片断化生境中滇桐传粉生物学和繁育系统

通过野外观察和人工授粉试验方法,对云南省文山州西畴县法斗分布区片断化生境中濒危植物滇桐(Craigia yunnanensis)的传粉生物学特征和繁育系统进行研究.结果表明:滇桐每个聚伞花序有2～9朵两性花,单花花期为3～4d,单花雌雄蕊在时空上有一定的隔离;杂交指数(OCI) ＞4,花粉-胚珠比(P/O)为1381±53;有效传粉昆虫为大头丽蝇(Chrysomyia megacephala);同株异花授粉结实率低;滇桐的繁育系统属于异交为主,部分自交亲和,传粉过程需要传粉者;自然状态下滇桐的座果率(56.67％±3.85％)和结籽率(6.26％±0.75％)均较低,而异株异花授粉均可显著提高座果率及结籽率(P＜0.01),这与当前片断化生境中传粉昆虫少、效率低下等有关,表明生境片断化正在影响滇桐植株的早期生殖成功.     

珍稀濒危植物细果秤锤树开花生物学特性和繁育系统

为了解珍稀濒危植物细果秤锤树（Sinojackia microcarpa）开花特征和有性繁殖,对其花部形态特征、开花动态、花粉胚珠比（P/O）、柱头可授性、花粉活力、套袋试验、访花昆虫及访花频率进行观测。结果表明：（1）细果秤锤树的杂交指数（OCI）为4,单花期5~7 d,种群花期可持续20 d左右。花粉与胚珠比为4 093.21±498.56。开花后第3天的花粉活力最高（76.21%）,而开花后第7天时花粉活力较低（18.37%）。细果秤锤树柱头最适可授期在开花后第2天。（2）套袋试验表明,细果秤锤树存在部分自交不亲和性,同时不存在无融合生殖,传粉昆虫是其完成生殖过程所必需的,且异株授粉能够提高其坐果率和结籽率。细果秤锤树的访花昆虫有3目5科7种,主要访花昆虫有黄胸木蜂（Xylocopa appendiculata）、熊蜂（Bombus sp.）、胡蜂（Vespa sp.）、中华蜜蜂（Apis cerana）、黑带食蚜蝇（Episyrphus balteatus）、其中熊蜂平均访花频率最高,达（8.67±0.21） 次·h^-1。对该物种开花生物学特征与繁育系统进行深入研究,有利于进一步探究其濒危机制,为后续珍稀濒危植物的研究提供理论依据和参考。     

秦岭岩白菜的传粉生物学特性与繁育系统

对珍稀濒危植物秦岭岩白菜(Bergenia scopulosa T.P.Wang)的开花特性、传粉适应及繁育系统进行了分析研究。结果表明:(1)秦岭岩白菜为蝎尾状聚伞花序,具(29±10)朵单花,单花期约15d,遇低温雨雪天气花冠闭合,可延长3~5d,花序花期约30d,种群花期近4个月。(2)花粉活力在花药开裂后的12h内最高(约90%),维持单花平均花粉活力在30%以上约6d;柱头在第1~4天内保持很强的可授性,维持可授性的时间约为9d。(3)秦岭岩白菜主要有效传粉昆虫为中华蜜蜂,平均访花频率为6.5朵/min,单花停留时间为(11.0±4.8)s。(4)秦岭岩白菜的花粉胚珠比(P/O)为589.8,杂交指数(OCI)为3;人工授粉实验显示,秦岭岩白菜不存在无融合生殖,自交亲和,主动自交罕见,生殖成功主要依赖传粉者。研究认为,秦岭岩白菜是兼性异交的繁育系统,胚珠受精过程中可能存在自交衰退。     

宁夏枸杞传粉生态学初步研究

田间观察检测了宁夏枸杞(Lycium barbarum L.)9个品系的开花物候、开花动态、花粉活力、柱头可授性及其繁育系统,并对花粉萌发、花粉管生长以及访花者的种类和访花行为进行了初步研究.结果表明,宁夏枸杞整个居群有长达5个月的花期,单花的花期仅为3～4 d,花粉活力可保持15 d左右,柱头可授性在72 h左右,雌雄生殖单位在持续时间上有较长的相遇期;花粉数量与胚珠比(P/O)为4 000左右,杂交指数(OCI)介于3或4,繁育系统为异交型,需要传粉者;荧光显微观察表明,参试的3个品系的花粉均能在柱头上萌发,2个有自交不亲和现象,其位置发生在花柱的上半部分和子房内.访花昆虫有17种,分属双翅目、膜翅目和鳞翅目,花的结构和开花式样适合以意大利蜜蜂和食蚜蝇为主的多种昆虫传粉.     

