紫萼（Hosta ventricosa）大小依赖性分配的初步研究
|
摘要
以一种顺序开花且每天只开一朵花(极少时开两朵)的多年生多次性生殖植物紫萼为研究材料,探讨了六个方面的问题:(1)是否存在一个繁殖的阈值;(2)个体水平上,果期性分配是否是大小依赖的;(3)花序内,性分配随位置的改变呈现怎样的变化;(4)雌性分配和雄性分配是否存在权衡关系;(5)种子大小与数量是否存在权衡关系(6)种群内种子大小变异以及引起变异的原因。主要研究结果如下:
1用个体总叶面积表征植株个体大小,分别对开花个体的叶长与不开花个体叶长以及两者的叶宽之间进行方差分析,结果它们的p值分别为:0.103和0.995。说明紫萼不存在一个大小繁殖的阈值。
2紫萼同一花序内,每花胚珠数在5个部位无显著变化(t=0.230,P=0.818),每花花粉数、P/O、结实率均从花序的基部到顶部逐渐下降(t=-6.435,P＜0.01;t=-7.493,P＜0.01;t=-3.967,p＜0.01),每果结籽率则从基部到顶部显著上升(t=2.453,p=0.15)。这可能与紫萼开花早期传粉昆虫较少有关。
3个体水平上,分析结果表明个体结实率不随个体总叶面积、个体高度、个体总花数的变化而变化。(r=0.125,p=0.383;r=0.163,p=0.252;r=0.248,p=0.080);个体结籽率与个体总叶面积、个体高度、个体总花数(代表个体大小)不相关(r=0.014,p=0.925;r=0.120,p=0.402,r=0.166,p=0.245);每花种子数与个体总叶面积不存在显著相关(r=0.014,p=0.112),每花种子数与株高、个体总花数显著相关。(r=0.310,p=0.027;r=0.358,p=0.010)。
4首先计算了每个个体平均胚珠数和平均花粉数,相关性分析发现两者之间无显著性相关(r=0.990,P＜0.01),然后分别计算了两者与个体总叶面积、个体总花数、茎高的残差,再分别计算两者残差的Pearson相关,相应的分析结果为:r=0.717,P＜0.01;r=0.680,P＜0.01;r=0.752,P＜0.01。即紫萼的胚珠数与花粉数无负相关,雌、雄性投入之间的权衡并未得到证实。
5紫萼个体水平上种子大小与数量不相关(无权衡R=0.091,P=0.640)。在29个个体内,22个个体每果种子大小与数量不存在显著关系,在6个体内有显著或极显著的负相关关系(即权衡),在1个个体内极显著负相关。在种群内单个果实水平上,每果种子大小与数量间存在较弱的负相关(即权衡R=0.241,P=0.007)。
6紫萼种子大小变异系数为19.1%,变幅高达7.3倍;种子大小变异主要来源于个体内58.4%(果实间41.2%,果实内17.2%),其次是个体间41.6%。
Sex allocation is one of the most important and success theories in the area of evolution-ecology, it has been used to explain the evolution of breeding systems, sex ratio in population and the formation of selfing. Presently, most studies have focused on the four levels of species, population, individual, inflorescences. The degree of selfing can affect sex allocation of species and populations. Within population, if fitness gain through male and female functions is differently influenced by size or status, it is expected that organisms will adjust sex allocation according to size; Furthermore, sex allocation often varies temporally and spatially among flowers within inflorescences. Ovule or pistil production per flower generally declines from basal(or early-opening)to distal(or late-opening) flowers, whereas pollen or stamen production per flower often remains constant or declines at a lesser degree.
Sex allocation theory predicts that there is trade-off between resource allocation to male vs female function or to size and number at multiple levels. For example, ovule production vs pollen production, seed size vs number. However, the size is different among seeds.
We examined the sex allocation among plants and flowers within inflorescences in a perennial hermaphrodite species with 1-day flowers, hosta ventricosa (Liliaceae). Flowering progresses sequentially from basal to distal positions within an inflorescence with little flowering overlap among flowers. The results are as follows:
(1) There is not a threshold of reproductive in hosta ventricosa.
(2) At plant level, the sex allocation is not correlated with plant size significantly, maybe due to the absence of geitonogamy.
(3) Within inflorescences ovules per flower remained constant, regardless of floral position. Pollen production per flower, the pollen to ovule ratio of individual flowers and seed to flower ratio declined from basal to distal position.
(4) There is no trade-off between ovule production and pollen production per flower at the individual level.
(5) No relation between seed size and number at individual level was detected, both negative and positive or neutral relations between seed size and number per fruit within individual plant was detected, there is the weak significant negative relation between seed size and number at the individual fruit level.
(6) Variable coefficient of the seed size is 19.1%; seed-size variation is mainly based onintra-individual, inter-individual followed.
