In Pursuit of the Gene

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Secondary spermatocytes Spermatogonium Primary spermatocyte Spermatids Diploid (2N) The process of spermatogenesis. Illustration by Tamara L. Clark. CELL BIOLOGY © 161 Haploid (1N) transformation of the spermatogonia into spermatocytes, only joining the other 11 tetrads at the moment before the first division. Unlike the other chromosomes, however, each of which was separated into two halves that were sent to two daughter cells, the accessory went undivided into only one of the two secondary spermatocytes. In the final stage, the accessory chromosome was divided and distributed to two (of four) daughter spermatids.
162 ¨ CELL BIOLOGY The clear depiction of the processes of the formation of the “reduced” sperm cells, containing only half of the chromosome number of all the other cells of the body, as well as the details of inheritance of the accessory chromosome, were major accomplishments, but Sutton had deduced something of even greater significance. As Wilson would later recall, in the early spring of 1902 Sutton came into his office and told him that he had finally understood why “the yellow dog was yellow.” With great humility, Wilson would later confess that he did not at first “fully comprehend his conception or realize its entire weight.” 50 Over a summer filled with swimming, sailing, and science in North Carolina and Maine, Sutton gradually brought Wilson around. As Sutton acknowledged in the first sentence of his extraordinary paper written in the fall of 1902, he was building on Boveri’s recent demonstration that each chromosome was a unique individual containing a unique set of hereditary determinants. 51 Applying Boveri’s principle, Sutton declared that each of the different size groups in the lubber must correspond to a definite set of characters, and that a given size chromosome in the sperm nucleus corresponded to the same size chromosome in the egg nucleus. When the two members of each chromosome pair were joined together in the spermatocyte, the maternal and paternal hereditary qualities were localized in a single continuous chromosome mass for the first and only time in the lifetime of an organism, and when, during the process of sex cell formation, they were again separated, so were the corresponding qualities. 52 This then was the basis for the combining of the factors in Mendel’s hybrids and the segregation into pure types seen among the gametes. A model of understatement, Sutton’s paper concluded with one of the central insights of biology: I may finally call attention to the probability that the association of paternal and maternal chromosomes in pairs and their subsequent separation during the reducing division as indicated above may constitute the physical basis of the Mendelian law of heredity. 53 The first paper Sutton wrote in Wilson’s lab, which was submitted for publication on October 17, 1902, was in many ways a direct outgrowth of Boveri’s work, but the second one, which was completed two months later,
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IN PURSUIT OF THE GENE
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To my wife, Ann Hochschild, whose u
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ILLUSTRATIONS Charles Darwin and Fr
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PREFACE ¨ SCIENCE IS FUNDAMENTALLY
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PREFACE © xi diana University, and
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PREFACE © xiii each of the more co
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CHAPTER 1 Viva Pangenesis ¨ IN THE
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VIVA PANGENESIS © 3 ber of small d
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VIVA PANGENESIS © 5 sumption that
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VIVA PANGENESIS © 7 university car
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VIVA PANGENESIS © 9 banish obsessi
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VIVA PANGENESIS © 11 Military Coll
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VIVA PANGENESIS © 13 tions of pang
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VIVA PANGENESIS © 15 Still playing
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VIVA PANGENESIS © 17 Although Galt
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VIVA PANGENESIS © 19 and Louisa tr
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VIVA PANGENESIS © 21 tion]. What l
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CHAPTER 2 Reversion to the Mean ¨
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REVERSION TO THE MEAN © 25 a price
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REVERSION TO THE MEAN © 27 the pop
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REVERSION TO THE MEAN © 29 In fact
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REVERSION TO THE MEAN © 31 jects h
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[To view this image, refer to the p
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REVERSION TO THE MEAN © 37 regular
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REVERSION TO THE MEAN © 41 self ha
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REVERSION TO THE MEAN © 43 cape re
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CHAPTER 3 Galton’s Disciples ¨ I
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GALTON’S DISCIPLES © 47 In the s
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GALTON’S DISCIPLES © 49 fection
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GALTON’S DISCIPLES © 51 treme va
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GALTON’S DISCIPLES © 53 popular
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GALTON’S DISCIPLES © 55 species
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GALTON’S DISCIPLES © 57 see the
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GALTON’S DISCIPLES © 59 ety meet
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GALTON’S DISCIPLES © 61 that no
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GALTON’S DISCIPLES © 63 lished r
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GALTON’S DISCIPLES © 65 lights m
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70 ¨ PANGENES he took a position a
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72 ¨ PANGENES In the following yea
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74 ¨ PANGENES his own. In the plac
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76 ¨ PANGENES transported from the
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78 ¨ PANGENES sized: The first he
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80 ¨ PANGENES tical study of the l
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82 ¨ PANGENES ber,” which posite
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84 ¨ PANGENES peared in April 1900
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88 ¨ MENDEL younger Mendel having
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90 ¨ MENDEL lower grades. The phys
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92 ¨ MENDEL brids, following the i
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94 ¨ MENDEL invisible, and the sam
