高山植物全缘叶绿绒蒿在不同海拔地区的传粉生态学研究北大核心CSCD

为了探明高山植物全缘叶绿绒蒿(Meconopsis integrifolia)的繁育系统特点和其对高山气候环境的繁殖适应特征,我们沿海拔梯度选择了5个样地(样地1(4 452 m)、样地2(4 215 m)、样地3(4 081 m)、样地4(3 841 m)、样地5(3 681 m))对其传粉生态学进行了连续2年的观察试验。结果发现,样地1、2的全缘叶绿绒蒿的单花寿命显著长于样地3、4和5。花开放早期柱头高于花药,之后花药不断伸长,并在开放中后期与柱头接触,说明全缘叶绿绒蒿具有不完全雌雄异位的花部特征。自然状态下,柱头可授能力持续期约8天(雌蕊先熟),但花药于开花第5天才散粉,花粉寿命约2天,说明全缘叶绿绒蒿为雌雄异熟,但存在一定的重叠期。人工授粉试验表明,全缘叶绿绒蒿自交部分亲和,且不存在无融合生殖现象。各样地中自然对照的结实率显著低于人工异交处理的结实率,说明自然状态下全缘叶绿绒蒿存在一定程度的传粉限制。传粉昆虫观察发现,样地1和2的传粉昆虫主要是蝇类,样地3、4和5的传粉昆虫主要是蝇类和蓟马(Thripidae spp.),蝇类在不同植株间的活动能够保证异花传粉结实,同时,蝇类和蓟马在花内的活动会引起"协助自交"。全缘叶绿绒蒿约65%的观察个体存在"自动自交"。蝇类在各样地的访花频率存在显著差异,样地1访花频率最低,样地2访花频率最高。各样地的结实由于异花传粉者的不足而受到传粉限制。两种不同类型的自交机制恰恰为该植物异花传粉者不足提供了一定程度的繁殖补偿。全缘叶绿绒蒿不分泌花蜜,当环境温度降低时,采取为昆虫提供保温庇护场所的方式来吸引传粉者。     

高山植物全缘叶绿绒蒿在不同海拔地区的传粉生态学研究

为了探明高山植物全缘叶绿绒蒿(Meconopsis integrifolia)的繁育系统特点和其对高山气候环境的繁殖适应特征, 我们沿海拔梯度选择了5个样地(样地1 (4452 m)、样地2 (4215 m)、样地3 (4081 m)、样地4 (3841 m)、样地5 (3681 m))对其传粉生态学进行了连续2年的观察试验。结果发现, 样地1、2的全缘叶绿绒蒿的单花寿命显著长于样地3、4和5。花开放早期柱头高于花药, 之后花药不断伸长, 并在开放中后期与柱头接触, 说明全缘叶绿绒蒿具有不完全雌雄异位的花部特征。自然状态下, 柱头可授能力持续期约8天(雌蕊先熟), 但花药于开花第5天才散粉, 花粉寿命约2天, 说明全缘叶绿绒蒿为雌雄异熟, 但存在一定的重叠期。人工授粉试验表明, 全缘叶绿绒蒿自交部分亲和, 且不存在无融合生殖现象。各样地中自然对照的结实率显著低于人工异交处理的结实率, 说明自然状态下全缘叶绿绒蒿存在一定程度的传粉限制。传粉昆虫观察发现, 样地1和2的传粉昆虫主要是蝇类, 样地3、4和5的传粉昆虫主要是蝇类和蓟马(Thripidae spp.), 蝇类在不同植株间的活动能够保证异花传粉结实, 同时, 蝇类和蓟马在花内的活动会引起“协助自交”。全缘叶绿绒蒿约65%的观察个体存在“自动自交”。蝇类在各样地的访花频率存在显著差异, 样地1访花频率最低, 样地2访花频率最高。各样地的结实由于异花传粉者的不足而受到传粉限制。两种不同类型的自交机制恰恰为该植物异花传粉者不足提供了一定程度的繁殖补偿。全缘叶绿绒蒿不分泌花蜜, 当环境温度降低时, 采取为昆虫提供保温庇护场所的方式来吸引传粉者。     

