川西风毛菊(Saussurea dzeurensis)花展示的变异:检验花及花序水平大小-数量的权衡

植物花展示对传粉昆虫吸引和植物繁殖成功至关重要。理论研究指出,花展示的组成部分(如花的大小和数量)之间由于资源的限制而存在权衡关系,从而制约花展示的进化,但相关实验研究并不经常能检测到花的大小-数量之间的权衡关系。本文以广泛分布于青藏高原的川西风毛菊(Saussurea dzeurensis)为对象,研究不同海拔种群间花展示(花序水平和花水平)的变异,检验花大小-数量的权衡关系。结果表明:每株的头状花序数、单个头状花序重、单花的数量/头状花序与地上生物量呈显著的正相关,说明花展示大小(包括花序水平和花水平)依赖于个体的资源状况;高海拔种群个体更小,平均花序数更少,但单个头状花序的重量、单花的数量及重量/头状花序与低海拔种群相比没有显著差异;在花序水平上头状花序的大小和数量间并没有权衡关系,但在花序内的花水平上,高海拔种群花的大小和数量呈显著负相关,表现出权衡关系,而在低海拔种群并没有权衡关系;当控制川西风毛菊个体大小的影响后,花序水平大小-数量及花大小-数量间的相关均没有发生改变,说明资源获取能力的差异,即个体大小的差别并没有影响川西风毛菊花序水平或花水平大小-数量间的权衡关系;植物花大小-数量的权衡关系可能较为复杂,今后的研究需要同时考虑多个组织水平,如种群、个体、花序水平的花大小与数量的关系,有助于全面揭示植物花的资源分配的模式及花展示大小的进化机制。     

新疆郁金香营养生长、个体大小和开花次序对繁殖分配的影响

植物有性繁殖与资源分配的关系研究对于揭示植物生活史特征及繁育系统进化具有重要意义。新疆郁金香(Tulipa sinkiangensis)是新疆天山北坡荒漠带特有的一种多年生早春短命植物。在自然生境中,该物种仅以有性繁殖产生后代,每株能产生1-8朵花,且不同植株上的花数及果实数以及花序不同位置上的花与果实大小明显不同。本文通过对新疆郁金香有性繁殖与营养生长及植株大小的关系以及花序中不同位置花及果实间的资源分配研究,旨在揭示营养生长、个体大小及开花次序对其繁殖分配的影响。结果表明:在开花和果实成熟阶段,新疆郁金香植株分配给营养器官(鳞茎和地上营养器官)与繁殖器官的资源间均存在极显著的负相关关系(P<0.01),说明其植株的营养生长与生殖生长间存在权衡关系。多花是新疆郁金香的一个稳定性状,其植株上花数目、花生物量、果实生物量和种子数量与植株生物量间均呈极显著的正相关关系(P<0.01),说明新疆郁金香植株的繁殖分配存在大小依赖性。在具2-5朵花的新疆郁金香植株中,花序内各花的生物量、花粉数和胚珠数、结实率、果实生物量、结籽数、结籽率及种子百粒重按其开花顺序依次递减,说明花序内各花和果实的资源分配符合资源竞争假说。植株通过减少晚发育的花或果实获得的资源来保障早发育的花或果实获得较多的资源,从而达到繁殖成功。     

高寒退化草地狼毒种群不同海拔花大小-数量的权衡关系

权衡关系是植物生活史对策理论的基础, 花大小-数量的权衡关系对理解花的生物量分配具有重要的意义。该文利用实验生态学方法, 研究了祁连山北坡高寒退化草地4个不同海拔狼毒(Stellera chamaejasme)种群花大小与花数量间的关系。结果表明: 随着海拔升高, 草地群落的高度、密度和地上生物量均呈先升高后降低的倒U型分布, 狼毒花大小呈逐渐增大的趋势, 而花数量呈相反的变化趋势; 不同海拔样地狼毒花大小与花数量间的相关性存在差异, 海拔2700、2900和3000 m样地狼毒花大小和花数量之间均存在极显著的负相关关系(p < 0.01), 海拔2800 m样地二者之间存在显著的负相关关系(p < 0.05), 狼毒植株分配给花大小与花数量的资源间存在着“此消彼长”的权衡关系; 海拔2800 m样地是狼毒植株花大小与花数量权衡关系由低海拔的花数量多而单花生物量低向高海拔的花数量少而单花生物量高转变的区域。狼毒植株通过合理权衡花数量和花大小的资源配置, 以补偿在海拔梯度上不利因素对种群繁衍带来的影响, 从而实现种群的繁殖成功, 反映了毒杂草对异质性生境的表型可塑性机制。     