雄全异株流苏树的花部特征及繁育系统研究

雄全异株是自然界罕见的繁育系统。通过野外观察和人工授粉实验对雄全异株植物流苏树传粉生物学特征及繁育系统进行研究。结果表明:流苏树的雄花与两性花的雄蕊发育过程基本一致,均能产生功能花粉粒。两性花的两个心皮原基愈合分化形成雌蕊,雄花的两个心皮原基愈合后形成一个空室并停止发育至整体退化。雌蕊先熟,柱头可授期长,花粉在花药开裂后具有活力,室温下,活力维持在10%以上约2周。流苏树靠风和昆虫(主要是蓟马和食蚜蝇)传粉。控制授粉30 d后,自然对照结实率为34.36%;两性花不存在无融合生殖现象,自交亲和,但自发自交的结实率仅10.70%;人工授粉下杂交结实率显著高于自交(同株异花);有性生殖受到传粉者限制;是混合交配系统。证实流苏树是木犀科又一功能性的雄全异株,其依靠雄株增加异交花粉的数量和质量,避免自交衰退;同时两性花的自交亲和保障生殖成功。流苏树雄花的雌蕊退化,从另一个角度证明木犀科的雄全异株是两性株向雌雄异株进化的过渡状态。     

贯叶连翘的开花动态与繁育系统研究

野外观察贯叶连翘的开花进程和花部形态特征,运用花粉萌发、杂交指数、花粉-胚珠比等方法测定其繁育系统。结果显示:贯叶连翘雌雄异熟,柱头先花药成熟,雌雄蕊无明显异位。单花花期4～5d。仅在开花当日有昆虫传粉,蜜蜂为主要传粉者。花粉在花药开裂1h后活力最大,萌发率达40.10%,花粉在柱头萌发3h后接近子房。根据杂交指数(OCI)推测其繁育系统属于异交,部分自交亲和,有时需要传粉者。花粉-胚珠比(P/O)则表明贯叶连翘的繁育系统为兼性异交。贯叶连翘结实率低,可能与花粉活力,花粉管的生长速度及花粉在柱头的竞争有关。     

黄梅秤锤树孤立居群的遗传多样性及其小尺度空间遗传结构

研究濒危植物片断化孤立居群不同年龄阶段植株的遗传多样性和小尺度空间遗传结构有助于认识残存居群动态和制订保育策略。本研究选取黄梅秤锤树(Sinojackia huangmeiensis)的一个片断化孤立居群(面积160m×80m)为研究对象,对居群内60株成年个体、175株幼树和198株幼苗全部定位,采用8个微卫星位点检测了居群内不同生活史阶段植株的遗传多样性、空间遗传结构,并分析了花粉和种子传播距离和式样。结果表明,黄梅秤锤树居群3种不同年龄阶段植株的遗传多样性之间无显著差异;居群出现显著的杂合子缺失,可能是由近交造成的;在10m以内成年个体、幼树和幼苗植株均呈现出显著的空间遗传结构,说明种子扩散限制于成年母树周边;种子和花粉传播的平均距离分别为9.07±13.38和23.81±23.60m,且花粉和种子传播式样均呈"L"型分布;种子雨重叠少、有限的基因流、自疏以及近亲繁殖是造成各年龄阶段出现空间遗传结构的主要原因。本研究结果启示,在采集秤锤树迁地保护材料时个体间距离应超过10m,以降低采样个体的遗传相似性;同时在就地保护过程中需要人为促进基因流和加强幼苗管理,以降低近交风险。     