红棕杜鹃不同海拔种群的表型多样性研究

为了揭示高山杜鹃群体在不同海拔梯度上的表型变异程度和变异规律,以云南轿子山国家级自然保护区天然分布的5个海拔梯度红棕杜鹃种群为研究对象,通过对其12个性状的测量,分析不同海拔梯度下红棕杜鹃种群的表型多样性。结果表明:(1)调查的12个性状中5个存在显著差异,占总表型性状的41.67%;各性状变异系数(CV)的大小分布在8.51%~34.32%之间;相对极差(R′)分布在38.04%~100%之间,低海拔种群(P1和P2)的表型性状变异丰富、极端变异高,而中、高海拔种群(P3、P4、P5)的表型性状变异和极端变异程度较低。(2)Shannon-Weaver多样性指数分析表明,轿子山红棕杜鹃不同海拔种群存在丰富的表型多样性,5个海拔种群的平均表型多样性指数为2.453 4,12个表型性状的平均多样性指数为4.076 3。(3)相关性分析表明,花序宽和花柄长与海拔呈显著负相关,其它性状与海拔相关性并不显著。(4)聚类分析表明,红棕杜鹃不同海拔种群可以聚为明显的2支。研究认为,轿子山自然保护区红棕杜鹃不同海拔种群的环境异质性是影响其表型变异的主要因素。     

高山植物全缘叶绿绒蒿叶片形态及解剖结构对海拔的响应

为了阐明高山植物全缘叶绿绒蒿(Meconopsis integrifolia)对海拔及高山环境的形态学适应特征,沿海拔梯度选择了5个海拔的全缘叶绿绒蒿分布区域(3681、3841、4081、4215和4452 m),采用常规石蜡制片技术和显微观察方法,对其形态及叶片解剖结构进行了研究。结果表明:随着海拔的升高,株型方面,全缘叶绿绒蒿的株高、基径逐渐减小;叶片形态方面,叶片逐渐变小,叶形逐渐狭长,具体表现为叶片长度、宽度逐渐减小,叶片长宽比逐渐增大;叶片解剖结构方面,叶片厚度、表皮细胞厚度、海绵组织厚度、组织疏松度及中脉直径呈减小趋势;栅栏组织厚度、栅海厚度比、组织紧密度呈增大趋势;叶表皮结构方面,叶片表皮毛密度、气孔密度及气孔指数均呈增大趋势。此外,解剖结构指标之间大多呈现出明显的协同进化,各形态指标对海拔的变化表现出较大的可塑性。全缘叶绿绒蒿形态及叶片解剖结构在不同海拔上表现出的这种差异,可能是植物长期适应高山复杂环境的结果。     

茶条槭不同海拔种群的表型多样性

为了揭示茶条槭（Acerginnala）不同海拔种群表型变异程度和变异规律,以山西七里峪天然分布的茶条槭为研究对象,调查不同海拔种群的种实和叶表型性状,采用方差分析、相关分析、聚类分析等方法,分析种群的表型多样性。结果表明：17个表型性状中16个存在显著差异,占总表型性状的94.12%。在物种水平上各个性状表现出较丰富的变异,变异系数（CV）在7.05%～38.12%之间。茶条槭种群具有高的表型多样性（1.9253）,5个不同海拔种群的平均表型多样性指数为1.9022～1.9837。种群间的表型分化系数均值（13.79%）小于种群内变异（82.71%）,种群内的变异是表型变异的主要来源。各表型性状及表型多样性指数与土壤中的N、K、AN、AK、AP、OR、MC表现出显著或极显著的相关关系（P〈0.05）,但与海拔高度呈现出不显著的相关性。UPGMA聚类分析显示5个种群形成明显的两组,与其地理分布相一致。不同海拔种群所处微生境的异质性是引起种群间差异的主要原因。茶条槭种群内、种群间变异的利用对其遗传改良具有重要的意义。     

