. Mahoney 173 Mrs. Helen D. Hill 174 J. Ben Hill 175J. Sanders I76 E. J. Gumbel 177 RenP Vandendries 178 A. Zulueta 179 Jacques Rousseau I80 L. J. Cole 181 E. N. Wentwonh
182 F. B. Morrison 183 C. J. Lynch 184 Wilhelmina F. Dunning 185 A. Schmid 186 E. W. Sinnott 187 A. F. Blakeslee 188 R. Ruggles Gates 189 H. H. Strandskov I90 Edward C. Colin 191 M. N. Weismann 192 Nathan Kaliss 193 A. C. Scott 194 Haig Dermen 195 G . B. Durham 196 Reginald H. Painter I97 B. Nebel 198 Lillian Hollingshead Hill 199 GeorgeS . Carter 200 R. J. Garber 201 Max M. Hoover 202 J. A. B. Nolla 203 V. W. Jackson 204 JohnT. Corfts 205 W. H. Longley 206 A. Franklin Shull 207 J. T. Patterson
208 D. E. Lancefield 209 F. A. Varrelman 2 10 Herbert S . Warren 2 I I Virgene Warbritton 2 I 2 G . Pincus 213 C. B. Davenport 2 14 R. S. McEwen 215 E. C. MacDowell 216 E.M.FASI 2 17 Fred N. Briggs 218 J. L. Collins 2 I9 E. Eleanor Carothers 220 P. W. Whiting 22 I Roger d e Vilmorin 222 J. W . MacArthur 223 R. K. Nabours 224 JohnJ. Bittner 225 Wm. H. Gates 226 Mrs. Wm. H. Gates 227 Ruth Marshall 228 0. Winge 229 A. M. Banta 230 S. C. Harland 231 0. E. White 232 R. A. Brink 233 D. N. Shoemaker
234 Paul A. Warren 235 A. F. Swanson 236 E. P. Hume 237 0.D. Smith 238 T. Hawryluk 239 G . A. Lebedeff 240 C. W. Metz 24 1 Ralph E. Cleland 242 Karl Sax 243 B. 0. Dodge 244 B.L. Warwick 245 Mrs. Adeline Van Lone 246 E.E. Van Lone 247 J. E. Bowstead 248 D. G . Steele 249 H. R. Albrecht 250 D. C. Cooper 251 Ruth H. Lindsay 252 P. H. Senn 253 N. P. Neal 254 M. C. Parker 255 Henri Prat 256 L. C. Strong 257 J. W. Lesley 258 H. U. Good 259 E. H. Hart
60 H. D. King 61 L.F. Whitney 62 E. G . Ritzman 63 Pall1 Popenoe 64 Mrs. W. E. Castle 6 5 Mrs. F. S. Tulloss 66 Beatrice Johnson-Little 67 Robert Cook 68 Hugh C. McPhee 69 Elmer Roberts 70 B. P. Kaufmann 71 Burch H. Schneider 72 L. R. Watson 73 J. R. Livermore 74 R. R. Hinman 7 5 W. Neely 76 Arruro Roque 77 Carlos A. Krug 78 Tage Kemp 79 J. Clausen 8 0 L. C. Thomas 81 W. F. Hanna 8 2 L. E. Kirk 83 0.McConkey 84 G . P. McRostie 85 T. R. Wood
286 Mary J. Brown 287 Mary Eleanor Davis 288 Donald W. Davis 289 Martha H. Scott 290 Donald F. Jones 291 M. Demerec 292 R. G . Wiggans 293 A. M. Brunson 294 S.G . Smith 295 H. D. Goodale 296 H. E. Warfel 297 Mrs. B. P. Kaufmann 298 Ernest C. Diver 299 A. E. Clarke 300 Alan Deakin 301 A. H. Hersh 302 T. W. Whitaker 303 Ladley Husted 304 Wilbur M. Luce 305 Merle T. Jenkins 306 W. R. Singleton 307 T. L. Smith 308 H. M. Showalter 309 S. H. Yarnell 310 H. L. Ibsen 31 I Albert Lorz
3 12 Alfred Marshak 3 1 3 S. 0. Burhoe 3 14 P. C. Mangelsdorf 3 15 Edgar Anderson 3 16 Kenneth Kopf 3 17 W. J. Duchemin 318 W. E. Castle 319 C. C. Little 320 N. F. Waters 32 1 C. V. Green 322 Horace Feldman 323 J. M. Murray 324 J. B. Park 325 A. E. Waller 326 Alan Boyden 327 H. H. Plough 328 A. H. Sturtevant 329 C. R. Plunkett 330 E. W. Erlanson 33 I H. A. Senn 332 Eva M.Tully 333 S. H. Emerson 334 Ruth E. Lenderking 335 Emerson G . Knowles 336 Philippe L'Heritier 337 Charles Zeleny
338 M. R. Irwin 339 J. 1. Kendall 340 G . D. Snell 341 H. Bentley Glass 342 E.M. Perry 343 M.M. Perry 344 Mrs. B. 0.Dodge 345 Hally J. Sax 346 Wm. H. Brittingham 347 S. K. Ru 348 OttoS . Margolis 349 A. L. Baron 350 George P. Child 351 L. H. Hamilton 352 M. H. Harnly 353 R. 0.Earl 354 E. Grace White 355 M. A. Hayden 356 L. W. Taylor 357 N. A. MacRae 358 T. J. Arnason 359 W. H. McGibbon 360 Howard B. Frost 361 Floyd lngersoll 362 W. V. Lambert 363 C, R. Burnham
364 F. A. Coffman 365 A. J. G . Maw 366 F.B. Meacham 367 J. B. Cotner 368 C. H. Bostian 369 R. A. Derick 370 A. G . Whiteside 371 J. R. G . Sutherland 372 Edward W. Shrigley 373 H. J. Fitzpatrick 374 W. G . McGregor 375 J. M. Armstrong 376 G . H. Stringfield 377 W. K. Smith 378 P. B. Sawin 379 F. H. Clark 380 S. C. Reed 381 Everett B. Clark 383 R. Cumming Robb 384 F. A. Krantz 385 Elliot 386 T. M. Currence 387 A. N. Wilcox 388 G . H. Rieman 389 Herben P. Riley