濒危植物裸芸香的繁育系统及在植物园迁地保护环境下的传粉研究

迁地保护是防止濒危物种灭绝的一种重要的辅助手段。为了有效保护和管理在中国科学院武汉植物园迁地保护环境下的濒危植物裸芸香（Psilopeganum sinense）,通过定位观察、人工授粉实验等方法对裸芸香的开花生物学、繁育系统和传粉过程进行了研究。结果表明：裸芸香单花花期6～8 d,雄性先熟,单花开放进程中性表达经历了雄性功能期、两性功能期和雌性功能期。裸芸香不存在无融合生殖和风媒传粉,其繁育系统类型为兼性异交,但在无传粉昆虫时能自发自交结实。在武汉植物园内的主要传粉者与宜昌野生居群的传粉者相同,均为双翅目的灰带管蚜蝇（Eristalis cerealis）、细腹食蚜蝇（Sphaerophoria sp.）、黑带食蚜蝇（Episyrphus balteatus）和膜翅目的两种遂蜂（Halictidae）。在植物园自然状况下裸芸香的结实率为90%,不存在生殖障碍。自然自发自交及兼性异交的繁育系统显示裸芸香具有一定的适应生境片断化及环境干扰的潜力。     

珍稀濒危植物陕西羽叶报春的开花及传粉生物学研究

陕西羽叶报春为中国珍稀濒危植物,研究其开花物候、传粉者及访花行为、花粉活力、柱头可授性、繁育系统,以明确陕西羽叶报春的传粉特性及制濒机制。以勉县陕西羽叶报春野生居群为研究对象,通过野外观察和人工授粉实验首次对陕西羽叶报春的传粉生物学进行研究。结果表明:(1)陕西羽叶报春种群花期为2-4月中旬,单花花期一般为14~17 d;单花开花进程中花部形态有6个明显发育变化:全部被萼片包裹、花苞伸出并为深粉色、花朵打开为淡粉色、花冠为白色、花朵枯萎、花冠脱落花萼宿存。(2)陕西羽叶报春为异花授粉植物,蜜蜂属为其主要传粉者。(3)柱头在花粉未开裂前就具有可授性,可授性逐渐增强;花药在花瓣打开后的第3天才开始开裂、散粉,在第4天活力最高,为98.18%,此后逐渐减弱。(4)繁育系统检测显示,长、短柱型花的花粉/胚珠比分别为902.26和831.48,杂交指数为4;套袋实验显示,陕西羽叶报春异型交配可育,同型交配不可育。由此表明,陕西羽叶报春的繁育系统为异交型,自交不亲和,需要传粉者。综上所述,陕西羽叶报春传粉者种类少、自交不亲和、胚珠败育现象可能是导致陕西羽叶报春濒危的重要原因。     

濒危植物凹叶厚朴的花部综合特征和繁育系统

为了解凹叶厚朴(Magnolia officinalis subsp.biloba)濒危的生殖生物学原因,通过野外观测,运用杂交指数、花粉-胚珠比、花粉活力和柱头可授性观测、人工授粉和套袋实验等方法,对凹叶厚朴种群的开花生物学、繁育系统进行了研究。结果表明:凹叶厚朴单花花期一般为4～5d,单花花期根据形态和散粉时间可分为4个时期:蕾期、初开期、盛开期和凋零期;种群于4—5月开花,一般开花历时30d左右,但不同地点、年份间有所差异;种群无开花高峰,单株平均开花数量为171.4～188.8朵,平均开花振幅为5～6朵·株-1·d-1;凹叶厚朴杂交指数为4,花粉胚珠比为5728.86,结合人工套袋和授粉实验结果可确定,繁育系统为异交,传粉过程需要传粉者;花粉活力较高(94%),且花粉活力可维持至散粉后的第6天。散粉后,柱头可授性显著下降,花粉高活力期与柱头可授期有5～6h的重叠;自然状况下结果率和单果出种百分率低,异株异花授粉可明显提高结果率和单果出种百分率,二者相对自然状况分别提高14.25%和43.66%。表明自然状态下传粉效率低、柱头最佳可授期短、同株自花授粉败育严重是其濒危的主要生殖生物学原因。     

