종분화 실험

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맥아당과 전분 배지에서 두 줄의 초파리를 사육하는 동종성 종분화 실험

종분화에 대한 실험은 잡종화, 강화, 창시자 효과 등 모두에 대해 수행되었다. 대부분의 실험은 파리, 특히 초파리로 이루어졌다.[1] 그러나 더 최근의 연구에서는 효모, 곰팡이, 심지어 바이러스까지도 이용하여 실험했다.

실험실 실험(Laboratory experiment)은 개체군 크기가 작고 세대가 제한되어 있기 때문에 변종 종분화(동종 및 주변)에 도움이 되지 않는다고 제안되었다.[2] 자연 연구에서 얻은 대부분의 추정치는 종분화에 수십만 년에서 수백만 년이 걸린다는 것을 나타낸다.[3] 다른 한편으로, 많은 종은 3000세대 미만에서 동종분화를 하고 있는 원양, 해저 지역에서 산란하는 유럽 가자미(Platichthys flesus)와 같이 더 빠르고 최근에 종분화한 것으로 생각된다.[4]

실험 목록

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현재 확인할 수 있는 6개의 출판물이 종분화에 대한 실험적 연구를 편집, 검토 및 분석하려고 시도했다.

  1. 1985년 존 링고, 데이비드 우드, 로버트 록웰, 해롤드 다우즈[5]
  2. 1993년 William R. Rice 와 Ellen E.[6]
  3. 2002년 Ann-Britt Florin 과 Anders Ödeen.[7]
  4. 2002년 Mark Kirkpatrick 과 Virginie Ravigné.[8]
  5. 2004년 Jerry A. Coyne 및 H. Allen Orr[1]
  6. 2009년 제임스 D. 프라이.[9]

표

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이 표는 이 간행물에서 검토한 연구 및 데이터를 요약한 것이다. 또한 여러 현대적 실험을 참조하며 포괄적이지 않다.

표에서 세대 열에서 세미콜론으로 구분된 여러 숫자는 여러 실험이 수행되었음을 나타낸다. 복제(괄호 안)는 실험에 사용된 모집단의 수, 즉 실험이 복제된 횟수를 나타낸다. 다양한 유형의 선택이 실험 모집단에 부과되었으며 선택 유형 열로 표시된다. 각 실험의 음성 또는 양성 결과는 생식 분리 컬럼에 의해 제공된다.

접합 전 생식 격리(Pre-zygotic)는 개체군에서 생식하는 개체가 자손을 낳을 수 없음을 의미한다(효과적으로 긍정적인 결과). 접합 후 격리(Post-zygotic)는 번식하는 개체가 자손을 낳을 수 있었지만 불임 상태이거나 생존할 수 없었음을 의미한다(양성 결과도 마찬가지임). 음성 결과는 "없음"으로 표시된다. 즉, 실험에서 생식적 격리가 발생하지 않았다는 뜻이다.

표의 내용과 용어는 영문판을 그대로 사용하였다.

종분화 실험에 대한 표[1][10][11][12][9]
종 특성 세대 테스트 선택형 유전적 부동 생식적 격리 출처 실험한 해
Drosophila melanogaster Escape response 18 Vicariant, reinforcement, parapatric/

sympatric

Indirect; divergent Yes Pre-zygotic Grant & Mettler[13] 1969
D. melanogaster Locomotion 112 Vicariant Indirect; divergent No Pre-zygotic Burnet & Connolly[14] 1974
D. melanogaster Temperature, humidity 70–130 Vicariant Indirect; divergent Yes Pre-zygotic Kilias et al.[15] 1980
D. melanogaster DDT adaptation 600 [25 years, +15 years] Vicariant Direct No Pre-zygotic Boake et al.[16] 2003
D. melanogaster 17, 9, 9, 1, 1, 7, 7, 7, 7 Vicariant; parapatric/

