性别分配及鸟类子代性比研究进展
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摘要
性别分配涉及物种生存策略、个体适合度以及种群动态等多方面因素影响,一直受到研究人员的广泛关注.鸟类是性别分配研究的理想对象.当前,有关鸟类性别分配已有较多研究,其中,子代性比作为性别分配最直接的观测指标受到研究者的普遍关注,其影响因素对于性别分配问题的研究具有指向性的意义,也因此成为该领域的一个研究热点.有关子代性比影响因素的研究主要集中在环境、亲代和子代三大方面,但是目前对于关键影响因素缺乏一般性结论,亲鸟对子代性比的控制能力及其内在作用机制也尚不明确,相关问题仍需不断探索.本文以性别分配研究得以发展的4个重要观点作为基础,对鸟类子代性比的相关研究及其进展进行了评述.
Sex allocation has caught much attention because it is related to life strategies,individual fitness and population dynamics.Birds,a class of vertebrates with extremely high species diversity and abundance,are ideal subjects for the study of sex allocation.Several classic hypotheses have been proposed,such as evolutionary stable strategy and Trivers-Willard hypothesis.Offspring sex ratio is a direct observation of sex allocation,so the factors influencing offspring sex ratio are key to the understanding of sex allocation between males and females.Currently,the field focuses on the research of environmental,parental and offspring factors.However,up to date it is still an open question which factor determines the offspring sex ratio,and much work needs to be done before we know to what extent parents can regulate offspring sexes.In this review,we summarize the progress in the factors influencing the offspring sex ratio,presenting a spectrum of offspring sex ratio studies from the classic hypotheses to empirical studies.
Sex allocation has caught much attention because it is related to life strategies,individual fitness and population dynamics.Birds,a class of vertebrates with extremely high species diversity and abundance,are ideal subjects for the study of sex allocation.Several classic hypotheses have been proposed,such as evolutionary stable strategy and Trivers-Willard hypothesis.Offspring sex ratio is a direct observation of sex allocation,so the factors influencing offspring sex ratio are key to the understanding of sex allocation between males and females.Currently,the field focuses on the research of environmental,parental and offspring factors.However,up to date it is still an open question which factor determines the offspring sex ratio,and much work needs to be done before we know to what extent parents can regulate offspring sexes.In this review,we summarize the progress in the factors influencing the offspring sex ratio,presenting a spectrum of offspring sex ratio studies from the classic hypotheses to empirical studies.
引文
1 Fisher R A.The Genetical Theory of Natural Selection.Oxford:Oxford University Press,1930
2 Darwin C.The Descent of Man and Selection In Relation To Sex.2nd ed.London:John Murray,1871
3 Kokko H,Jennions M D.Parental investment,sexual selection and sex ratios.J Evol Biol,2008,21:919-948
4 Li J Q.The breeding behavior and sex ratio of black-throated tit(Aegithalos concinnus)and long-tailed tit(Aeigithalos caudatus).Dissertation.Beijing:Beijing Normal Univ,2010[李建强.红头长尾山雀(Aegithalos concinnus)和银喉长尾山雀(Aeigithalos caudatus)的繁殖行为与性比的研究.博士学位论文.北京:北京师范大学,2010]
5 Booksmythe I,Mautz B,Davis J,et al.Facultative adjustment of the offspring sex ratio and male attractiveness:a systematic review and metaanalysis.Biol Rev,2017,92:108-134
6 Charnov E L.Simultaneous hermaphroditism and sexual selection.Proc Natl Acad Sci USA,1979,76:2480-2484
7 Komdeur J,Pen I.Adaptive sex allocation in birds:the complexities of linking theory and practice.Philos Trans R Soc Lond B Biol,2002,357:373-380
8 Donald P F.Adult sex ratios in wild bird populations.Ibis,2007,149:671-692
9 Benvenuti B,Walsh J,O’Brien K M,et al.Annual variation in the offspring sex ratio of saltmarsh sparrows supports Fisher’s hypothesis.Auk,2018,135:342-358
10 Vanthournout B,Busck M M,Bechsgaard J,et al.Male spiders control offspring sex ratio through greater production of female-determining sperm.Proc R Soc B,2018,285:20172887