山莨菪(茄科)的传粉生物学

茄科的多数种类具有自交不亲和的特点, 主要通过异花传粉结实; 但是, 一些物种或者物种内的部分种群或者个体却高度自交亲合, 转变为自交的繁育系统。该科植物山莨菪(Anisodus tanguticus)主要分布在青藏高原, 开花较早, 比其他晚开花的植物种类更加缺少有效的异花传粉昆虫。我们选择了位于不同海拔高度的2个种群进行比较研究, 主要目的是检验该物种的繁育系统是否在极端环境下由于传粉者的缺乏而发生了部分改变。研究发现,山莨菪的花不完全雌性先熟, 柱头和花药间的平均距离随着花开放时间的延长而不断缩小, 但两者在多数花的单花花期结束时并没有发生接触。因此, 山莨菪花主要表现为适应异花传粉的雌雄异位特征。然而, 少数花 (4.9％)的柱头和花药发生接触, 为“自动自交”的传粉解除了空间隔离。2个种群的多数个体存在自交不亲和机制, 应具有异花传粉的繁育系统; 但是部分个体具有明显的自交亲和能力, 为自交提供了生理基础。高海拔种群的传粉昆虫主要是厕蝇(Fannia sp.), 它们在不同植株间的活动能够保证异花传粉结实; 同时该种群的部分个体存在“自动自交”。低海拔种群的主要访花昆虫是蚂蚁, 它们在花内的活动导致花粉在同一朵花内传递, 而引起“协助自交”; 而异花传粉昆虫厕蝇的访花频率则较高海拔种群低。两个种群的结实均由于异花传粉者不足而受到传粉限制。因此两种不同类型的自交机制为该早期开花植物异花访花昆虫的不足提供了一定程度上的繁殖补偿。     

高山植物扁蕾的延迟自交机制

扁蕾(Gentianopsis barbata)具有鲜艳的花和显著的腺体,并且花开放的前5 d柱头和花药始终处于不同的位置(雌雄异位),这些花综合征表明该植物应为异花传粉。为检验这一假设,我们对青藏高原植物扁蕾的海北站种群进行了3年的传粉生物学研究实验。与花综合征所表明的繁育系统相反,两年的野外观察发现昆虫的访花频率十分低,不去雄并隔离昆虫处理也能产生大量种子,说明这一种群的繁殖主要是依赖于自花传粉。尽管利用种子结实评价的柱头可授性从花开放4 d后开始下降,但随着花的发育进程,雄蕊的伸长能使得花药与柱头完全接触。实验也证明,柱头可授性和花粉活力都超过5 d,说明花药和柱头的接触能够发生自花授粉。扁蕾的这种自花传粉机制应属于典型的延迟自交类型。自花授粉发生在单花花期快要结束前,自交之前仍然保持异交传粉机制,这种延迟自交避免了自交与异交竞争造成的花粉或者种子折损,并为扁蕾在青藏高原极端环境下由于访花昆虫缺乏造成的异交失败提供了繁殖保障。     

慈姑花的开放式样及其花粉流

对一野生慈姑（ＳａｇｉｔａｒｉａｔｒｉｆｏｌｉａＬ．）居群的性表达状态及其花粉流散布规律进行了观测。慈姑雌雄同株,单株雄花数是雌花数的３．６倍。雌雄花单花期均为１ｄ,但二者开放式样不同。尽管雌花的数目较少,但在１ｄ内开放较多的花,其花的开放速率比雄花快。在居群水平上表现出雌雄花比例随时间大幅度波动。在传粉者进行的２６１２次访花活动中,传粉者表现出偏“雌性”访问,对每朵雌花的访问是雄花的１．４倍。用番红染色法标记慈姑的花粉运动表明,花的不同开放式样影响花粉流的散布。当雄花数占高比例时,花粉主要在邻近的个体间散布;而雄花的比例较低、雌花相对较多时,花粉远距离散布的强度加大。     

早春短命植物黑鳞顶冰花的花粉呈现时序及其适应性

花粉的时序呈现是指植物花药按一定次序释放花粉的现象, 被认为是对传粉者访问频率的一种适应。在传粉者充足的环境中, 植物通过限制花粉1次被移出的数量, 使花粉供体能作为多个父本, 从而提高雄性适合度。该文从开花习性、花部特征、传粉者及繁育系统等方面对早春短命植物黑鳞顶冰花(Gagea nigra)的花粉呈现时序及其适应性进行研究, 结果表明, 黑鳞顶冰花单花期约5-7天; 白天开放, 晚上闭合, 花药次序开裂, 呈拉链式散粉, 散粉期4-6天。黑鳞顶冰花以异交为主, 部分自交亲和。蝇类和食蚜蝇为主要传粉者, 访花频率为(0.141±0.078) flower∙h^-1。在雄蕊的时序散粉过程中, 雌蕊持续生长, 经历了从低于雄蕊到等高、再到高于雄蕊阶段。在等高阶段, 单花早晚的开闭, 使得雌雄蕊紧靠在一起, 促成了自动自花授粉。在传粉者缺乏的环境中, 黑鳞顶冰花的花粉时序呈现延长了散粉期, 在等待传粉者和分摊风险方面具重要作用。这种花粉渐次呈现的策略, 在新疆的早春开花植物中可能广泛存在。     