Trade-off between the biomass and number of flowers in Stellera chamaejasme along an elevation gradient in a degraded alpine grassland

权衡关系是植物生活史对策理论的基础, 花大小-数量的权衡关系对理解花的生物量分配具有重要的意义。该文利用实验生态学方法, 研究了祁连山北坡高寒退化草地4个不同海拔狼毒(Stellera chamaejasme)种群花大小与花数量间的关系。结果表明: 随着海拔升高, 草地群落的高度、密度和地上生物量均呈先升高后降低的倒U型分布, 狼毒花大小呈逐渐增大的趋势, 而花数量呈相反的变化趋势; 不同海拔样地狼毒花大小与花数量间的相关性存在差异, 海拔2700、2900和3000 m样地狼毒花大小和花数量之间均存在极显著的负相关关系(p < 0.01), 海拔2800 m样地二者之间存在显著的负相关关系(p < 0.05), 狼毒植株分配给花大小与花数量的资源间存在着“此消彼长”的权衡关系; 海拔2800 m样地是狼毒植株花大小与花数量权衡关系由低海拔的花数量多而单花生物量低向高海拔的花数量少而单花生物量高转变的区域。狼毒植株通过合理权衡花数量和花大小的资源配置, 以补偿在海拔梯度上不利因素对种群繁衍带来的影响, 从而实现种群的繁殖成功, 反映了毒杂草对异质性生境的表型可塑性机制。     

高寒退化草地不同海拔狼毒种群花大小与叶大小、叶数量的关系

植物花大小与叶大小、叶数量的关系反映了植物长期与环境相互作用形成的外在形态方面的适应对策,不同生境中其生长关系的变异体现了植物对异质环境的适应方式。本文研究了祁连山北坡高寒退化草地4个不同海拔狼毒(Stellera chamaejasme)种群花大小与叶大小、叶数量的关系。结果表明:随着海拔的升高,草地群落的高度、密度和地上生物量均呈先升高后降低的倒U型分布;狼毒的地上生物量、株高和叶大小均呈减小趋势,而繁殖分配、花大小和叶数量均呈增大的变化趋势;在海拔梯度上,狼毒花大小与叶数量呈极显著正相关(P0.01),与叶大小呈不显著负相关(P0.05),说明狼毒种群花大小与叶数量具有显著的依赖关系;生境对狼毒花大小与叶数量之间的依赖关系产生显著影响,高海拔的环境胁迫导致狼毒个体减小,植株通过增加花大小和叶数量、减小叶大小的资源分配策略来保证物种的繁衍。     

红姜花(姜科)同步大量开花的适应意义

两性花植物同步大量开花增加了个体花展示大小,能吸引更多的传粉者,使植株获得更多的交配机会,但也不可避免地带来不同程度的同株异花授粉,进而对被子植物的性资源分配、花部性状以及雌雄异株的进化等产生重要的影响.本文以姜科红姜花(Hedychium coccineum)为研究对象,通过人工授粉和操控试验、传粉动物的观察、自然结果率和自然居群的大小和密度的调查,探讨红姜花同步大量开花的适应意义.结果表明:(1)红姜花的穗状花序由57.33±1.68(n=30)个蝎尾状聚伞小花序组成,每个小花序具3.8±0.15(n=30)朵花,同花序上所有的小花序同步开花,不同小花序内的花按轮次同步开放,使花序在整个开花期间保持有数十朵花同时开放,花序水平上的花展示极为显著;(2)红姜花为自交亲和植物,但不存在自动自花授粉,自然居群存在严重的传粉者限制,其结实也存在着资源限制;(3)多型蓝凤蝶(Papilio memnon)、小矩翅粉蝶(Dercas lycorias)和黑角尖翅粉蝶云南亚种(Appias indra aristoxemus)是红姜花的有效传粉者,3种蝴蝶对红姜花不同大小花序的访问频率都显著不同,随着花序花展示大小的增加其访问频率也显著增加;(4)红姜花同一个体小花序内不同轮次花之间的胚珠数随轮次显著减少,而花粉数前3轮随轮次显著增加.胚珠数的递减及结实存在着资源限制,说明红姜花一个花序开大量的花并不是为了增加结实,而花粉数前3轮的递增,有利于花粉的分散输出,提高个体的雄性适合度.     