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96 ¨ MENDEL A less sinister interp
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98 ¨ MENDEL way had their position
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100 ¨ MENDEL In the same paper, Me
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102 ¨ MENDEL sequestration if Mend
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106 ¨ REDISCOVERY planning to publ
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108 ¨ REDISCOVERY which each pange
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Page 155 and 156: 140 ¨ MENDEL WARS his research, Hu
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Page 181 and 182: 166 ¨ SEX CHROMOSOMES were two kin
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OENOTHERA RECONSIDERED © 211 winge
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OENOTHERA RECONSIDERED © 213 succe
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OENOTHERA RECONSIDERED © 215 tatio
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OENOTHERA RECONSIDERED © 217 compl
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OENOTHERA RECONSIDERED © 219 outcr
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CHAPTER 12 X-Rays ¨ WITH THE RESOL
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X-RAYS © 223 through, due to the i
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X-RAYS © 225 Suddenly Muller jumpe
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X-RAYS © 227 C � B experiments a
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X-RAYS © 229 Bateson’s plenary a
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X-RAYS © 231 Bateson’s famous fo
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X-RAYS © 233 Cramped, noisy, overh
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X-RAYS © 235 ing. “It was funny
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X-RAYS © 237 birth of the baby, wh
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X-RAYS © 239 spontaneous lethals (
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X-RAYS © 241 the Fifth Internation
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CHAPTER 13 Triumph of the Modern Ge
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TRIUMPH OF THE MODERN GENE © 245 D
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TRIUMPH OF THE MODERN GENE © 247 l
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TRIUMPH OF THE MODERN GENE © 249 w
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TRIUMPH OF THE MODERN GENE © 251 t
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+ + B f B f + B B fu + B B + f + TR
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TRIUMPH OF THE MODERN GENE © 255 f
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TRIUMPH OF THE MODERN GENE © 259 F
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TRIUMPH OF THE MODERN GENE © 263 O
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TRIUMPH OF THE MODERN GENE © 265 m
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TRIUMPH OF THE MODERN GENE © 269 p
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TRIUMPH OF THE MODERN GENE © 271 p
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TRIUMPH OF THE MODERN GENE © 273 s
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EPILOGUE ¨ IN FEBRUARY 1944, Oswal
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locks were laid down that accounted
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EPILOGUE © 281 In November 1951, W
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EPILOGUE © 283 The discovery in 19
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EPILOGUE © 285 characterization of
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EPILOGUE © 287 master gene on chro
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EPILOGUE © 289 the genome could be
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NOTES ACKNOWLEDGMENTS INDEX
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294 ¨ NOTES TO PAGES 4-8 help in p
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296 ¨ NOTES TO PAGES 19-28 58. Let
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298 ¨ NOTES TO PAGES 36-42 35. Let
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300 ¨ NOTES TO PAGES 50-58 F.R.S.
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302 ¨ NOTES TO PAGES 70-81 2. Ibid
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304 ¨ NOTES TO PAGES 84-89 27. Ibi
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306 ¨ NOTES TO PAGES 93-95 29. The
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308 ¨ NOTES TO PAGES 106-111 (1900
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310 ¨ NOTES TO PAGES 114-120 24. I
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312 ¨ NOTES TO PAGES 125-132 15. A
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314 ¨ NOTES TO PAGES 137-146 55. W
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316 ¨ NOTES TO PAGES 150-153 15. E
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318 ¨ NOTES TO PAGES 157-160 38. M
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320 ¨ NOTES TO PAGES 167-171 6. Ib
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322 ¨ NOTES TO PAGES 173-176 33. A
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324 ¨ NOTES TO PAGES 182-186 of Tw
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326 ¨ NOTES TO PAGES 190-192 multi
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330 ¨ NOTES TO PAGES 202-210 Franc
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332 ¨ NOTES TO PAGES 213-215 13. I
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334 ¨ NOTES TO PAGES 218-222 28. B
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336 ¨ NOTES TO PAGES 225-228 14. T
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338 ¨ NOTES TO PAGES 235-239 “Ob
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340 ¨ NOTES TO PAGES 245-254 9. H.
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342 ¨ NOTES TO PAGES 258-262 51. J
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344 ¨ NOTES TO PAGES 266-272 84. S
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346 ¨ NOTES TO PAGES 276-283 123.
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348 ¨ NOTES TO PAGES 288-289 Becau
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350 ¨ ACKNOWLEDGMENTS also deeply
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“Accessory Chromosome, The: Sex D
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ell curve. See normal distribution
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34, 50, 60-61; and study of plant m
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“fly room.” See Columbia Univer
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ics; evolution; gemmules; genes; hy
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y, 306n37; early life of, 87-88; fu
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222-223; Russian geneticists and, 2
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studies of, 79-85; Wichura and, 100
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Stalin, Joseph, 258, 262, 267, 269,
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In Pursuit of the Gene

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