Effects of pollination and pollen source on the seed set ofPedicularis palustris

Habitat alteration can deteriorate plant-pollinator interactions and thereby increase the risk of population extinction. As part of a larger study on the effects of changes in land use on fen grassland vegetation, factors influencing the seed set of a short-lived, endangered wetland plant,Pedicularis palustris, were studied. We conducted field pollination experiments in one large and one small population. To investigate the effect of pollen source on seed set, individual flowers of caged plants were left unpollinated or were pollinated with pollen from the same flower, the same population or another population. To study pollen limitation and flower display, whole plants were subjected to pollinator exclosure, hand pollination or natural pollination. Self-compatibility was high, but differed between populations (61% and 97% of seed set after cross-pollination within populations). Cross-pollination between populations did not significantly alter seed number per capsule. Pollinator exclosure resulted in a very low seed set (<15% of natural seed set), despite high self-compatibility. The most likely explanations for high self-compatibility in combination with low autofertility are geitonogamy as reproductive assurance, selective neutrality of self-compatibility and phylogenetic constraints. Because of low autofertility, the seed set inP. palustris depends on pollinators. In the study populations, natural pollination was clearly sufficient for maximum seed production per plant, but seed set per capsule was significantly pollen-limited in the smaller population. Plants in this population also had a higher maximum percentage of simultaneously open flowers than those of the large population (31% vs. 13%), while flower longevity was generally extended without pollination. It is concluded thatP. palustris may influence pollinator behaviour and therefore the risk of pollen limitation by flower display.P. palustris showed a flexible reaction to differing pollination regimes without losses in overall seed set in the study populations.     

Sexual differences in reproductive characters and pollinator attractiveness in gynodioecious Glechoma longituba (Lamiaceae)

We quantified the differences in floral characters and attractiveness to flower visitors under natural conditions between the sexual types in the gynodioecious plant Glechoma longituba. We also manipulated flowers by altering corolla size or nectar volume, or by removing anthers, to examine the effect of these primary and secondary attractants (i.e. rewards and advertisements) on attractiveness. A change in corolla size and shape reduced visiting frequency and pollen load. Removal of anthers did not affect visiting rates, but significantly reduced pollination rates and stigmatic pollen load. A decrease in the nectar volume of a flower was associated with a reduction in handling time and pollen loads on stigmas. These results show that corolla size is an important advertisement to pollinators (particularly at greater distance), which associate hermaphrodite flowers with a larger corolla and a larger volume of nectar than female flowers. We found that artificial changes in population structure affected the behavior of pollinators as well as the pollination rates of flowers. We suggest that the pattern of distribution of hermaphrodite and female clones in a population may serve to avoid pollen limitation in a female clone or patch. This effect may ensure female reproductive success and allow for the maintenance of female individuals in natural populations of this gynodioecious plant.     

草豆蔻传粉生物学的研究

比较研究了花柱卷曲性植物草豆蔻 Alpinia hainanensis两种表型(上举型和下垂型)的传粉生物学特性。结果表明:草豆蔻的花柱卷曲运动节律与其他已报道的山姜属Alpinia植物基本一致,而下垂型花的花柱卷曲运动明显滞后于上举型花约2 h。草豆蔻的花柱卷曲运动中存在一些不同步的现象,甚至在同一花序上的两朵花的花柱运动节律亦会表现出不一致的现象,但上举型花的花药开裂时间却严格同步,都发生在同类型个体的柱头全部位于花药上方之后进行。草豆蔻上举型花的花蜜分泌量、单花花粉量、花粉/胚珠比率(P/O)均明显比下垂型花多,而两种表型的胚珠数、花蜜糖含量、氨基酸含量无显著差异。在草豆蔻的单花期内不同时间段进行人工授粉,上举型花在其柱头位于花药下方时进行人工授粉,其结实率明显比柱头位于花药上方时人工授粉的处理高,下垂型花则没有显著差异。在自然居群中,草豆蔻的主要传粉者是无垫蜂Ameglla sp.和两种木蜂Xylocopa sp.,但存在传粉者不足而影响结实率的现象。完全套袋、去雄和去雌(去柱头)处理的均不结实,表明草豆蔻中不存在无融合生殖、主动自花授粉和滞后自交的生殖保障现象。而人工自交和异交均具有较高的结实率,表明草豆蔻为自交亲和植物。草豆蔻的繁育系统是具花柱卷曲性运动的异花授粉的交配系统。     