1用个体总叶面积表征植株个体大小,分别对开花个体的叶长与不开花个体叶长以及两者的叶宽之间进行方差分析,结果它们的p值分别为:0.103和0.995。说明紫萼不存在一个大小繁殖的阈值。
2紫萼同一花序内,每花胚珠数在5个部位无显著变化(t=0.230,P=0.818),每花花粉数、P/O、结实率均从花序的基部到顶部逐渐下降(t=-6.435,P＜0.01;t=-7.493,P＜0.01;t=-3.967,p＜0.01),每果结籽率则从基部到顶部显著上升(t=2.453,p=0.15)。这可能与紫萼开花早期传粉昆虫较少有关。
3个体水平上,分析结果表明个体结实率不随个体总叶面积、个体高度、个体总花数的变化而变化。(r=0.125,p=0.383;r=0.163,p=0.252;r=0.248,p=0.080);个体结籽率与个体总叶面积、个体高度、个体总花数(代表个体大小)不相关(r=0.014,p=0.925;r=0.120,p=0.402,r=0.166,p=0.245);每花种子数与个体总叶面积不存在显著相关(r=0.014,p=0.112),每花种子数与株高、个体总花数显著相关。(r=0.310,p=0.027;r=0.358,p=0.010)。
4首先计算了每个个体平均胚珠数和平均花粉数,相关性分析发现两者之间无显著性相关(r=0.990,P＜0.01),然后分别计算了两者与个体总叶面积、个体总花数、茎高的残差,再分别计算两者残差的Pearson相关,相应的分析结果为:r=0.717,P＜0.01;r=0.680,P＜0.01;r=0.752,P＜0.01。即紫萼的胚珠数与花粉数无负相关,雌、雄性投入之间的权衡并未得到证实。
5紫萼个体水平上种子大小与数量不相关(无权衡R=0.091,P=0.640)。在29个个体内,22个个体每果种子大小与数量不存在显著关系,在6个体内有显著或极显著的负相关关系(即权衡),在1个个体内极显著负相关。在种群内单个果实水平上,每果种子大小与数量间存在较弱的负相关(即权衡R=0.241,P=0.007)。
6紫萼种子大小变异系数为19.1%,变幅高达7.3倍;种子大小变异主要来源于个体内58.4%(果实间41.2%,果实内17.2%),其次是个体间41.6%。
Sex allocation is one of the most important and success theories in the area of evolution-ecology, it has been used to explain the evolution of breeding systems, sex ratio in population and the formation of selfing. Presently, most studies have focused on the four levels of species, population, individual, inflorescences. The degree of selfing can affect sex allocation of species and populations. Within population, if fitness gain through male and female functions is differently influenced by size or status, it is expected that organisms will adjust sex allocation according to size; Furthermore, sex allocation often varies temporally and spatially among flowers within inflorescences. Ovule or pistil production per flower generally declines from basal(or early-opening)to distal(or late-opening) flowers, whereas pollen or stamen production per flower often remains constant or declines at a lesser degree.
Sex allocation theory predicts that there is trade-off between resource allocation to male vs female function or to size and number at multiple levels. For example, ovule production vs pollen production, seed size vs number. However, the size is different among seeds.
We examined the sex allocation among plants and flowers within inflorescences in a perennial hermaphrodite species with 1-day flowers, hosta ventricosa (Liliaceae). Flowering progresses sequentially from basal to distal positions within an inflorescence with little flowering overlap among flowers. The results are as follows:
(1) There is not a threshold of reproductive in hosta ventricosa.
(2) At plant level, the sex allocation is not correlated with plant size significantly, maybe due to the absence of geitonogamy.
(3) Within inflorescences ovules per flower remained constant, regardless of floral position. Pollen production per flower, the pollen to ovule ratio of individual flowers and seed to flower ratio declined from basal to distal position.
(4) There is no trade-off between ovule production and pollen production per flower at the individual level.
(5) No relation between seed size and number at individual level was detected, both negative and positive or neutral relations between seed size and number per fruit within individual plant was detected, there is the weak significant negative relation between seed size and number at the individual fruit level.
(6) Variable coefficient of the seed size is 19.1%; seed-size variation is mainly based onintra-individual, inter-individual followed.
引文
[1]钟章成.植物种群生态研究进展.西南师范大学学报,2001,26:230-236
[2]Wu,Zheng-yi,Hengduan mountain flora and her significance.Journal of Janpanese Botany,1988,63:1-311
[3]李锡文.中国特有种子植物属在云南的两大多样性中心及特征.云南植物研究16(3):221-227
[4]张荣祖,郑度,杨勤业,刘燕华。横断山区自然地理。北京,科学出版社1997:1-12
[5]李锡文,李捷.横断山脉地区种子植物区系的初步研究.云南植物研究,1993,15(3):217-231
[6]祖元刚,王文杰,杨逢建等,植物生活史型多样性及其动态研究,生态学报,2002,22(11):1811-1818
[7]孙儒泳.生活史对策.生物学通报,1997.32(5):2-4.
[8]钟章成.植物种群生态研究进展.西南师范大学学报,2001,26:230-236
[9]张大勇 植物生活史进化与繁殖生态学[M].北京:科学出版社.2004,P:44-46
[10]Gadgil M,and Solbig O.T.The concept of r-k-selection evidence from wild flowers and some theoretical considerations,Amer.Nat.,1972,106:14-13
[11]Abrahamson W.G,Raproductive strategies in dewberries,Ecology,1976,56:712-726.
[12]Newell S.J.and Tramer E.J.Reproductive strategies in herbaceous plant communities during succession.Ecology,1978.59:228-234.
[13]Klinkhamer P.G.L.Life history tactics of annual organiums:the joint effects of dispersal and delayed germination.Theor.Popul.Biol.,1987.32:127-156.
[14]Loehle C.Tree life history strategies:the role of defenses.Can.J.For.Res.,1988.18:209-222
[15]Pendleton B.K.Life history featires of three sexual morphs of Atriplex canescens (Chenopodiaceae)clonal growth in a common garden.Amer.Journal of Botany,1992.79(4):376-382.
[16]Eriksson,O.Reproduction and clonal growth in Potentilla anserina L.(Rosaceae):the relation between growth from and dry weight allocation.Oecologia.1985.66:378-380
[17]Barton N.H.and Turelli M.Genetieal Research.1987,49:157-173.
[18]Goldman,D.A.& Willson,M.F.Sex allocation in functionally hermaphroditic plants:a review and critique.Botanical Review.1986.52:157-194.