11 Hamilton W D.Extraordinary sex ratios.Science,1967,156:477-488
12 Trivers R L,Willard D E.Natural selection of parental ability to vary the sex ratio of offspring.Science,1973,179:90-92
13 West S A.Sex Allocation.Princeton:Princeton University Press,2009
14 Smith J M,Price G R.The logic of animal conflict.Nature,1973,246:15-18
15 Hamilton W D,May R M.Dispersal in stable habitats.Nature,1977,269:578-581
16 Greenwood P J.Mating systems,philopatry and dispersal in birds and mammals.Anim Behav,1980,28:1140-1162
17 Lessells C M.Natal and breeding dispersal of Canada geese Branta canadensis.Ibis,1985,127:31-41
18 Gowaty P A.Differential dispersal,local resource competition,and sex ratio variation in birds.Am Nat,1993,141:263-280
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20 Komdeur J,Daan S,Tinbergen J,et al.Extreme adaptive modification in sex ratio of the Seychelles warbler’s eggs.Nature,1997,385:522-525
21 Charnov E L,Bull J J,Maynard Smith J.Why be an hermaphrodite?Nature,1976,263:125-126
22 Charnov E L,Los-den Hartogh R L,Jones W T,et al.Sex ratio evolution in a variable environment.Nature,1981,289:27-33
23 West S A,Sheldon B C.Constraints in the evolution of sex ratio adjustment.Science,2002,295:1685-1688
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25 Trivers R L,Hare H.Haploidploidy and the evolution of the social insect.Science,1976,191:249-263
26 Hamilton W D.The genetical evolution of social behavior.II.J Theor Biol,1964,7:17-52
27 Sheldon B C.Recent studies of avian sex ratios.Heredity,1998,80:397-402
28 Pen I.Sex Allocation:In a Life History Context.Groningen:Rijksuniversiteit Groningen,2000
29 Charnov E L.The Theory of Sex Allocation.Princeton:Princeton University Press,1982
30 Clutton-brock T H.Sex ratio variation in birds.Ibis,2008,128:317-329
31 Bull J J,Charnov E.How fundamental are Fisherian sex ratios?Oxf Surv Evol Bio,1988,5:96-135
32 Hasselquist D,Kempenaers B.Parental care and adaptive brood sex ratio manipulation in birds.Philos Trans R Soc Lond B Biol,2002,357:363-372
33 Ding Z,Ji F,Huang Q,et al.Brood sex ratio in the Yellow-bellied Prinia(Prinia flaviventris).Avian Res,2017,8:15
34 Griffiths R,Double M C,Orr K,et al.A DNA test to sex most birds.Mol Ecol,1998,7:1071-1075
35 Alonso-Alvarez C.Manipulation of primary sex-ratio:an updated review.Avian Poul Biol Rev,2006,17:1-20
36 Stubblefield J W,Orzack S H.Resource transfers and evolution:helpful offspring and sex allocation.Theor Popul Biol,2013,83:64-81
37 Ewen J G,Cassey P,M?ller A P.Facultative primary sex ratio variation:a lack of evidence in birds?Proc R Soc B,2004,271:1277-1282
38 Mainwaring M C,Lucy D,Hartley I R.Parentally biased favouritism in relation to offspring sex in zebra finches.Behav Ecol Sociobiol,2011,65:2261-2268
39 Dijkstra C,Daan S,Buker J B.Adaptive seasonal variation in the sex ratio of kestrel broods.Funct Ecol,1990,4:143-147
40 Olsent P D,Cockburn A.Female-biased sex allocation in peregrine falcons and other raptors.Behav Ecol Sociobiol,1991,28:417-423
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47 Andersson M,Wallander J,Oring L,et al.Adaptive seasonal trend in brood sex ratio:test in two sister species with contrasting breeding systems.J Evol Biol,2003,16:510-515
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2 Darwin C.The Descent of Man and Selection In Relation To Sex.2nd ed.London:John Murray,1871
3 Kokko H,Jennions M D.Parental investment,sexual selection and sex ratios.J Evol Biol,2008,21:919-948
4 Li J Q.The breeding behavior and sex ratio of black-throated tit(Aegithalos concinnus)and long-tailed tit(Aeigithalos caudatus).Dissertation.Beijing:Beijing Normal Univ,2010[李建强.红头长尾山雀(Aegithalos concinnus)和银喉长尾山雀(Aeigithalos caudatus)的繁殖行为与性比的研究.博士学位论文.北京:北京师范大学,2010]