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。     

高海拔地区不同海拔对菊花植株生长与品质的影响

通过田间测定引种栽培于四川省宝兴县不同海拔高度菊花植株的株高、地径、分枝数等营养生长指标及花数量、花重量、花直径、花厚度等生殖生长指标,采用高效液相色谱法测定菊花样品的绿原酸含量、木犀草苷含量、异绿原酸A含量等主要品质指标,并对测得数据进行方差分析和相关性分析,研究比较了高海拔地区不同海拔高度对菊花植株生长与品质的影响。结果表明:海拔梯度对菊花株高、花数、花重量、花直径、花厚度、木犀草苷含量和异绿原酸A含量均有显著影响(P0.05),对地径、分枝数和绿原酸含量无显著影响(P0.05),其中株高、花数和花直径与海拔梯度相关性显著(P0.05)。除分枝数和地径外,高海拔地区不同海拔高度对菊花植株的生长影响较大,在一定海拔范围内(2051~2 405m),海拔高度对菊花品质形成的影响较小,据2010版《药典净对菊花有效成分最低限量标准,海拔2051~2598 m所产的菊花均符合药用条件,以海拔2 329 m左右所产的菊花品质较佳。研究结果可为高海拔地区引种栽培菊花提供理论依据。     

麦瓶草种群中的花药黑粉菌感染：花的形态学变异、菌孢子散布式样及病原菌—传媒调节性选择

花药黑粉菌（Microbotryum violaceum)可系统侵染其寄主植物麦瓶草（Silene dioica),使其不育而代之以在植物花中布满菌孢子,这些菌孢子主要由花传媒昆虫带到健康植株,花传媒昆虫对花部特征变异的行为反应不仅可能影响花粉的输入/输出率,而且影响菌孢子的着落率和植株的发病与否,为研究S.dioica雌株花部性状特征与花粉传布,菌孢子着落及由此而导致的植株染病之间的相关性,用采自4个不同S.dioica种群的植物进行了移栽实验,讠4个种群均位于瑞典北部的Skeppsvik群岛,代表了从健康（岛1,种群1）,低度发病（岛2,种群2）到高度发病（岛3、4,种群3、4）的发病梯度,从上述4个不同种群中各采50个植株移栽至岛4的中部,来自健康种群的植株较来自高发病种群的植株具有较大的花,较长的花柱,研究发现,着落在花上的花粉粒数和菌孢子数呈强正相关,来自健康种群的植株每朵花上着落的花粉粒和菌孢子数分别是高发病种群（种群4）植株的4倍和9倍,导致来自4个不同种群的植株的发病率存在着显差异。种群1、2、3、4的发病率分别为37%,20%,18%,0。在涉及10个种群的田间调查研究中,发现种群中健康植株的平均花柱长度（与花冠大小,胚珠数目正相关）与植株的发病率显负相关,讨论了病原体-传媒调节的植物花部性状特征选择潜势及其导致的麦瓶草S.dioica种群音质 基因漂移。     

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。     

露蕊乌头花性状的表型选择

为解释植物的花性状的进化机制,分析选择作用的强度和方向是必须的方法之一。采取表型操控实验和表型选择分析,通过人为增加露蕊乌头侧萼片开度,本文揭示了露蕊乌头花性状与适合度的关系,探究了花性状的表型选择作用。结果发现,增加侧萼片开度后,传粉者首访的频率及访花的总次数降低,但对单花种子产量没有影响;露蕊乌头的株高、花数目、盔高及侧萼片宽度均受到总选择作用,而花数目和株高的选择梯度显著,受到了直接选择作用;而花数目则受到稳定选择,即中等开花数目具有更高的种子产量。以上结果表明,侧萼片宽度是露蕊乌头吸引传粉者的主要特征,而花序越高、花数目适中的植株,种子产量也越多。     