[19]Lalonde R.and Roitberg B.D.Mating system,life-history,and reproduction in Canada this He(Cirsium arvense).Amer.J.Bot.,1994.81:21-28.
[20]张大勇 理论生态学研究[M]北京:高等教育出版社2003.3
[21]Wilbur H.M.Life history evolution in seven milkweeds of the geneus ascleplas,Jour.of Ecol.,1976.64:223-240.
[22]Campbell,D.R.Experimental tests of sex-allocation theory in plants.Trends in Ecology and Evolution.2000.15:227-232.
[23]Brunet,J.Sex allocation in hermaphroditic plants.Trends in Ecology and Evolution.1992.7:79-84.
[24]Klinkhamer,P.G.L.De Jong,T.J.and Meta,H.Sex and size in cosexual plants.Trends in Ecology and Evolution,1997.12:260-265
[25]赵志刚,杜国祯,刘左军 雌雄同花植物性分配。生态学报,2005,25:2726-2728
[26]张大勇,姜新华,雷光春,Hanski I.理论生态学研究。北京:高等教育出版社2000.
[27]Klinkhamer,P.G.L.& de Jong,T.J.Size-dependent allocation to male and female reproduction.In Plant resouce allocation,ed.E.A.Bazzaz & J.Grace.Pp.1997.221-229.Sandiego:Academic Press.
[28]Lloyd D G.Selection of offspring size at independence and other size-versus-number strategies.American Naturalist,1987,129:800-817.
[29]Vonhof M J,Harder L D.Size-number trade-offs and pollen production by papilionaceous legumes [J].American Journal of Botany,1995,82:230-238.
[30]de Jong T,Klinkhamer P.Evolution Ecology of Plant Reproductive strategies[M].Cambridge University Press.New York.2005,1:127-128.
[31]武高林,杜国祯,尚占环.种子大小及其命运对植被更新贡献研究进展[J].应用生态学报.2006,17:1969-1972.
[32]Mendez M.Sources of Vriation in Sed Mss in Arumitalicum[J].International Journal of Plant Science.1997,158:298-305.
[33]Schaal B A.Reproductive capacity and seed size in Lupinus texensis[J].American Journal of Botany.1980,67:703-709.
[34]Winn A A,Werner P A,Regulation of seed yeild within and among populations of Prunella vulgaris [J].Ecology,1987,68:1224-1233
[35]Mehlman D W.Seed size and seed packaging variation in Baptisia lanceolata(Fabaceae)[J].American Journal of Botany,1993,80:735-742.
[36]Tom de Jong and Peter Klinkhamer.Evolution Ecology of Plant Reproductive strategies[M]. Cambridge University Press.New York.2005,1:127-128
[37]Ove Eriksson.Seed size variation and its effect on germination and seedling performance in the clonal herb Convallaria majalis[J].Acta Oecologica.1999,20(1):61-66
[38]王桔红,崔现亮,陈学林,杜国祯.中、旱生植物萌发特性及其与种子大小关系的比较[J].植物生态学报.2007,31(6):1037-1045
[39]张大勇,姜新华.植物交配系统的进化、资源分配对策与遗传多样性.植物生态学报,2001,25(2):130-143
[40]Thompson J N.Variation among individual seed masses in Lomatium grayi(Umbelliferae)under controlled conditions:m agnitude and partitioning of the variance.Ecol.,1984,65(2):626-631
[41]Westoby M.Jurado E,Leishman M.Comparative evolutionary ecology of seed sizes.Trends Ecol.Evol.,1992.7(11):368-372.
[42]Renata D.Wulff.Seed size variation in Desmodium Paniculatum[J].Journal of Ecology.1986,74:87-97
[43]Glenda vaughton and Mike ramsey.Seed mass variation in the Shrub Banksia spinulosa (Proteaceae):resource constraints and pollen source effects[J].Int.J.Plant Sci.1997,158(4):424-431
[44]Hammond D S,Brown V K.Seed size of woody-plants in relation to distrubance,dispersal,soil type in the wet neotropical forests[J].Ecology.1995,76:2544-2561
[45]Leishman M R,Wright I J,Moles A T,Westoby M.The evolutionary ecology o seed size[M].In:Fenner M.Seeds:The Ecology of Regeneration in Plant Communities.New York:CABI Publishing.2000
[46]Michaels H.I.Benner B,Hartgerink A P,Lee T D,Rice S.Seed size variation:Magnitude,distribution,and ecology correlates[J].Evolutionary Ecology.1988,2:157-166
[47]Mendez M.Sources of Vriation in Seed Mass in Arum italicum[J].International Journal of Plant Science.1997,158:298-305.
[48]宗文杰,刘坤,卜海燕等.高寒草甸51种菊科植物种子大小变异及其对种子萌发的影响研究[J].兰州大学学报(自然科学版).2006,42(5):52-55
[49]刘振恒,徐秀丽,卜海燕等.青藏高原尔部常见禾本科植物种子大小变异及其与萌发的关系[J].草业科学.2006,23(11):53-57
[50]Lloyd D GGender allocations in outcrossing cosexual plants[A} Perspectives on plant population ecology.Sinauer,Sunderland,Mass.1984.277-300
[51]Chamov. E.L . The Theory of Sex Allocation. Princeton University Press, Princeton NJ.1982.
[52]Charlesworth D,& B. Charlesworth. Allocation of resources to male and female function inhermaphrodites. Biological Journal of the Linnean Society, 1981, 15:57-74
[53]Charlesworth, D, and B.Charlesworth. The of investment in atractive structures on allocation to male and female function in plants. Evolution, 1987.41: 948-968.