sympatric

Direct, divergent Pre-zygotic in vicariance; none with gene flow Barker & Karlsson[17] 1974
D. melanogaster 40; 50 Reinforcement Direct; divergent Pre-zygotic Crossley[18] 1974
D. melanogaster Locomotion 45 Vicariant Direct; divergent No None van Dijken & Scharloo[19][20] 1979
D. melanogaster Reinforcement Direct; divergent Pre-zygotic Wallace[21] 1953
D. melanogaster 36; 31 Reinforcement Direct; divergent Pre-zygotic Knight[22] 1956
D. melanogaster EDTA adaptation 25, 25, 25, 14 Semi-allopatric, reinforcement Indirect; divergent No Post-zygotic Robertson[23][24] 1966
D. melanogaster 25 (8) Vicariant; reinforcement; parapatric; sympatric Direct None Hostert[25] 1997
D. melanogaster Abdominal chaeta

number

21–31 Vicariant Direct Yes None Santibanez & Waddington[26] 1958
D. melanogaster Sternopleural chaeta number 32 Vicariant, reinforcement, parapatric/

sympatric

Direct No None Barker & Cummins[27] 1969
D. melanogaster Phototaxis, geotaxis 20 Vicariant No None Markow[28][29] 1975; 1981
D. melanogaster Peripatric Yes Rundle et al.[30] 1998
D. melanogaster Vicariant; peripatric Yes Mooers et al.[31] 1999
D. melanogaster 12 Reinforcement Divergent Pre-zygotic Thoday & Gibson[32] 1962
D. melanogaster None Thoday & Gibson[33][34] 1970; 1971
D. melanogaster 16 Reinforcement Indirect None Spiess & Wilke[35] 1954
D. melanogaster Reinforcement Direct; divergent Pre-zygotic Ehrman[36][37][38][39] 1971; 1973; 1979; 1983
D. melanogaster Sternopleural chaeta number 5; 27; 27; 1; 1; 1; 1; 1 Parapatric/

sympatric

None Chabora[40] 1968
D. melanogaster None Scharloo[41] 1967
D. melanogaster 1, 1 Coyne & Grant[42] 1972
D. melanogaster 25 Rice[43] 1985
D. melanogaster 25 Disruptive Pre-zygotic Rice & Salt[44] 1988
D. melanogaster 35; 35 Sympatric Pre-zygotic Rice & Salt[45] 1990
D. melanogaster NaCl and CuSO4 levels in food [3 years in allopatry, 1 in sympatry] Allopatric; reinforcement; sympatric Pre-zygotic in allopatry, none in sympatry Wallace[46] 1982
D. melanogaster Reinforcement Ehrman et al.[47] 1991
D. melanogaster Reinforcement Fukatami & Moriwaki[48] 1970
Drosophila simulans Scutellar bristles, development speed, wing width; desiccation resistance, fecundity, ethanol resistance; courtship display, re-mating speed, lek behavior; pupation height, clumped egg laying, general activity [3 years] Vicariant; peripatric Yes Post-zygotic Ringo et al.[49] 1985
Drosophila paulistorum 131; 131 Reinforcement Direct Pre-zygotic Dobzhansky et al.[50] 1976
D. paulistorum [5 years] Vicariant Dobzhansky and Pavlovsky[51] 1966
Drosophila willistoni pH adaptation 34–122 Vicariant Indirect; divergent No Pre-zygotic Kalisz & Cordeiro[52] 1980
Drosophila pseudoobscura Carbohydrate source 12 Vicariant Indirect Yes Pre-zygotic Dodd[53] 1989
D. pseudoobscura Temperature adaptation 25–60 Vicariant Direct Ehrman[54][55][56][57][58] 1964;

1969

D. pseudoobscura Phototaxis, geotaxis 5–11 Vicariant Indirect No Pre-zygotic del Solar[59] 1966
D. pseudoobscura Vicariant; peripatric Pre-zygotic Powell[60][61] 1978; 1985
D. pseudoobscura Peripatric; vicariant Yes Galiana et al.[62] 1993
D. pseudoobscura Temperature photoperiod; food 37 (78) [33–34 months] Vicariant Divergent Yes None Rundle[63] 2003
D. pseudoobscura &

Drosophila persimilis

22; 16; 9 Reinforcement Direct; divergent Pre-zygotic Koopman[64] 1950
D. pseudoobscura &