雌全同株植物三脉紫菀花期偏雄的个体大小依赖的性别分配

经典的虫媒传粉植物个体大小依赖的性别分配模型通常预期:分配给雌性功能的资源比例将随着个体大小的增大而增加;但一些研究表明,花期个体大小依赖的性别分配模式表现出随个体大小增大而偏雄的趋势.我们以植株高度衡量个体大小,从花和花序两个水平上研究了雌花、两性花同株植物三脉紫菀(Aster ageratoides)花期个体大小依赖的性别分配策略.随着植株高度的增大,植株产生的头状花序数量增加,表明三脉紫菀投入到繁殖的资源不是固定不变的,而是随个体大小增大而增加的.在花和花序水平上,繁殖资源在雌雄性别功能之间的分配均表现为随个体大小的增大而更偏雄的模式,即花粉/胚珠比增加,产生花粉的两性花占两性花和雌花总花数的比例升高.这些结果与花期个体越大、性别分配越偏雄的预期一致.花期更偏雄的性别分配可能有助于植物在花期通过输出花粉提高雄性适合度,从而实现个体适合度的最大化.     

风毛菊花序、种子大小和数量之间的权衡:资源条件的影响

繁殖器官的大小和数量之间的权衡是植物繁殖对策研究的重要问题之一, 但是支持权衡关系的实验研究非常有限。该文以青藏高原东缘高寒草甸菊科风毛菊属优势植物种风毛菊(Saussurea japonica)为材料, 通过不同光照和营养处理, 研究其花序、种子大小和数量之间的关系。结果表明: 光照和营养处理对花序数目没有显著影响, 但对花的大小, 以及种子大小和数量均有显著影响。大小和数量之间的关系同样会受到处理条件的影响: 在对照和低营养处理下, 花序、种子的大小和数量之间呈显著的正相关关系, 而在低光照处理和低光照低营养处理下, 花序、种子的大小和数量之间的关系变为显著负相关, 即出现大小和数量之间的权衡。结果表明: 环境胁迫(比如有限的土壤养分和光资源的抑制)加剧了花序、种子大小和数量之间的权衡。风毛菊花序和种子的大小和数量之间的权衡关系的环境依赖性, 使其能够对外界环境的变化做出响应, 这种繁殖对策能提高其适合度。     

呼伦贝尔草原植物叶片大小与数量的权衡关系及其内在机制

叶片大小与数量的权衡关系是植物生长策略的基础,研究叶片大小与数量权衡关系的内在机制对于深刻理解植物生长策略具有重要意义。本研究以单个叶片干重表示叶大小,以单位茎上的叶片数量表示出叶强度,采用标准主轴回归分析方法研究呼伦贝尔草原植物叶片大小与数量的权衡关系以及内在机制。结果表明: 陈旗嵯岗(典型草原)和陈旗八一(草甸草原)叶片大小与出叶强度存在显著的负的等速生长关系,谢尔塔拉(草甸草原)的叶片大小与出叶强度存在显著的负的异速生长权衡关系。叶片大小和数量权衡关系的内在机制取决于叶片与茎生物量的分配机制以及茎组织密度的变化。     

利用腊叶标本初探豹子花属植物的性表达及其与个体大小的关系

植物性系统进化过程中很少有植物通过雌性败育的方式实现性系统由联合走向分离,然而雄花在百合科植物中大量存在,引起了研究者的关注。本研究利用百合科豹子花属植物的腊叶标本,获取它们的花性别表型、花大小、株高以及分布海拔等信息探讨豹子花属植物的性表达样式,并对样本量较多的开瓣豹子花、云南豹子花、多斑豹子花和豹子花四个种的性表达及其与个体大小的关系进行分析。研究结果表明:1)六种豹子花属植物标本中都有雄花存在;根据雄性花和两性花的不同组合,形成雄性个体、两性花个体,和雄花两性花同株个体三种植株类型;2)个体大小(株高)和分布海拔之间不存在相关关系,和植株性别类型相关,即雄性个体一般显著小于具有雌性功能的两性花个体和雄花两性花同株个体;3)个体大小与花大小、每个个体开花总数和雌花数量成显著的正相关关系,而与每个个体的雄花数量无显著的相关关系。本研究更正了一直以来对豹子花属植物性系统的错误认识,首次报道雄花在豹子花属植物中普遍存在,并指出雄花和雄性个体的出现是大小依赖性分配的结果。同时,以研究实例证明了利用腊叶标本开展植物性系统多样性的可行性。     