[54]Lovett Doust J, Cavers P B. Biomass allocation in hermaphrodite flowers. Canada Journalof botany, 1982.60:2530-2534
[55]Mckone M J. Sex allocation and outcrossing rate: a test of theoretical predictions using Bromegrasses (Bromus). Evolution, 1987.41(3): 591-598
[56]Parachnowttsch, A. L.. and E. EU. Variation in sex allocation and male-female trade-offs in six populations of Collinsia parviflora (Scrophulariaceae S.L). American Journal of Botany 2004.91: 1200-1207.
[57]Lloyd D G, Bawa K S. Modification of the gender of seed plants in varying conditions [J]. Evolutionary Biology, 1984, 17: 225-388.
[58]Akiko Sakai,Satoki Sakai. size-denpendent ESS sex allocation inwind-pollinated cosexual plants.fecundity vs.stature effects[J].Journal of Theoretical Biology .2003.222:283-295.
[59]Stephen I. Wright and Spencer C. H. Barrett, size-denpendent gender modification in a hermaphroditic perennial herb[J] .The royal society , 1999. 266:225-232.
[60]Burd M&Allen T F H.Sexualallocationstrategyinwind-pollinatedplants.Evolution,1988,42:403- 407.
[61] Sato T. Size-dependent sex allocation in hermaphroditic plants: the effects of resource pool and self-incompatibility. Journal of Theoretical Biology.2004, 227:265-275
[62]Cao G X, Kudo K. Size-dependent sex allocation in a monocarpic perennial herb, Cardiocrinum cordatum (Liliaceae) [J]. Plant Ecology, 2007, DOI.10.1007/s11258-007-9277-x (In press).
[63]Diggle, P. K. Architectural effects and the interpretation of patterns of fruit and seed development. Annual Review of Ecology and Systematics. 1995.26: 531-552
[64]Diggle, P. K. Ontogenetic contingency and floral morphology: the. effect of architecture and resource limitation. International Journal of Plant Science, 1997.158 (suppl.): 99-107
[65]Brunet J. Male reproductive success and variation in fruit and seed set in Aquilegia caerulea (R(?)mmculaceae)[J].Ecology, 1996,77:2458-2471
[66]Kudo G. and Molau U. Vareations in reproductive traits at inflorescence and flower levels of an arctic legume, Astragalus alponus L: Comparisous between a subalpone and an alpone population[J]. Plant Species Biology,1999,14:181-19
[67]SusIo D. J. and Lovett-Doust L. Effects of resource availability, and fruit and ovule position on components of fecundity in Alliarea petiolata (Brassicaceae) [J]. New phytol., 1999,144:295-306
[68]E.valltus Position-dependent reproductive success of flowers in Dactylorhiza maculate (Orchidaceae) [J]. Functional Ecology,2000,14:573-579
[69]Medrano M. Guitian P. and Guitian J. Patterns of fruit and seed set within inflorescences of pancratium maritimum(Amaryllidaceae): nonuniform pollination resource limitation or architectural effects [J]. American Journal of Botany,2000,87:493-501
[70]Buide M.L. Variation in Floral Traits and Reproductive Success of the Hermaphrodite Silene acutifolia [L].Annals of Botany,2004,94:441-448
[71] Guitian J. Medrano M. and OTI J. E. Variation in Floral Sex Allocation in Polygonatum odoratum (Liliaceae) [J]. Annals of Botany,2004, 94:433-440
[72]Cao Guo-xing,Yan Juan, Li Yan,et.al Intra-inflorescenceVariation in female fitness components and seed size-number trade-off in Hosta Plantaginea (Liliaceae)[J].Si chuang Agriculture university,2006
[73]Sutherland, S. Why hermaphroditic plants produce many more flowers than fruits: experimental tests with Agave mckelveyana [J].Evolution, 1987,41:750-759
[74]Goldingay, R. L., and R. J. WHELAN. The influence of pollinators on fruit positioning in the Australian shrub Telopea speciosissima (proteaceae)[J].Oikos, 1993,68:501-509
[75]Zimmerman J K. Aide T M. Pattrtns of fruit production in a neotropical orchid:pollinator vs.resource limitation [J]. American Journal of BOTANY,1989,76:67-73
[76]Stephenson, A. G. Flower and fruit abortion:proximate causes and ultimate functions[J] Annual review of Ecology and Systematics, 1981,12:253-279
[77] Lee,T.D. Patterns of fruit and seed production[J].Plant reproductive ecology:patterns and atrategies, 1988,179-202
[78]Ishii, H. S., and S. Sakai. Temporal variation in floral display size and individual floral sex allocation in racemes of Narthecium asiaticum (Liliaceae). American Journal of Botany, 2002. 89:441-446
[79]Vogler, D. W.. S. Peretz, and A. G. Stephenson. Floral plasticity in an iteroparous plant: the interactive effects of genotype,environment,and ontogeny in Campanula rapunculoides (Campanulaceae).American Journal of Botany,1999.86:482-494
[80]Garcia,M,B.Sex allocation in a long-lived monocarpic plant.Plant Biology.2003.5:203-209
[81]Stephenson A G.An evolutionary examination of the floral display of Catalpa speciosa (Bignoniaceae)[J].Evolution,1979,33:1200-1209
[82]Kliber A.and Eckert C.G.Sequential decline in allocation among flowers within inflorescences:proximate mechanisms and adaptive significance[J].Ecology,2004,85:1675-1687
[83]M.B.Garcia Sex Allocation in a Long-Lived Monocarpic Plant[J].Plant biology,2003,5:203-209
[84]Giggle.K.and Siller J.S.Architectural effects mimic floral sexual dimorphism in solanom (solanaceae)[J].American Journal of Botany,2004,91:2030-3040
[85]Ashman T.and Hitchens M.S.Dissecting the causes of variation in intra-inflorescence allocation in a sexually polymorphic apecies,fragaria virginiana(rosaceae)[J].American Journal of Botany,2000,87:197-204
[86]Cao Guo-xing.Intra-inflorescence Variation in female fitness components and seed size-number trade-off in Hosta Plantaginea(Liliaceae)[J].Si chuang Agriculture university,2006
[87]Brunet,J.and D.Charlesworth.Floral sex allocation in sequentially blooming plants[J].Evolution,1995,49:70-79
[88]Lee,T.D.Patterns of fruit and seed production[J].Plant reproductive ecology:patterns and atrategies,1988,179-202
[89]Berry,P.E.,and R.N.Calvo.Pollinator limitation and position dependent fruit set in the high Andean orchid Myrosmodes cochleare(Orchidaceae)[J].Plant Systematica and Evolution,1991,174:93-101
[90]唐进.中国植物志第14卷.科学出版社.1980.P49-51
[91]贾林平,将远胜,戎晓红.碧峰峡景区生态系统服务及其价值评价.国土资源科技管理.2006.P168-112
[92]Solomon,B.P.Sexual allocation and andromonoecy:resource investment in male and hermaphrodite flowers of Solanum carolinence(Solanaceae).American Journal Botany,1986.73:1215-1221.