D. persimilis

18 (4) Direct Pre-zygotic Kessler[65] 1966
Drosophila mojavensis 12 Direct Pre-zygotic Koepfer[66] 1987
D. mojavensis Development time 13 Divergent Yes None Etges[67] 1998
Drosophila adiastola Peripatric Yes Pre-zygotic Arita & Kaneshiro[68] 1974
Drosophila silvestris Peripatric Yes Ahearn[69] 1980
Musca domestica Geotaxis 38 Vicariant Indirect No Pre-zygotic Soans et al.[70] 1974
M. domestica Geotaxis 16 Vicariant Direct; divergent No Pre-zygotic Hurd & Eisenburg[71] 1975
M. domestica Peripatric Yes Meffert & Bryant[72] 1991
M. domestica Regan et al.[73] 2003
Bactrocera cucurbitae Development time 40–51 Divergent Yes Pre-zygotic Miyatake & Shimizu[74] 1999
Zea mays 6; 6 Reinforcement Direct; divergent Pre-zygotic Paterniani[75] 1969
Drosophila grimshawi Peripatric Jones, Widemo, & Arrendal[76] N/A
Saccharomyces cerevisiae Leu & Murry[77] 2006
D. melanogaster Reinforcement Harper & Lambert[78] 1983
Tribolium castaneum Pupal weight 15 (6) Disruptive Halliburton & Gall[79] 1983
D. melanogaster Geotaxis Divergent Lofdahl et al.[80] 1992
D. pseudoobscura [10 years] Moya et al.[81] 1995
Neurospora Divergent Dettman et al.[82] 2008
S. cerevisiae 500 Divergent Dettman et al.[83] 2007
Sepsis cynipsea 35 Martin & Hosken[84] 2003
D. melanogaster Wigby & Chapman[85] 2006
D. pseudoobscura Sexual conflict 48–52 (4; 4; 4) Bacigalupe et al.[86] 2007
D. serrata Rundle et al.[87] 2005
Drosophila serrata & D. birchii Mate recognition 9 (3; 3) Reinforcement Natural Pre-zygotic Higgie et al.[88] 2000
Enterobacteria phage λ Escherichia coli receptor exploitation 35 cycles (6) Vicariant, sympatric Pre-zygotic Meyer et al.[89] 2016
Tetranychus urticae Resistance to host plant toxin Overmeer[90] 1966
T. urticae Resistance to host plant toxin Fry[91] 1999
Helianthus annus × H. petiolaris and H. anomalus Hybrid Rieseburg et al.[92] 1996
S. cerevisiae Greig et al.[93] 2002
D. melanogaster Life history Ghosh & Joshi[94] 2012
Drosophila subobscura Mate behavior Bárbaro et al.[95] 2015
Digital organisms ~42,000; ~850 (20) Ecological Post-zygotic Anderson & Harmon[96] 2014
Schizosaccharomyces pombe Complete reproductive isolation Seike et al.[97] 2015
D. pseudoobscura Courtship song 130 Debelle et al.[98] 2014
Callosobruchus maculatus 40 (16) Debelle et al.[99] 2010