海拔对青藏高原东部毛茛科植物繁殖特征和资源分配的影响

通过野外采样对青藏高原东部高寒草甸上两个海拔间3种常见毛茛科植物条裂银莲花(Anemone trullifo-liavar.linearis)、粗距翠雀花(Delphinium pachycentrumHemsl.)和钝裂银莲花(Anemone obtusiloba)的繁殖性状和资源分配进行了研究,并对植物在极端环境下采取的繁殖策略及繁殖模式进行了探讨.结果显示:(1)3个物种的株高都随着海拔的升高而降低;同一物种的花大小在不同海拔间均无显著差异;条裂银莲花单个种子重随海拔的升高而增加,但虫食数/株、结籽率和种子数/株均随着海拔的升高而减少;钝裂银莲花的繁殖分配、虫食数/株、种子数/株和单个种子重均随着海拔的升高而减小.(2)不同海拔条裂银莲花的个体大小与单花重、雄蕊重、虫食数/株均呈显著正相关,且回归斜率在海拔间有显著差异.粗距翠雀花花期的雄蕊重与个体大小呈显著正相关,但这种异速关系不受海拔和个体大小的影响;钝裂银莲花的个体大小与花期的所有繁殖特征以及果期的种子重/果实均呈显著正相关,不同海拔间个体大小与种子重/果实的斜率差异显著,且与花期各繁殖性状异速关系的截距差异显著.研究表明海拔对植物的个体大小以及种群间的繁殖对策和繁殖成功率有着重要的影响.     

Selection by Pollinators on Floral Traits in Generalized Trollius ranunculoides (Ranunculaceae) along Altitudinal Gradients

Abundance and visitation of pollinator assemblages tend to decrease with altitude, leading to an increase in pollen limitation. Thus increased competition for pollinators may generate stronger selection on attractive traits of flowers at high elevations and cause floral adaptive evolution. Few studies have related geographically variable selection from pollinators and intraspecific floral differentiation. We investigated the variation of Trollius ranunculoides flowers and its pollinators along an altitudinal gradient on the eastern Qinghai-Tibet Plateau, and measured phenotypic selection by pollinators on floral traits across populations. The results showed significant decline of visitation rate of bees along altitudinal gradients, while flies was unchanged. When fitness is estimated by the visitation rate rather than the seed number per plant, phenotypic selection on the sepal length and width shows a significant correlation between the selection strength and the altitude, with stronger selection at higher altitudes. However, significant decreases in the sepal length and width of T. ranunculoides along the altitudinal gradient did not correspond to stronger selection of pollinators. In contrast to the pollinator visitation, mean annual precipitation negatively affected the sepal length and width, and contributed more to geographical variation in measured floral traits than the visitation rate of pollinators. Therefore, the sepal size may have been influenced by conflicting selection pressures from biotic and abiotic selective agents. This study supports the hypothesis that lower pollinator availability at high altitude can intensify selection on flower attractive traits, but abiotic selection is preventing a response to selection from pollinators.     

Factors affecting pollinator movement and plant fitness in a specialized pollination system

The rate of pollen exchange within and among flowers may depend on pollinator attraction traits such as floral display size and flowering plant density. Variations in these traits may influence pollinator movements, pollen receipt, and seed number. To assess how floral display size and flowering plant density affect parameters of pollinator visitation rate, pollen receipt per flower, seed number per fruit and the between-plant pollinator movements, we studied the self-incompatible plant, Nierembergia linariifolia. Per-flower pollinator visitation rate and bout length increased linearly with increasing floral display size. Pollen receipt per flower increased linearly with increasing flowering plant density. For seed number per fruit, a polynomial model describing an increased seed number per fruit at low density and a decreased seed number per fruit at high density provided a significant fit. Per-flower pollinator visitation rate was not associated with pollen receipt per flower and seed number per fruit. Bees visited plants located near to the center of the population more frequently than plants located at the periphery. Increases in both floral display size and flowering plant density led to an increased chance of a plant being chosen as the center of the pollinator foraging area. These results suggest that even though large floral displays and high flowering plant density are traits that attract more pollinators, they may also reduce potential mate diversity by restricting pollen movement to conspecific mates that are closely located.     

展毛翠雀的花性状表型选择

传粉者的选择作用是花表型性状进化的重要驱动力, 解析选择作用的强度是理解花进化的关键。通过表型操控实验和表型选择研究能够分析花性状与其适合度的关系, 探究花性状的表型选择作用。为揭示花性状变化对雌性适合度的影响, 本研究处理展毛翠雀(Delphinium kamaonense var. glabrescens)花萼片大小, 并进行表型选择分析。结果表明: 人为减小展毛翠雀花萼片显著降低了传粉者的访花频率, 但是并没有影响种子产量(种子数和结籽率), 说明展毛翠雀花萼片的大小不影响种子产量, 可能主要吸引传粉昆虫输出花粉。通过雌性适合度(种子数量)估计表型选择梯度, 发现花萼片大小(长和宽)没有受到显著的直接选择梯度。但是, 花距长受到显著的线性选差和选择梯度, 表明花距的延长能够增加种子产量。本研究表明展毛翠雀花性状受到选择的作用, 但萼片和花距有不同的功能, 分别影响传粉者访问频率和种子产量。     