[93]Stearns,S.C The evolution of life histories.Oxford University Press,New York.1992.
[94]Cao G X,Kudo K,Ida T Y.Floral sex allocation of a hermaphrodite herb with one-day flowers, Hosta rectifolia(Liliaceae)[J].Plant Species Biology,2007,22:191-196.
[95]Wright S I,Barrett S C H.Size-dependent gender modification in a hermaphroditic perennial herb [J].Proceedings of the Royal Society of London B,1999,266:225-232.
[96]Koelewijn H P,Hunscheid M P H.Intraspecific variation in sex allocation in hermaphroditic Plantago coronopus(L.)[J].Journal of Evolutionary Biology.2000,13:302-315.
[97]Mazer S J,Dawson K A.Size-dependent sex allocation within flowers of the annual herb Clarkia unguiculata(Onagraceae):ontogenetic and among-plant variation[J].American Journal of Botany,2001,88,819-831.
[98]Mazer S J,Delesalle V A,Neal P R.Responses of floral traits to selection on primary sexual investment in Spergularia marina:the battle between the sexes[J].Evolution,1999,53:717-731.
[99]Schaal B A.Reproductive capacity and seed size in Lupinus texensis[J].American Journal of Botany.1980,67:703-709.
[100]Winn A A,Werner P A.Regulation oi seed yeild within and among populations of Prunella vulgaris[J].Ecology,1987,68:1224-1233
[101]H J Michaels,B Benner,A P Hangerink,T D Lee,S Rice,M F Willson and R I Benin.Seed size variation:magnitude,distribution,and ecological correlates.Evolutionary Ecology,1988.2:157-166.
[102]Mehlman D W.Seed size and seed packaging variation in Baptisia lanceolata(Fabaceae)[J].American Journal of Botany,1993,80:735-742.
[103]Simons AM,Johnston MO Variation in seed traits of Lobelia inflata(Campanulaceae):sources and fitness consequences.American Journal of Botany,2000.87,124-132.
[104]Vaughton G,Ramsey M.Seed mass variation in the shrub Banksia spinulosa(Proteaceae):resource constraints and pollination effects.International Journal of Plant Science,1997.158,424-431.
[105]Lloyd D G.Sexual Strategies in Plants.Ⅰ.An Hypothesis of Serial Adjustment of Maternal Investment During One Reproductive Session,New Phytologist.1980,86:69-79
[106]Stacey L.Halpern.Sources and consequences of seed size variation in Lupinus perennis (fabaceae):adaptive and non-adaptive hypotheses[J].American Journal of Botany.2005,92(2):205-213
[107]Obeso J R.Seed mass variation in the perennial herb asphodelus aldus:Sources of varialion and position effect[J].Oecologia.1993,93:571-575
[108]张世挺,杜国祯卜,陈家宽.种子大小变异的进化生态学研究现状与展望[J].生态学报.2003,23(2):353-364
[109]D.Lawrence Venable. Size-number trade-offs and the variation o seed size with plant resource status[J].The American Naturalist. 1992,140(2):287-304
[2]Wu,Zheng-yi,Hengduan mountain flora and her significance.Journal of Janpanese Botany,1988,63:1-311
[3]李锡文.中国特有种子植物属在云南的两大多样性中心及特征.云南植物研究16(3):221-227
[4]张荣祖,郑度,杨勤业,刘燕华。横断山区自然地理。北京,科学出版社1997:1-12
[5]李锡文,李捷.横断山脉地区种子植物区系的初步研究.云南植物研究,1993,15(3):217-231
[6]祖元刚,王文杰,杨逢建等,植物生活史型多样性及其动态研究,生态学报,2002,22(11):1811-1818
[7]孙儒泳.生活史对策.生物学通报,1997.32(5):2-4.
[8]钟章成.植物种群生态研究进展.西南师范大学学报,2001,26:230-236
[9]张大勇 植物生活史进化与繁殖生态学[M].北京:科学出版社.2004,P:44-46
[10]Gadgil M,and Solbig O.T.The concept of r-k-selection evidence from wild flowers and some theoretical considerations,Amer.Nat.,1972,106:14-13
[11]Abrahamson W.G,Raproductive strategies in dewberries,Ecology,1976,56:712-726.