각주

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  1. ↑ 가 나 다 , Sinauer Associates  |제목=이(가) 없거나 비었음 (도움말)
  2. ↑ Florin, Ann-Britt & Ödeen, Anders (2002), “Laboratory environments are not conducive for allopatric speciation”, 《Journal of Evolutionary Biology》 15 (1): 10–19, doi:10.1046/j.1420-9101.2002.00356.x 
  3. ↑ Coyne, Jerry A.; Orr, H. Allen (1997), “"Patterns of Speciation in Drosophila" Revisited”, 《Evolution》 51 (1): 295–303, doi:10.1111/j.1558-5646.1997.tb02412.x, PMID 28568795 
  4. ↑ Momigliano, Paolo; Jokinen, Henri; Fraimout, Antoine; Florin, Ann-Britt; Norkko, Alf; Merilä, Juha (2017), “Extraordinarily rapid speciation in a marine fish” (PDF), 《PNAS》 114 (23): 6074–6079, doi:10.1073/pnas.1615109114, PMC 5468626, PMID 28533412 
  5. ↑ Ringo, John; Wood, David; Rockwell, Robert; Dowse, Harold (1985), “An Experiment Testing Two Hypotheses of Speciation”, 《The American Naturalist》 126 (5): 642–661, doi:10.1086/284445, S2CID 84819968 
  6. ↑ Rice, William R. & Hostert, Ellen E. (1993), “Laboratory Experiments on Speciation: What Have We Learned in 40 Years?”, 《Evolution》 47 (6): 1637–1653, doi:10.1111/j.1558-5646.1993.tb01257.x, PMID 28568007, S2CID 42100751 
  7. ↑ Florin, Ann-Britt & Ödeen, Anders (2002), “Laboratory environments are not conducive for allopatric speciation”, 《Journal of Evolutionary Biology》 15 (1): 10–19, doi:10.1046/j.1420-9101.2002.00356.x 
  8. ↑ Kirkpatrick, Mark & Ravigné, Virginie (2002), “Speciation by Natural and Sexual Selection: Models and Experiments”, 《The American Naturalist》 159: S22–S35, doi:10.1086/338370, PMID 18707367, S2CID 16516804 
  9. ↑ 가 나 Fry, James D. (2009). Laboratory Experiments on Speciation. In Garland, Theodore & Rose, Michael R. "Experimental Evolution: Concepts, Methods, and Applications of Selection Experiments". Pp. 631–656. doi:10.1525/california/9780520247666.003.0020
  10. ↑ Rice, William R. & Hostert, Ellen E. (1993), “Laboratory Experiments on Speciation: What Have We Learned in 40 Years?”, 《Evolution》 47 (6): 1637–1653, doi:10.1111/j.1558-5646.1993.tb01257.x, PMID 28568007, S2CID 42100751 
  11. ↑ Kirkpatrick, Mark & Ravigné, Virginie (2002), “Speciation by Natural and Sexual Selection: Models and Experiments”, 《The American Naturalist》 159: S22–S35, doi:10.1086/338370, PMID 18707367, S2CID 16516804 
  12. ↑ Florin, Ann-Britt & Ödeen, Anders (2002), “Laboratory environments are not conducive for allopatric speciation”, 《Journal of Evolutionary Biology》 15 (1): 10–19, doi:10.1046/j.1420-9101.2002.00356.x 
  13. ↑ Grant, B. S. & Mettler, L. E. (1969), “Disruptive and stabilizing selection on the" escape" behavior of Drosophila melanogaster”, 《Genetics》 62 (3): 625–637, doi:10.1093/genetics/62.3.625, PMC 1212303, PMID 17248452 
  14. ↑ Burnet, B. & Connolly, K. (1974). Activity and sexual behavior in Drosophila melanogaster. In Abeelen, J. H. V. F. (eds). The Genetics of Behaviour. North-Holland, Amsterdam. Pp. 201–258.
  15. ↑ Kilias, G., Alahiotis, S. N., & Pelecanos, M. (1980), “A Multifactorial Genetic Investigation of Speciation Theory Using Drosophila melanogaster”, 《Evolution》 34 (4): 730–737, doi:10.2307/2408027, JSTOR 2408027, PMID 28563991 
  16. ↑ Boake, C. R. B., Mcdonald, K., Maitra, S., Ganguly, R. (2003), “Forty years of solitude: life-history divergence and behavioural isolation between laboratory lines of Drosophila melanogaster”, 《Journal of Evolutionary Biology》 16 (1): 83–90, doi:10.1046/j.1420-9101.2003.00505.x, PMID 14635883 
  17. ↑ Barker, J. S. F. & Karlsson, L. J. E. (1974), “Effects of population size and selection intensity on responses to disruptive selection in Drosophila melanogaster”, 《Genetics》 78 (2): 715–735, doi:10.2307/2407287, JSTOR 2407287, PMC 1213230, PMID 4217303 