Variations of flower size and reproductive traits in self-incompatible <Emphasis Type="Italic">Trollius ranunculoides</Emphasis> (Ranunculaceae) among local habitats at Alpine Meadow

For plants that rely on animals for pollination, the ability to attract the animals to their flowers can be a crucial component of fitness. A large number of studies have documented pollinators to be important selective agents driving the evolution of flower size and correlated traits on a large scale. In this paper, we studied variations of reproductive traits in self-incompatible Trollius ranunculoides (Ranunculaceae) among local habitats at Alpine Meadow. The results showed significant variations of floral size, seed mass per fruit and sex allocation (male/female mass ratio) between different habitats, where floral size and seed mass was not explained fully by variation of plant size among habitats. It suggested that other factors unrelated to plant size might also influence floral variation. However, in our manipulated experiment, it showed no effects of manipulated floral size not only on visit rate of effective pollinators (bees and flies) but also on female success (seed set, seed mass per fruit), irrespective of flower density. Consequently, we could not conclude that the variation of floral size in T. ranunculoides was due to phenotypic plasticity, or natural selection. But if selection occurred, it should not be mediated by pollinators. It was likely that variation of sex allocation between habitats lead to changes of flower or corolla size, because plant invested much less to male function (female-biased sex allocation and larger single seed mass) in shade habitat (bottom of bush) than other exposed habitats, to gain higher fitness. In addition, high-floral density in T. ranunculoides had a negative effect on service of main pollinator (bees) and female success. This situation would influence the strength of selection on floral size.     

MEASURING POLLINATOR-MEDIATED SELECTION ON MORPHOMETRIC FLORAL TRAITS: BUMBLEBEES AND THE ALPINE SKY PILOT,POLEMONIUM VISCOSUM

Sweet-flowered plants of Polemonium viscosum in Colorado are visited by a fly-dominated pollinator fauna at timberline (krummholz), but almost exclusively by bumblebees in higher-elevation tundra habitats. Significant increases in flower size and height are associated with increasing elevation along this habitat gradient. This paper presents the results of an experiment designed to test whether bumblebees exert sufficient selection on morphometric floral phenotypes to account for the clinal shifts seen in natural populations. Two populations of sweet-flowered plants of krummholz origin were established: one randomly pollinated, the other solely bumblebee-pollinated. I tested the effects of two independent axes of floral variation, obtained by principal-components analysis, on mean seed set per flower of plants in each population. PC1, with strong correlations to corolla diameter, corolla length, and stem height, explained a significant amount of variance in seed set for bumblebee-pollinated plants but had no bearing on that of randomly pollinated plants. PC2, with strong correlation to flower number, did not influence seed set in either population. Bumblebee behavior was correlated with variation in PC1 scores of the selected population, yielding positive directional selection on morphometric floral traits associated with PC1. Selection coefficients for PC1, corolla length, corolla diameter, and inflorescence height were estimated, respectively, as 0.11, 0.09, 0.07, and 0.06 (P < 0.025 in all cases). These results support the hypothesis that pollinator-mediated selection can bring about changes in floral form, and can explain shifts in floral morphology of P. viscosum along natural habitat gradients.     