[12]Newell S.J.and Tramer E.J.Reproductive strategies in herbaceous plant communities during succession.Ecology,1978.59:228-234.
[13]Klinkhamer P.G.L.Life history tactics of annual organiums:the joint effects of dispersal and delayed germination.Theor.Popul.Biol.,1987.32:127-156.
[14]Loehle C.Tree life history strategies:the role of defenses.Can.J.For.Res.,1988.18:209-222
[15]Pendleton B.K.Life history featires of three sexual morphs of Atriplex canescens (Chenopodiaceae)clonal growth in a common garden.Amer.Journal of Botany,1992.79(4):376-382.
[16]Eriksson,O.Reproduction and clonal growth in Potentilla anserina L.(Rosaceae):the relation between growth from and dry weight allocation.Oecologia.1985.66:378-380
[17]Barton N.H.and Turelli M.Genetieal Research.1987,49:157-173.
[18]Goldman,D.A.& Willson,M.F.Sex allocation in functionally hermaphroditic plants:a review and critique.Botanical Review.1986.52:157-194.
[19]Lalonde R.and Roitberg B.D.Mating system,life-history,and reproduction in Canada this He(Cirsium arvense).Amer.J.Bot.,1994.81:21-28.
[20]张大勇 理论生态学研究[M]北京:高等教育出版社2003.3
[21]Wilbur H.M.Life history evolution in seven milkweeds of the geneus ascleplas,Jour.of Ecol.,1976.64:223-240.
[22]Campbell,D.R.Experimental tests of sex-allocation theory in plants.Trends in Ecology and Evolution.2000.15:227-232.
[23]Brunet,J.Sex allocation in hermaphroditic plants.Trends in Ecology and Evolution.1992.7:79-84.
[24]Klinkhamer,P.G.L.De Jong,T.J.and Meta,H.Sex and size in cosexual plants.Trends in Ecology and Evolution,1997.12:260-265
[25]赵志刚,杜国祯,刘左军 雌雄同花植物性分配。生态学报,2005,25:2726-2728
[26]张大勇,姜新华,雷光春,Hanski I.理论生态学研究。北京:高等教育出版社2000.
[27]Klinkhamer,P.G.L.& de Jong,T.J.Size-dependent allocation to male and female reproduction.In Plant resouce allocation,ed.E.A.Bazzaz & J.Grace.Pp.1997.221-229.Sandiego:Academic Press.
[28]Lloyd D G.Selection of offspring size at independence and other size-versus-number strategies.American Naturalist,1987,129:800-817.
[29]Vonhof M J,Harder L D.Size-number trade-offs and pollen production by papilionaceous legumes [J].American Journal of Botany,1995,82:230-238.
[30]de Jong T,Klinkhamer P.Evolution Ecology of Plant Reproductive strategies[M].Cambridge University Press.New York.2005,1:127-128.
[31]武高林,杜国祯,尚占环.种子大小及其命运对植被更新贡献研究进展[J].应用生态学报.2006,17:1969-1972.
[32]Mendez M.Sources of Vriation in Sed Mss in Arumitalicum[J].International Journal of Plant Science.1997,158:298-305.
[33]Schaal B A.Reproductive capacity and seed size in Lupinus texensis[J].American Journal of Botany.1980,67:703-709.
[34]Winn A A,Werner P A,Regulation of seed yeild within and among populations of Prunella vulgaris [J].Ecology,1987,68:1224-1233
[35]Mehlman D W.Seed size and seed packaging variation in Baptisia lanceolata(Fabaceae)[J].American Journal of Botany,1993,80:735-742.
[36]Tom de Jong and Peter Klinkhamer.Evolution Ecology of Plant Reproductive strategies[M]. Cambridge University Press.New York.2005,1:127-128
[37]Ove Eriksson.Seed size variation and its effect on germination and seedling performance in the clonal herb Convallaria majalis[J].Acta Oecologica.1999,20(1):61-66
[38]王桔红,崔现亮,陈学林,杜国祯.中、旱生植物萌发特性及其与种子大小关系的比较[J].植物生态学报.2007,31(6):1037-1045
[39]张大勇,姜新华.植物交配系统的进化、资源分配对策与遗传多样性.植物生态学报,2001,25(2):130-143
[40]Thompson J N.Variation among individual seed masses in Lomatium grayi(Umbelliferae)under controlled conditions:m agnitude and partitioning of the variance.Ecol.,1984,65(2):626-631
[41]Westoby M.Jurado E,Leishman M.Comparative evolutionary ecology of seed sizes.Trends Ecol.Evol.,1992.7(11):368-372.
[42]Renata D.Wulff.Seed size variation in Desmodium Paniculatum[J].Journal of Ecology.1986,74:87-97
[43]Glenda vaughton and Mike ramsey.Seed mass variation in the Shrub Banksia spinulosa (Proteaceae):resource constraints and pollen source effects[J].Int.J.Plant Sci.1997,158(4):424-431
[44]Hammond D S,Brown V K.Seed size of woody-plants in relation to distrubance,dispersal,soil type in the wet neotropical forests[J].Ecology.1995,76:2544-2561
[45]Leishman M R,Wright I J,Moles A T,Westoby M.The evolutionary ecology o seed size[M].In:Fenner M.Seeds:The Ecology of Regeneration in Plant Communities.New York:CABI Publishing.2000
[46]Michaels H.I.Benner B,Hartgerink A P,Lee T D,Rice S.Seed size variation:Magnitude,distribution,and ecology correlates[J].Evolutionary Ecology.1988,2:157-166
[47]Mendez M.Sources of Vriation in Seed Mass in Arum italicum[J].International Journal of Plant Science.1997,158:298-305.