Phylogenetic evidence for a flower size and number trade-off

The size and number of flowers displayed together on an inflorescence (floral display) influences pollinator attraction and pollen transfer and receipt, and is integral to plant reproductive success and fitness. Life history theory predicts that the evolution of floral display is constrained by trade-offs between the size and number of flowers and inflorescences. Indeed, a trade-off between flower size and flower number is a key assumption of models of inflorescence architecture and the evolution of floral display. Surprisingly, however, empirical evidence for the trade-off is limited. In particular, there is a lack of phylogenetic evidence for a trade-off between flower size and number. Analyses of phylogenetic independent contrasts (PICs) of 251 angiosperm species spanning 63 families yielded a significant negative correlation between flower size and flower number. At smaller phylogenetic scales, analyses of individual genera did not always find evidence of a trade-off, a result consistent with previous studies that have examined the trade-off for a single species or genus. Ours is the first study to support an angiosperm-wide trade-off between flower size and number and supports the theory that life history constraints have influenced the evolution of floral display.     

荒漠植物准噶尔无叶豆结实、结籽格局及其生态适应意义

连续2a采用野外记录的方式对荒漠植物准噶尔无叶豆的结实、结籽格局进行了比较研究。结果表明:植株内花序生成格局表现为不同大小的花序在总花序中的比重与其结实率存在显著正相关,其中含有中等花数目的花序所占的比重及其结实率均最高,此种分布格局能够最大程度的保证繁殖成功;果序内果实生成格局表现为单花着生位置(从近柄端算起)与其结实量占总结实量的比例成线性负相关,花序基部的结实比例最高,顶部结实比例近为0;荚果内种子生成格局表现为:中间位置的胚珠败育率最低。另外,年际间荚果内仅有(1.08±0.03)粒和(1.07±0.03)粒种子能够完全成熟,此种结籽格局是准噶尔无叶豆保证后代质量的最佳策略。     

Branch length mediates flower production and inflorescence architecture of Fouquieria splendens (ocotillo)

The capacity of individual branches to store water and fix carbon can have profound effects on inflorescence size and architecture, thus on floral display, pollination, and fecundity. Mixed regression was used to investigate the relation between branch length, a proxy for plant resources, and floral display of Fouquieria splendens (ocotillo), a woody, candelabraform shrub of wide distribution in arid North America. Long branches produced three times as many flowers as short branches, regardless of overall plant size. Long branches also had more complex panicles with more cymes and cyme types than short branches; thus, branch length also influenced inflorescence architecture. Within panicles, increasing the number of cymes by one unit added about two flowers, whereas increasing the number of cyme types by one unit added about 21 flowers. Because flower production is mediated by branch length, and because most plants have branches of various lengths, the floral display of individual plants necessarily encompasses a wide range of inflorescence size and structure.     

Genetic constraints on floral evolution in a sexually dimorphic plant revealed by artificial selection

Sexual dimorphism is one of the most widespread and recognizable patterns of phenotypic variation in the biotic world. Sexual dimorphism in floral display is striking in the dioecious plant Silene latifolia, with males making many, small flowers compared to females. We investigated this dimorphism via artificial selection on two populations to determine whether genetic variation exists within populations for flower size and the extent of the between-sex correlation, whether a flower size and number trade-off exists within each sex, and whether pollen and ovule production vary with flower size. We selected for decreased flower size (calyx width) in females and increased flower size in males and measured the response to selection in size and correlated responses in flower dry mass, flower number, and pollen or ovule number per flower. Four bouts of selection in each of two selection programs were performed, for a total of three selection lines to decrease size, three to increase it, and two control lines. Flower size always significantly responded to selection and we always found a significant correlated response in the sex not under selection. Selection decreased but did not eliminate the sexual dimorphism in flower dry mass and number. A negative relationship between flower size and number within each sex was revealed. Whereas ovule number showed a significant correlated response to selection on flower size, pollen number did not. Our results indicate that although substantial additive genetic variation for flower size exists, the high between-sex genetic correlation would likely constrain flower size from becoming more sexually dimorphic. Furthermore, floral display within each sex is constrained by a flower size and number trade-off. Given this trade-off and lack of variation in pollen production with flower size, we suggest that sexual dimorphism evolved via sexual selection to increase flower number in males but not females.     