Variation in and adaptive plasticity of flower size and drought-coping traits

Although phenotypic plasticity of morphological and physiological traits in response to drought could be adaptive, there have been relatively few tests of plasticity variation or of adaptive plasticity in drought-coping traits across populations with different moisture availabilities. We measured floral size, vegetative size, and physiological traits in four field populations of Leptosiphon androsaceus (Polemoniaceae) that were distributed across a rainfall gradient in California, USA. Measurements were made over 5 years that varied in precipitation. We also conducted a growth chamber experiment in which half-sibs from three populations were divided equally among a well-watered and a drought treatment. We tested for selection on traits in each of the watering treatments, and evaluated whether traits exhibited plasticity. In the field, plant traits exhibited substantial variation across populations and years. Flower size, leaf size, and water-use efficiency (WUE) were generally higher for populations that received greater average rainfall. However, in dry years, we observed a decrease in flower and leaf size, but an increase in WUE across the populations. In the growth chamber experiment, leaf and physiological traits exhibited plasticity, with smaller leaves and higher WUE found in the drought, as compared to the well-watered treatment. Only specific leaf area exhibited differentiation in plasticity among populations. Although there was no observed plasticity in floral size, selection favored smaller flowers in the drought treatment and larger flowers in the well-watered treatment. Our results suggest that moisture availability has led to trait variation in L. androsaceus via a combination of selection and phenotypic plasticity.     

Floral stability,pollination efficiency,and experimental manipulation of the corolla phenotype in Nemophila menziesii (Hydrophyllaceae)

The present study examines the degree of phenotypic stability in vegetative and floral traits in a population of the annual, self-compatible, and protandrous Nemophila menziesii (Hydrophyllaceae) and examines whether local pollinators select for a uniform floral phenotype in this population. I found a strong positive association between flower number, leaf size, shoot length, and pedicel length, implying a general effect of overall plant vigor on this suite of traits, while characters reflecting corolla shape were phenotypically stable across a broad range of plant sizes. However, results of a manipulative experiment indicate that individual flowers maintain high pollination success despite drastic changes in the size and shape of the corolla. This suggests that pollinator-mediated selection does not directly favor floral stability in N. menziesii or that selection was weak during the course of this study. Seed number per flower was positively correlated with plant size, which varied greatly from one patch to another.     

MEASUREMENTS OF NATURAL SELECTION ON FLORAL TRAITS IN WILD RADISH (RAPHANUS RAPHANISTRUM). I. SELECTION THROUGH LIFETIME FEMALE FITNESS

Although the role of natural selection in the evolution of floral traits has been of great interest to biologists since Darwin, studies of selection on floral traits through differences in lifetime fitness have been rare. We measured selection acting on flower number, flower size, stigma exsertion, and ovule number per flower using field data on lifetime female fitness (seed production) in wild radish, Raphanus raphanistrum. The patterns of selection were reasonably consistent across three field seasons, with strong directional selection for increased flower production in all three years, weaker selection for increased ovule number per flower in two years, and selection for increased flower size in one year. The causes of the selection were investigated using path analysis combined with multiplicative fitness components. Increased flower production increased fruit production directly, and increased numbers of ovules per flower increased the number of seeds per fruit in all three years; pollinator visitation did not influence either of these fitness components. Increased flower size was associated with increases in both the number of fruit and the number of seeds per fruit in one year, with the latter relationship being stronger. Total lifetime seed production was affected more strongly by differences in fruit production than by differences in either the number of seeds per fruit or the proportion of fertilized seeds that were viable, but all three fitness components were positively correlated with total seed production.     

The quantitative genetics of floral trait variation in Lobelia: potential constraints on adaptive evolution

Abstract Although pollinator-mediated natural selection has been measured on many floral traits and in many species, the extent to which selection is constrained from producing optimal floral phenotypes is less frequently studied. In particular, negative correlations between flower size and flower number are hypothesized to be a major constraint on the evolution of floral displays, yet few empirical studies have documented such a trade-off. To determine the potential for genetic constraints on the adaptive evolution of floral displays, I estimated the quantitative genetic basis of floral trait variation in two populations of Lobelia siphilitica . Restricted maximum likelihood (REML) analyses of greenhouse-grown half-sib families were used to estimate genetic variances and covariances for flower number and six measures of flower size. There was significant genetic variation for all seven floral traits in both populations. Flower number was negatively genetically correlated with four measures of flower size in one population and three measures in the other. When the genetic variance-covariance matrices were combined with field estimates of phenotypic selection gradients, the predicted multivariate evolutionary response was less than or opposite in sign to the selection gradient for flower number and five of six measures of flower size, suggesting genetic constraints on the evolution of these traits. More generally, my results indicate that the adaptive evolution of floral displays can be constrained by tradeoffs between flower size and number, as has been assumed by many theoretical models of floral evolution.