[48]宗文杰,刘坤,卜海燕等.高寒草甸51种菊科植物种子大小变异及其对种子萌发的影响研究[J].兰州大学学报(自然科学版).2006,42(5):52-55
[49]刘振恒,徐秀丽,卜海燕等.青藏高原尔部常见禾本科植物种子大小变异及其与萌发的关系[J].草业科学.2006,23(11):53-57
[50]Lloyd D GGender allocations in outcrossing cosexual plants[A} Perspectives on plant population ecology.Sinauer,Sunderland,Mass.1984.277-300
[51]Chamov. E.L . The Theory of Sex Allocation. Princeton University Press, Princeton NJ.1982.
[52]Charlesworth D,& B. Charlesworth. Allocation of resources to male and female function inhermaphrodites. Biological Journal of the Linnean Society, 1981, 15:57-74
[53]Charlesworth, D, and B.Charlesworth. The of investment in atractive structures on allocation to male and female function in plants. Evolution, 1987.41: 948-968.
[54]Lovett Doust J, Cavers P B. Biomass allocation in hermaphrodite flowers. Canada Journalof botany, 1982.60:2530-2534
[55]Mckone M J. Sex allocation and outcrossing rate: a test of theoretical predictions using Bromegrasses (Bromus). Evolution, 1987.41(3): 591-598
[56]Parachnowttsch, A. L.. and E. EU. Variation in sex allocation and male-female trade-offs in six populations of Collinsia parviflora (Scrophulariaceae S.L). American Journal of Botany 2004.91: 1200-1207.
[57]Lloyd D G, Bawa K S. Modification of the gender of seed plants in varying conditions [J]. Evolutionary Biology, 1984, 17: 225-388.
[58]Akiko Sakai,Satoki Sakai. size-denpendent ESS sex allocation inwind-pollinated cosexual plants.fecundity vs.stature effects[J].Journal of Theoretical Biology .2003.222:283-295.
[59]Stephen I. Wright and Spencer C. H. Barrett, size-denpendent gender modification in a hermaphroditic perennial herb[J] .The royal society , 1999. 266:225-232.
[60]Burd M&Allen T F H.Sexualallocationstrategyinwind-pollinatedplants.Evolution,1988,42:403- 407.
[61] Sato T. Size-dependent sex allocation in hermaphroditic plants: the effects of resource pool and self-incompatibility. Journal of Theoretical Biology.2004, 227:265-275
[62]Cao G X, Kudo K. Size-dependent sex allocation in a monocarpic perennial herb, Cardiocrinum cordatum (Liliaceae) [J]. Plant Ecology, 2007, DOI.10.1007/s11258-007-9277-x (In press).
[63]Diggle, P. K. Architectural effects and the interpretation of patterns of fruit and seed development. Annual Review of Ecology and Systematics. 1995.26: 531-552
[64]Diggle, P. K. Ontogenetic contingency and floral morphology: the. effect of architecture and resource limitation. International Journal of Plant Science, 1997.158 (suppl.): 99-107
[65]Brunet J. Male reproductive success and variation in fruit and seed set in Aquilegia caerulea (R(?)mmculaceae)[J].Ecology, 1996,77:2458-2471
[66]Kudo G. and Molau U. Vareations in reproductive traits at inflorescence and flower levels of an arctic legume, Astragalus alponus L: Comparisous between a subalpone and an alpone population[J]. Plant Species Biology,1999,14:181-19
[67]SusIo D. J. and Lovett-Doust L. Effects of resource availability, and fruit and ovule position on components of fecundity in Alliarea petiolata (Brassicaceae) [J]. New phytol., 1999,144:295-306
[68]E.valltus Position-dependent reproductive success of flowers in Dactylorhiza maculate (Orchidaceae) [J]. Functional Ecology,2000,14:573-579
[69]Medrano M. Guitian P. and Guitian J. Patterns of fruit and seed set within inflorescences of pancratium maritimum(Amaryllidaceae): nonuniform pollination resource limitation or architectural effects [J]. American Journal of Botany,2000,87:493-501
[70]Buide M.L. Variation in Floral Traits and Reproductive Success of the Hermaphrodite Silene acutifolia [L].Annals of Botany,2004,94:441-448
[71] Guitian J. Medrano M. and OTI J. E. Variation in Floral Sex Allocation in Polygonatum odoratum (Liliaceae) [J]. Annals of Botany,2004, 94:433-440
[72]Cao Guo-xing,Yan Juan, Li Yan,et.al Intra-inflorescenceVariation in female fitness components and seed size-number trade-off in Hosta Plantaginea (Liliaceae)[J].Si chuang Agriculture university,2006
[73]Sutherland, S. Why hermaphroditic plants produce many more flowers than fruits: experimental tests with Agave mckelveyana [J].Evolution, 1987,41:750-759
[74]Goldingay, R. L., and R. J. WHELAN. The influence of pollinators on fruit positioning in the Australian shrub Telopea speciosissima (proteaceae)[J].Oikos, 1993,68:501-509
[75]Zimmerman J K. Aide T M. Pattrtns of fruit production in a neotropical orchid:pollinator vs.resource limitation [J]. American Journal of BOTANY,1989,76:67-73
[76]Stephenson, A. G. Flower and fruit abortion:proximate causes and ultimate functions[J] Annual review of Ecology and Systematics, 1981,12:253-279
[77] Lee,T.D. Patterns of fruit and seed production[J].Plant reproductive ecology:patterns and atrategies, 1988,179-202
