종분화 실험
생물학 관련 문서 |
진화생물학 |
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원리 |
자연의 역사 |
진화론의 역사 |
응용 분야 |
사회적 영향 |
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
현재 확인할 수 있는 6개의 출판물이 종분화에 대한 실험적 연구를 편집, 검토 및 분석하려고 시도했다.
- 1985년 존 링고, 데이비드 우드, 로버트 록웰, 해롤드 다우즈[5]
- 1993년 William R. Rice 와 Ellen E.[6]
- 2002년 Ann-Britt Florin 과 Anders Ödeen.[7]
- 2002년 Mark Kirkpatrick 과 Virginie Ravigné.[8]
- 2004년 Jerry A. Coyne 및 H. Allen Orr[1]
- 2009년 제임스 D. 프라이.[9]
표
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.
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 & | 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
- ↑ 가 나 다 , Sinauer Associates
|제목=
이(가) 없거나 비었음 (도움말) - ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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.
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ Wallace, B. (1953), “Genetic divergence of isolated populations of Drosophila melanogaster”, 《Proceedings of the Ninth International Congress of Genetics》 9: 761–764
- ↑ 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
- ↑ Robertson, Forbes W. (1966), “A test of sexual isolation in Drosophila”, 《Genetical Research》 8 (2): 181–187, doi:10.1017/S001667230001003X, PMID 5922518
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ Thoday, J. M. & Gibson, J. B. (1962), “Isolation by disruptive selection”, 《Nature》 193 (4821): 1164–1166, doi:10.1038/1931164a0, PMID 13920720, S2CID 5156234
- ↑ 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
- ↑ Scharloo, W. (1971), “Reproductive isolation by disruptive selection: Did it occur?”, 《American Naturalist》 105 (941): 83–86, doi:10.1086/282706, S2CID 84204545
- ↑ 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
- ↑ Ehrman, Lee (1971), “Natural selection and the origin of reproductive isolation”, 《American Naturalist》 105 (945): 479–483, doi:10.1086/282739, S2CID 85401244
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
- ↑ 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
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