Perspectives the hybrid meioses producedchromosomerings, showing that the chromosome arms had been extensively shuffled by translocations-a most convincing demonstration that translocations were an important part of the evolutionary process. A high point of the 1932 Congress was the paper by N. I. VAVILOV,in which hereported extensive geographical studiesof the wild relatives of cultivated plants. He described a series of polyploid potatoes in South America, wheat varieties in Abyssinia, and many others. In those premolecular days, he realized that one could compare noncrossable species by looking for homologous chromosome changesand genetic variants. He emphasized thatthefuture of plant breeding must rely on wild varieties as sources of useful genetic variability and established foundation stocks in widely different latitudes in the USSR. Alas, VAVILOV’Smethods promised only hard work, more geographical expeditions, and slow (but certain) improvement of cultivated crops. In contrast LYSENKO’S expansive promises based on his eccentric Lamarckian views caught Stalin’s eye. It is ironic that, inhis Congress paper, VAVILOV called attention to the “remarkable discovery” by LYSENKO of “simple physiological methods of shortening the period of growth, of transforming winter varieties into spring ones and late varieties into early ones by inducing processes of fermentation in the seeds before sowing them,” thereby building up the man who would later be his ruination.’VAvrLov was the first of four speakers in a session on evolution. The other three were R. A. FISHER, J. B. S. HALDANE and SEWALL WRIGHT.This was one of the few times, if not the only one, that this triumvirate who founded the genetical theory of evolution appeared on the same platform. T h e session was organized by E. M. EAST, who asked each of the speakersto give anonmathematicalpresentation. HALDANE asked, “Can evolution be explained in terms of known genetical facts?” He concluded that a great many facts can be explained qualitatively and quantitatively, and “while we cannot yet explain all evolutionary phenomena in terms of known genetical facts, thenumber of phenomena so explicable increases
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VAVILOVwas named president of the1939 Congress in Edinburgh. Shortly before the opening, he sent a letter noting that the Congress had been postponed for a year by the Academy of Sciences of the USSR so as to make better arrangements, and added, “TheInternational Committee, however, postponed the Seventh International Congress of Genetics until 1939 and chose as its place of meeting not the USSR but another country. Under such circumstances Soviet geneticists and plant and animal breeders do not consider it possible to take part in the Congress.“ Nobody who knew VAVILOV thought this represented his true feelings. F. A. E. CREWwas then chosen President and, with his usual grace, said, “1 understand that in those places where films are made, everystar has his shadow (technically known, I think, as a ‘stand-in’)who is required to look more or less like his principal and to take his place in the more arduous parts of his role. I would suggest to you that at the moment this is exactly what I am-a stand-in for a star. You invite me to play a part that VAVlLOV would have so adorned. Around my unwilling shoulders you drape his robes, and if in them I seem to walk ungainly, you will not forget that this mantle was tailored for a bigger man” (PUNNETT 1939). Soon after, VAVILOVwas arrested and died in prison,
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every year, and there is no sign that the possibilities of explanation are reaching a limit.” FISHERartfully noted that his title might well have been, in antiparallelism to HALDANE’S, “Can genetical phenomena be explained in terms of known evolutionary causes?” and discussed the evolution of such genetic fundamentals as dominance and linkage. He accepted EAST’Sadvice to suppress the mathematics and said, “As I am a mathematician by trade, perhaps I should explain that I shall use no mathematics, partly because I recognize that the first duty of amathematician, rather like that of a lion tamer, is to keep his mathematics in their place.” WRIGHT’S paper has turned out to be the most influential of the three. This is partly because FISHERand HALDANE had both recentlycompleted books that developed their