[78]Ishii, H. S., and S. Sakai. Temporal variation in floral display size and individual floral sex allocation in racemes of Narthecium asiaticum (Liliaceae). American Journal of Botany, 2002. 89:441-446
[79]Vogler, D. W.. S. Peretz, and A. G. Stephenson. Floral plasticity in an iteroparous plant: the interactive effects of genotype,environment,and ontogeny in Campanula rapunculoides (Campanulaceae).American Journal of Botany,1999.86:482-494
[80]Garcia,M,B.Sex allocation in a long-lived monocarpic plant.Plant Biology.2003.5:203-209
[81]Stephenson A G.An evolutionary examination of the floral display of Catalpa speciosa (Bignoniaceae)[J].Evolution,1979,33:1200-1209
[82]Kliber A.and Eckert C.G.Sequential decline in allocation among flowers within inflorescences:proximate mechanisms and adaptive significance[J].Ecology,2004,85:1675-1687
[83]M.B.Garcia Sex Allocation in a Long-Lived Monocarpic Plant[J].Plant biology,2003,5:203-209
[84]Giggle.K.and Siller J.S.Architectural effects mimic floral sexual dimorphism in solanom (solanaceae)[J].American Journal of Botany,2004,91:2030-3040
[85]Ashman T.and Hitchens M.S.Dissecting the causes of variation in intra-inflorescence allocation in a sexually polymorphic apecies,fragaria virginiana(rosaceae)[J].American Journal of Botany,2000,87:197-204
[86]Cao Guo-xing.Intra-inflorescence Variation in female fitness components and seed size-number trade-off in Hosta Plantaginea(Liliaceae)[J].Si chuang Agriculture university,2006
[87]Brunet,J.and D.Charlesworth.Floral sex allocation in sequentially blooming plants[J].Evolution,1995,49:70-79
[88]Lee,T.D.Patterns of fruit and seed production[J].Plant reproductive ecology:patterns and atrategies,1988,179-202
[89]Berry,P.E.,and R.N.Calvo.Pollinator limitation and position dependent fruit set in the high Andean orchid Myrosmodes cochleare(Orchidaceae)[J].Plant Systematica and Evolution,1991,174:93-101
[90]唐进.中国植物志第14卷.科学出版社.1980.P49-51
[91]贾林平,将远胜,戎晓红.碧峰峡景区生态系统服务及其价值评价.国土资源科技管理.2006.P168-112
[92]Solomon,B.P.Sexual allocation and andromonoecy:resource investment in male and hermaphrodite flowers of Solanum carolinence(Solanaceae).American Journal Botany,1986.73:1215-1221.
[93]Stearns,S.C The evolution of life histories.Oxford University Press,New York.1992.
[94]Cao G X,Kudo K,Ida T Y.Floral sex allocation of a hermaphrodite herb with one-day flowers, Hosta rectifolia(Liliaceae)[J].Plant Species Biology,2007,22:191-196.
[95]Wright S I,Barrett S C H.Size-dependent gender modification in a hermaphroditic perennial herb [J].Proceedings of the Royal Society of London B,1999,266:225-232.
[96]Koelewijn H P,Hunscheid M P H.Intraspecific variation in sex allocation in hermaphroditic Plantago coronopus(L.)[J].Journal of Evolutionary Biology.2000,13:302-315.
[97]Mazer S J,Dawson K A.Size-dependent sex allocation within flowers of the annual herb Clarkia unguiculata(Onagraceae):ontogenetic and among-plant variation[J].American Journal of Botany,2001,88,819-831.
[98]Mazer S J,Delesalle V A,Neal P R.Responses of floral traits to selection on primary sexual investment in Spergularia marina:the battle between the sexes[J].Evolution,1999,53:717-731.
[99]Schaal B A.Reproductive capacity and seed size in Lupinus texensis[J].American Journal of Botany.1980,67:703-709.
[100]Winn A A,Werner P A.Regulation oi seed yeild within and among populations of Prunella vulgaris[J].Ecology,1987,68:1224-1233
[101]H J Michaels,B Benner,A P Hangerink,T D Lee,S Rice,M F Willson and R I Benin.Seed size variation:magnitude,distribution,and ecological correlates.Evolutionary Ecology,1988.2:157-166.
[102]Mehlman D W.Seed size and seed packaging variation in Baptisia lanceolata(Fabaceae)[J].American Journal of Botany,1993,80:735-742.
[103]Simons AM,Johnston MO Variation in seed traits of Lobelia inflata(Campanulaceae):sources and fitness consequences.American Journal of Botany,2000.87,124-132.
[104]Vaughton G,Ramsey M.Seed mass variation in the shrub Banksia spinulosa(Proteaceae):resource constraints and pollination effects.International Journal of Plant Science,1997.158,424-431.
[105]Lloyd D G.Sexual Strategies in Plants.Ⅰ.An Hypothesis of Serial Adjustment of Maternal Investment During One Reproductive Session,New Phytologist.1980,86:69-79
[106]Stacey L.Halpern.Sources and consequences of seed size variation in Lupinus perennis (fabaceae):adaptive and non-adaptive hypotheses[J].American Journal of Botany.2005,92(2):205-213
[107]Obeso J R.Seed mass variation in the perennial herb asphodelus aldus:Sources of varialion and position effect[J].Oecologia.1993,93:571-575
[108]张世挺,杜国祯卜,陈家宽.种子大小变异的进化生态学研究现状与展望[J].生态学报.2003,23(2):353-364
[109]D.Lawrence Venable. Size-number trade-offs and the variation o seed size with plant resource status[J].The American Naturalist. 1992,140(2):287-304