  18. ↑ Crossley, Stella A. (1974), “Changes in Mating Behavior Produced by Selection for Ethological Isolation Between Ebony and Vestigial Mutants of Drosophila melanogaster”, 《Evolution》 28 (4): 631–647, doi:10.1111/j.1558-5646.1974.tb00795.x, PMID 28564833, S2CID 35867118 
  19. ↑ van Dijken, F. R. & Scharloo, W. (1979), “Divergent selection on locomotor activity in Drosophila melanogaster. I. Selection response”, 《Behavior Genetics》 9 (6): 543–553, doi:10.1007/BF01067350, PMID 122270, S2CID 39352792 
  20. ↑ van Dijken, F. R. & Scharloo, W. (1979), “Divergent selection on locomotor activity in Drosophila melanogaster. II. Test for reproductive isolation between selected lines”, 《Behavior Genetics》 9 (6): 555–561, doi:10.1007/BF01067351, PMID 122271, S2CID 40169222 
  21. ↑ Wallace, B. (1953), “Genetic divergence of isolated populations of Drosophila melanogaster”, 《Proceedings of the Ninth International Congress of Genetics》 9: 761–764 
  22. ↑ Knight, G. R., Robertson, Alan, & Waddington, C. H. (1956), “Selection for sexual isolation within a species”, 《Evolution》 10 (1): 14–22, doi:10.1111/j.1558-5646.1956.tb02825.x 
  23. ↑ Robertson, Forbes W. (1966), “A test of sexual isolation in Drosophila”, 《Genetical Research》 8 (2): 181–187, doi:10.1017/S001667230001003X, PMID 5922518 
  24. ↑ Robertson, Forbes W. (1966), “The ecological genetics of growth in Drosophila 8. Adaptation to a New Diet”, 《Genetical Research》 8 (2): 165–179, doi:10.1017/S0016672300010028, PMID 5922517 
  25. ↑ Hostert, Ellen E. (1997), “Reinforcement: a new perspective on an old controversy”, 《Evolution》 51 (3): 697–702, doi:10.1111/j.1558-5646.1997.tb03653.x, PMID 28568598 
  26. ↑ Koref Santibañez, S. & Waddington, C. H. (1958), “The origin of sexual isolation between different lines within a species”, 《Evolution》 12 (4): 485–493, doi:10.2307/2405959, JSTOR 2405959 
  27. ↑ Barker, J. S. F. & Cummins, L. J. (1969), “The effect of selection for sternopleural bristle number in mating behaviour in Drosophila melanogaster”, 《Genetics》 61 (3): 713–719, doi:10.1093/genetics/61.3.713, PMC 1212235, PMID 17248436 
  28. ↑ Markow, Therese Ann (1975), “A genetic analysis of phototactic behavior in Drosophila melanogaster”, 《Genetics》 79 (3): 527–534, doi:10.1093/genetics/79.3.527, PMC 1213291, PMID 805084 
  29. ↑ Markow, Therese Ann (1981), “Mating preferences are not predictive of the direction of evolution in experimental populations of Drosophila”, 《Science》 213 (4514): 1405–1407, doi:10.1126/science.213.4514.1405, PMID 17732575, S2CID 15497733 
  30. ↑ Rundle, H. D., Mooers, Arne Ø. & Whitlock, Michael C. (1998), “Single founder-flush events and the evolution of reproductive isolation”, 《Evolution》 52 (6): 1850–1855, doi:10.1111/j.1558-5646.1998.tb02263.x, PMID 28565304, S2CID 24502821 
  31. ↑ Mooers, Arne Ø., Rundle, Howard D. & Whitlock, Michael C. (1999), “The effects of selection and bottlenecks on male mating success in peripheral isolates”, 《American Naturalist》 153 (4): 437–444, doi:10.1086/303186, PMID 29586617, S2CID 4411105 
  32. ↑ Thoday, J. M. & Gibson, J. B. (1962), “Isolation by disruptive selection”, 《Nature》 193 (4821): 1164–1166, doi:10.1038/1931164a0, PMID 13920720, S2CID 5156234 
  33. ↑ Thoday, J. M. & Gibson, J. B. (1970), “The probability of isolation by disruptive selection”, 《Nature》 104 (937): 219–230, doi:10.1086/282656, S2CID 85333360 
  34. ↑ Scharloo, W. (1971), “Reproductive isolation by disruptive selection: Did it occur?”, 《American Naturalist》 105 (941): 83–86, doi:10.1086/282706, S2CID 84204545 