ideas more completely, whereas WRIGHThad only writtenapaper that hardly anyone understood. His paper at theCongress was his first attemptto explain verbally the importance of population structure, random drift, and differential migration-what he later called the “shifting balance” theory of evolution, as controversial today as it wasin 1932. WRIGHT spent much of the remainder ofhis long life restating the theory and arguing for it, but hardly changing it. My pleasure in writing this essaywas enhanced by working with WRIGHT’Swell-worn copy of the Proceedings and inferring from his annotations on the abstracts which talks interested him most. In addition tothe plenaryspeakers, there were about200papers. T. S. PAINTERand MULLER reporteda cytological map of Drosophila. This was madefrom metaphase chromosomes, salivary chromosomes having not yet been discovered, and showed a large variation in gene density in different chromosome regions. C. C. LITTLE,the founderofThe Jackson Laboratory,argued against a highly publicized view that cancer in mice was a single recessive. DOBZHANSKY and STURTEVANT discussed variegated position effects produced by translocations. GEORGE SNELL,laterto win a Nobel Prize for his work on histocompatibility, reported fertility reduction in irradiated mice, presumably the resultof translocations. H. H. NEWMANdescribed 10 sets of identical twins who hadbeen rearedapart. D. F. JONES reported using two mutant genesto create a heterosexual strain of maize. (Some years later I explained the fundamentals of genetics to him, not knowing who he was. He was a quick study.) LILLIAN V. MORGANdescribed the properties of a ring chromosome, including the predicted absenceof single exchanges. There were scores of papers using plants and animals other than Drosophila and maize. There was also a paper entitled “Genetical Engineering,” meaning the application of genetic principles to animal and plant breeding. It is fascinating to see what kinds of problems were attracting attention in those days and what kinds of methods
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were used. The variety of animal and plant species discussed was much greater than ata genetics meeting today. What about theday-to-day aspects of this Congress? Remember that 1932 was the worst of the depression. Almost everybody was poor, and therewere no grants to pay travel and living expenses. Nevertheless, 856 registered. The total expenses of the Congress were $17,583.58. For comparison, the 1988 TorontoCongress spent Can$1,396,701.16 (=US$1,135,000) and 3702 attended. The advance registration fee was $10 for full members and $6 for students. Those who couldn’t afford the whole fee at one time could pay $5 down and the balance on arrival. Rooms in the residence halls at Cornell were $1.75 perday and rates in private rooming houses in the campus area ranged from $1 .OO to $1.50.Those who traveled by car were told that “thereare several very attractivecamping places within thirty minutes’ drive of Ithaca.” Railroad fare from NewYork to Ithaca was $8.93 and attendees could get a round trip with various excursion privi-
leges for 50% more. But the hard times took their toll; of 856 registered, only 562 were able to attend. Despite great advances around the periphery, the central question of genetics-the nature of the gene, and how it replicates and mutates-was still elusive. In a review of the Congress, R. C. COOK(1932) said that “Oceans of words were spilled in formal and informal gatherings to discuss the vital question: ‘What is the gene?’ but that important entity is still elusive. Perhaps in 1937 the answer may be forthcoming.” He was too optimistic; it would be two decades before WATSON and CRICK turned the trick. LITERATURE CITED
K. C., 1932 The GeneticsCongress. J . Hered. 23: 355960. JONES, I). F,,ED., 1932 Proceedings of the Sixth International Congress of Genetics. Vol. 1. Transactions and General Addresses. Vol 2. Condensed Articles and Descriptions ofExhibits. Brooklyn Botanic Garden, Brooklyn, N Y . I’UNNE’I-I‘, K. C . , ED. 1939 Proceedings of the Seventh International (hzetical Congress. Cambridge University Press, Cambridge. COOK,