5 Booksmythe I,Mautz B,Davis J,et al.Facultative adjustment of the offspring sex ratio and male attractiveness:a systematic review and metaanalysis.Biol Rev,2017,92:108-134
6 Charnov E L.Simultaneous hermaphroditism and sexual selection.Proc Natl Acad Sci USA,1979,76:2480-2484
7 Komdeur J,Pen I.Adaptive sex allocation in birds:the complexities of linking theory and practice.Philos Trans R Soc Lond B Biol,2002,357:373-380
8 Donald P F.Adult sex ratios in wild bird populations.Ibis,2007,149:671-692
9 Benvenuti B,Walsh J,O’Brien K M,et al.Annual variation in the offspring sex ratio of saltmarsh sparrows supports Fisher’s hypothesis.Auk,2018,135:342-358
10 Vanthournout B,Busck M M,Bechsgaard J,et al.Male spiders control offspring sex ratio through greater production of female-determining sperm.Proc R Soc B,2018,285:20172887
11 Hamilton W D.Extraordinary sex ratios.Science,1967,156:477-488
12 Trivers R L,Willard D E.Natural selection of parental ability to vary the sex ratio of offspring.Science,1973,179:90-92
13 West S A.Sex Allocation.Princeton:Princeton University Press,2009
14 Smith J M,Price G R.The logic of animal conflict.Nature,1973,246:15-18
15 Hamilton W D,May R M.Dispersal in stable habitats.Nature,1977,269:578-581
16 Greenwood P J.Mating systems,philopatry and dispersal in birds and mammals.Anim Behav,1980,28:1140-1162
17 Lessells C M.Natal and breeding dispersal of Canada geese Branta canadensis.Ibis,1985,127:31-41
18 Gowaty P A.Differential dispersal,local resource competition,and sex ratio variation in birds.Am Nat,1993,141:263-280
19 Legge S,Heinsohn R,Double M C,et al.Complex sex allocation in the laughing kookaburra.Behav Ecol,2001,12:524-533
20 Komdeur J,Daan S,Tinbergen J,et al.Extreme adaptive modification in sex ratio of the Seychelles warbler’s eggs.Nature,1997,385:522-525
21 Charnov E L,Bull J J,Maynard Smith J.Why be an hermaphrodite?Nature,1976,263:125-126
22 Charnov E L,Los-den Hartogh R L,Jones W T,et al.Sex ratio evolution in a variable environment.Nature,1981,289:27-33
23 West S A,Sheldon B C.Constraints in the evolution of sex ratio adjustment.Science,2002,295:1685-1688
24 Benito M M,Schielzeth H,González-Solís J,et al.Sex ratio adjustments in common terns:influence of mate condition and maternal experience.JAvian Biol,2013,44:179-188
25 Trivers R L,Hare H.Haploidploidy and the evolution of the social insect.Science,1976,191:249-263
26 Hamilton W D.The genetical evolution of social behavior.II.J Theor Biol,1964,7:17-52
27 Sheldon B C.Recent studies of avian sex ratios.Heredity,1998,80:397-402
28 Pen I.Sex Allocation:In a Life History Context.Groningen:Rijksuniversiteit Groningen,2000
29 Charnov E L.The Theory of Sex Allocation.Princeton:Princeton University Press,1982
30 Clutton-brock T H.Sex ratio variation in birds.Ibis,2008,128:317-329
31 Bull J J,Charnov E.How fundamental are Fisherian sex ratios?Oxf Surv Evol Bio,1988,5:96-135
32 Hasselquist D,Kempenaers B.Parental care and adaptive brood sex ratio manipulation in birds.Philos Trans R Soc Lond B Biol,2002,357:363-372
33 Ding Z,Ji F,Huang Q,et al.Brood sex ratio in the Yellow-bellied Prinia(Prinia flaviventris).Avian Res,2017,8:15
34 Griffiths R,Double M C,Orr K,et al.A DNA test to sex most birds.Mol Ecol,1998,7:1071-1075
35 Alonso-Alvarez C.Manipulation of primary sex-ratio:an updated review.Avian Poul Biol Rev,2006,17:1-20
36 Stubblefield J W,Orzack S H.Resource transfers and evolution:helpful offspring and sex allocation.Theor Popul Biol,2013,83:64-81
37 Ewen J G,Cassey P,M?ller A P.Facultative primary sex ratio variation:a lack of evidence in birds?Proc R Soc B,2004,271:1277-1282
38 Mainwaring M C,Lucy D,Hartley I R.Parentally biased favouritism in relation to offspring sex in zebra finches.Behav Ecol Sociobiol,2011,65:2261-2268