  35. ↑ Spiess, E. B. & Wilke, C. M. (1984), “Still another attempt to achieve assortive mating by disruptive selection in Drosophila”, 《Evolution》 38 (3): 505–515, doi:10.2307/2408700, JSTOR 2408700, PMID 28555983 
  36. ↑ Ehrman, Lee (1971), “Natural selection and the origin of reproductive isolation”, 《American Naturalist》 105 (945): 479–483, doi:10.1086/282739, S2CID 85401244 
  37. ↑ Ehrman, Lee (1973), “More on natural selection and the origin of reproductive isolation”, 《American Naturalist》 107 (954): 318–319, doi:10.1086/282835, S2CID 83780632 
  38. ↑ Ehrman, Lee (1979), “Still more on natural selection and the origin of reproductive isolation”, 《American Naturalist》 113 (1): 148–150, doi:10.1086/283371, S2CID 85237458 
  39. ↑ Ehrman, Lee (1983), “Fourth report on natural selection for the origin of reproductive isolation”, 《American Naturalist》 121 (2): 290–293, doi:10.1086/284059, S2CID 83654887 
  40. ↑ Chabora, Alice J. (1968), “Disruptive selection for sternopleural chaeta number in various strains of Drosophila melanogaster”, 《American Naturalist》 102 (928): 525–532, doi:10.1086/282565, S2CID 84885812 
  41. ↑ Scharloo, W., Hoogmoed, M. S. & Kuile, A. T. (1967), “Stabilizing and disruptive selection on a mutant character in Drosophila. I. The phenotypic variance and its components.”, 《Genetics》 56 (4): 709–726, doi:10.1093/genetics/56.4.709, PMC 1211648, PMID 6061662 
  42. ↑ Coyne, Jerry A. & and Grant, Bruce (1972), “Disruptive selection on I-maze activity in Drosophila melanogaster”, 《Genetics》 71 (1): 185–188, doi:10.1093/genetics/71.1.185, PMC 1212770, PMID 17248572 
  43. ↑ Rice, W. R. (1985), “Disruptive selection on habitat preference and the evolution of reproductive isolation: an exploratory experiment”, 《Evolution》 39 (3): 645–656, doi:10.1111/j.1558-5646.1985.tb00401.x, PMID 28561974 
  44. ↑ Rice, William R. & Salt, George, W. (1988), “Speciation via disruptive selection on habitat preference”, 《American Naturalist》 131 (6): 911–917, doi:10.1086/284831, S2CID 84876223 
  45. ↑ Rice, William R. & Salt, George, W. (1990), “The evolution of reproductive isolation as a correlated character under sympatric conditions: experimental evidence”, 《Evolution》 44 (5): 1140–1152, doi:10.2307/2409278, JSTOR 2409278, PMID 28563894 
  46. ↑ Wallace, B. (1982), “Drosophila melanogaster populations selected for resistances to NaCl and CuSO4 in both allopatry and sympatry”, 《Journal of Heredity》 73 (1): 35–42, doi:10.1093/oxfordjournals.jhered.a109572, PMID 6802898 
  47. ↑ Ehrman, Lee, White, Marney A. & Wallace, B. (1991). A long-term study involving Drosophila melanogaster and toxic media. In Hecht, M. K., Wallace, B., & Maclntyre, R. J. (eds). Evolutionary biology, vol. 25. Plenum Press, New York. Pp. 175–209
  48. ↑ Fukatami, A & Moriwaki, D. (1970), “Selection for sexual isolation in Drosophila melanogaster by a modification of Koopman's method”, 《The Japanese Journal of Genetics》 45 (3): 193–204, doi:10.1266/jjg.45.193 
  49. ↑ Ringo, John; Wood, David; Rockwell, Robert; Dowse, Harold (1985), “An Experiment Testing Two Hypotheses of Speciation”, 《The American Naturalist》 126 (5): 642–661, doi:10.1086/284445, S2CID 84819968 
  50. ↑ Dobzhansky, Theodosius; Pavlovsky, O.; Powell, J. R. (1976), “Partially Successful Attempt to Enhance Reproductive Isolation Between Semispecies of Drosophila paulistorum”, 《Evolution》 30 (2): 201–212, doi:10.2307/2407696, JSTOR 2407696, PMID 28563045 
  51. ↑ Dobzhansky, Theodosius & Pavlovsky, O. (1966), “Spontaneous origin of an incipient species in the Drosophila paulistorum complex”, 《Proceedings of the National Academy of Sciences》 55 (4): 723–733, doi:10.1073/pnas.55.4.727, PMC 224220, PMID 5219677 
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