The effect of flower position on male and female reproductive success in a deceptively pollinated tropical orchid

In many plants, including orchids, differential fruit set along the inflorescence has been attributed to pollinator behaviour. For instance, the pollinator, moving up the inflorescence, becomes satiated with the resources and leaves before visiting the upper flowers. Consequently, the pollinators do not visit flowers as frequently higher up the inflorescence. Alternatively, flower size may vary along the inflorescence, making pollination ineffective as flowers decrease in size. I tested for the presence of differential pollination along the inflorescence in a pollinator-limited tropical epiphyte, Lepanthes rupestris Stimson, and determined the likely cause of the observed pattern. As this species has inflorescences with sequential flowering, pollinator behaviour, moving up the inflorescence as in synchronous multiflowering inflorescences, can be discounted as an explanation for differential fruit set. Fruit set is shown to be more frequent at the base of the inflorescence, but male reproductive success through pollinarium removal is basically independent of flower position. Moreover, cross-pollination by hand at variable flower positions along the inflorescence results in equal fruit set, suggesting that resources are not limiting and cannot explain the cause of differential fruit production along the inflorescence in natural populations. Furthermore, flower size is shown to diminish along the inflorescence, suggesting that the pollinator(s) may be ineffective at depositing the pollinarium in the smaller higher flowers. Consequently, pollinator behaviour and its interaction with flower size, and not resource limitation, is likely to be the main cause of differential fruit set along the inflorescence in L. rupestris .  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 405–410.     

云南草寇花序内不同部位的性分配

两性花植物花序内的性分配常存在差异,资源竞争、结构效应、交配环境(雌雄异熟、传粉者定向访花行为等)或授粉不均匀等几种假说可以解释这种现象。为验证上述假说,该研究以云南草寇两种表型(雄先熟型和雌先熟型)为材料,分析了其花序内不同部位(基部、中部和顶部)的每花花粉数、胚珠数、花粉/胚珠比、结实率和结籽率,花序内传粉者的定向访花行为,以及人工辅助授粉和去花处理对结实率和结籽率的影响。结果表明:两种表型花序内每花花粉数不随部位而变化,每花胚珠数、结实率和结籽率由基部到顶部依次降低,每花花粉/胚珠比由基部到顶部依次增加,表明顶部花存在偏雄的性分配。人工辅助授粉后,结实率、结籽率仍由基部到顶部依次降低,表明授粉不均匀假说不能解释云南草寇花序内不同部位结实率、结籽率的差异。去除基部和中部花后,顶部花人工辅助授粉条件下的结实率、结籽率与基部花人工辅助授粉条件下的结实率、结籽率无差异,表明云南草寇花序内不同部位结实率、结籽率的差异主要由资源竞争引起。雌先熟表型每花花粉数、花粉/胚珠比高于雄先熟表型,表明两种表型存在性分配差异。传粉者主要先访问云南草寇基部的花,然后向顶部移动。云南草寇花序内顶部偏雄的性分配可能是由资源竞争和传粉者定向访花造成的。     

Sexual dimorphism and sex ratios of two Australian dioecious species of alpine pineapple grass, Astelia alpina var. novaehollandiae and Astelia psychrocharis (Asteliaceae)

Sexual dimorphism and male biased sex ratios have been predicted for dioecious plants experiencing the limited conditions for growth and reproduction found in many alpine environments. To test these predictions, the reproductive ecology of two congeneric, co-occurring, dioecious, clonal, species was examined in the subalpine and alpine zones of Kosciuszko National Park, southeastern Australia. Specifically, plant size (vegetative cover of plants in quadrats), floral display (number of flowers per inflorescence, number of inflorescences per quadrat) and sex ratios (proportion of females in quadrats with flowers) were examined in ten populations of Astelia psychrocharis (Asteliaceae) and nine populations of Astelia alpina var. novae-hollandiae (Asteliaceae). Sexual dimorphism did occur, with males having more flowers per inflorescence (106% more flowers for A.alpina males and 12% more for A.psychrocharis males compared to females) and more inflorescences per quadrat than females (78% more inflorescences for A.alpina males and 46% more inflorescences for A.psychrocharis males compared to females). Plant size did not differ between male and female quadrats of either species, nor were there male biased sex ratios. However, plant size was related to flowering status in A.psychrocharis with the 65 quadrats that did not flower having lower vegetative cover than the 175 flowering quadrats indicating that there may be a minimum size/ cover required prior to flowering in this species. For A.alpina, all but two of the 185 quadrats randomly sampled flowered. There was no effect of altitude on plant size and very little effect of altitude on floral display for either species, apart from a slight increase in the number of inflorescences per quadrat with increasing altitude for A.psychrocharis, and slight decrease in number of flowers per inflorescence with increasing altitude for A.alpina females.