39 Dijkstra C,Daan S,Buker J B.Adaptive seasonal variation in the sex ratio of kestrel broods.Funct Ecol,1990,4:143-147
40 Olsent P D,Cockburn A.Female-biased sex allocation in peregrine falcons and other raptors.Behav Ecol Sociobiol,1991,28:417-423
41 Daan S,Dijkstra C,Weissing F J.An evolutionary explanation for seasonal trends in avian sex ratios.Behav Ecol,1996,7:426-430
42 Smallwood P D,Smallwood J.Seasonal shifts in sex ratios of fledgling American kestrels(Falco sparverius paulus):the early bird hypothesis.Evol Ecol,1998,12:839-853
43 Mora O,del Mar Delgado M,Penteriani V.Secondary sex ratio in Eurasian Eagle-owls:early-breeding females produce more daughters.J Raptor Res,2010,44:62-65
44 Howe H F.Sex-ratio adjustment in the common grackle.Science,1977,198:744-746
45 Korpimaki E,May C A,Parkin D T,et al.Environmental-and parental condition-related variation in sex ratio of kestrel broods.J Avian Biol,2000,31:128-134
46 Que P J.Breeding success and population genetic structure of Kentish plover(Charadrius alexandrinus)in China(in Chinese).Dissertation for Doctoral Degree.Beijing:Beijing Normal Univ,2015[阙品甲.中国环颈鸻(Charadrius alexandrinus)的繁殖成功率与种群遗传结构研究.博士学位论文.北京:北京师范大学,2015]
47 Andersson M,Wallander J,Oring L,et al.Adaptive seasonal trend in brood sex ratio:test in two sister species with contrasting breeding systems.J Evol Biol,2003,16:510-515
48 Chen W.Breeding strategies of Tibetan ground tit,(Pseudopodoces humilis)from the northern Tibetan Plateau(in Chinese).Dissertation for Doctoral Degree.Wuhan:Wuhan Univ,2011[陈伟.藏北高原地山雀繁殖策略研究.博士学位论文.武汉:武汉大学,2011]
49 Szekely T,Cuthill I C,Yezerinac S,et al.Brood sex ratio in the Kentish plover.Behav Ecol,2004,15:58-62
50 Suorsa P,Helle H,Huhta E,et al.Forest fragmentation is associated with primary brood sex ratio in the treecreeper(Certhia familiaris).Proc RSoc B,2003,270:2215-2222
51 Rutz C.Brood sex ratio varies with diet composition in a generalist raptor.Biol J Linn,2012,105:937-951
52 Katzner T E,Jackson D S,Ivy J,et al.Variation in offspring sex ratio of a long-lived sexually dimorphic raptor,the Eastern Imperial Eagle Aquila heliaca.Ibis,2014,156:395-403
53 Santoro S,Green A J,Speakman J R,et al.Facultative and non-facultative sex ratio adjustments in a dimorphic bird species.Oikos,2015,124:1215-1224
54 Prior G L,Evans D M,Redpath S,et al.Birds bias offspring sex ratio in response to livestock grazing.Biol Lett,2011,7:958-960
55 Bouvier J C,Boivin T,Charmantier A,et al.More daughters in a less favourable world:breeding in intensively-managed orchards affects tertiary sex-ratio in the great tit.Basic Appl Ecol,2016,17:638-647
56 Bell S C,Owens I P F,Lord A M.Quality of breeding territory mediates the influence of paternal quality on sex ratio bias in a free-living bird population.Behav Ecol,2014,25:352-358
57 Griffin A S,Sheldon B C,West S A.Cooperative breeders adjust offspring sex ratios to produce helpful helpers.Am Nat,2005,166:628-632
58 Ewen J G,Crozier R H,Cassey P,et al.Facultative control of offspring sex in the cooperatively breeding bell miner,Manorina melanophrys.Behav Ecol,2003,14:157-164
59 Khwaja N,Hatchwell B J,Freckleton R P,et al.Sex allocation patterns across cooperatively breeding birds do not support predictions of the repayment hypothesis.Am Nat,2017,190:547-556
60 Valone T J,Templeton J J.Public information for the assessment of quality:a widespread social phenomenon.Philos Trans R Soc Lond B Biol,2002,357:1549-1557
61 Qin Y F.Effects of“Public information”on offspring sex ratio in great tit(Parus major)(in Chinese).Dissertation for Master’s Degree.Changchun:Northeast Normal Univ,2010[秦玉芳.“公众信息”对大山雀子代性比的影响.硕士学位论文.长春:东北师范大学,2010]
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