Change of floral orientation affects pollinator diversity and their relative importance in an alpine plant with generalized pollination system,Geranium refractum (Geraniaceae)

Floral orientation may affect pollinator attraction and pollination effectiveness, and its influences may differ among pollinator species. We, therefore, hypothesized that, for plant species with a generalized pollination system, changes in floral orientation would affect the composition of pollinators and their relative contribution to pollination. Geranium refractum, an alpine plant with downward floral orientation was used in this study. We created upward-facing flowers by altering the flower angle. We compared the pollinator diversity, pollination effectiveness, and pollinator importance, as well as female reproductive success between flowers with downward- and upward-facing orientation. Results indicated that the upward-facing flowers were visited by a wider spectrum of pollinators (classified into functional groups), with higher pollinator diversity than natural flowers. Moreover, due to influences on visitation number and pollen removal, the pollinator importance exhibited by the main pollinator groups differed between flower types. Compared with natural flowers, the pollination contribution of principal pollinators (i.e., bumblebees) decreased in upward-facing flowers and other infrequent pollinators, such as solitary bees and muscoid flies, removed more pollen. Consequently, stigmatic pollen loads were lower in upward- than in downward-facing flowers. These findings reveal that floral orientation may affect the level of generalization of a pollination system and the relative importance of diverse pollinators. In this species, the natural downward-facing floral orientation may increase pollen transfer by effective pollinators and reduce interference by inferior pollinators.     

Reproductive Biology of the Andean Shrub Discaria nana (Rhamnaceae)

Abstract: The widely-held view that the frequency of self-compatibility increases at higher elevations has been questioned for communities of the southern Andes. The study of pollination biology of obligate outcrossers with wide altitudinal range may provide clues on how plants can remain reproductively successful in increasingly hostile environments without resorting to selfing. I studied the phenology, pollinator assemblages, breeding system and maternal success of the perennial entomophile outcrosser, Discaria nana, across a 1200-m elevation span in the Andes of Mendoza and Neuquén, Argentina (34° - 37°S). D. nana behaved as self-incompatible throughout the studied gradient. At the higher site proportionally fewer flowers were pollinated, stigmatic loads were smaller and number of visitor species was lower than at low altitude; however, natural fruit set was 3.6 times higher and seed set 3.3 times higher. As a non-exclusive explanation, it is proposed that increased maternal success reflected better pollination quality at the higher site, to which several factors seemed to contribute. At higher altitude D. nana a) exhibited mutualisms with a high proportion of the available pollen vectors, b) had a higher rate of specialized pollinators (Lepidoptera, obligate nectar feeders) to unspecialized ones (Diptera, pollen and mixed feeders) in the visitor assemblage which would reduce ineffective (i.e. self) pollination because nectar feeders visited both male and female phase flowers and pollen feeders favoured male phase flowers, c) had an extended floral longevity (especially the female phase), and d) showed an increased floral display (as number of flowers per unit plant area). Differential rewarding through more concentrated nectar may explain the change in assemblage composition. The flowers' reverse herkogamy (i.e. the location of the stigma below the anthers) seemed to enhance the beneficial effects of the abundance of more effective pollinators.     

FLORAL PARAMETERS,BREEDING SYSTEM AND POLLINATOR TYPE IN TRICHOSTEMA (LABIATAE)

Diversity in flower size and degree of exsertion of anthers and stigma from the corolla in the California species of Trichostema appear to be correlated with breeding system and pollinator type (bee vs. bird). Autogamous (self-pollinating) species unlike xenogamous (cross-pollinating) species lack spatially separate anthers and stigma and nototribic pollination. The outcrossing species have significantly larger flowers, significantly larger nectar volumes and significantly higher pollen-ovule ratios than do selfing species. Because autogamous species are less reliant on flower visitors to facilitate pollination, there may be relaxed selection for large nectar producing flowers. Pollen-ovule ratios are correlated with breeding system and reflect the efficiency of pollen transport. Data on floral parameters suggest xenogamous species expend more energy per floral unit and are less efficient seed producers than closely related autogamous species.