雄蕊合生植物半边莲的花部综合征与繁育系统

 为了解花内雄蕊合生结构的繁殖适应意义, 初步研究了雄蕊合生植物半边莲(Lobelia chinensis)的花部综合征、传粉特性和繁育系统。半边莲花大且鲜艳, 花瓣中下部弯折并合生成背部有裂缝的不封闭的花冠筒。5雄蕊的花药紧密合生成花药筒, 花丝中上半部也合生在一起, 只有花丝基部分离插生于花冠筒上。柱头被包裹在花药筒内。半边莲单花寿命可达5 d左右。雄性先熟, 柱头在伸出花药筒之后才具活性。花的主要访问者为蚂蚁、食蚜蝇和苍蝇类等小型昆虫。半边莲单花花粉数约为(5 200±130)粒、胚珠数约为(55±6)颗, 花粉胚珠比为94.54, 应属于兼性自交繁育系统, 但异交指数和其它特征都显示其以异交为主, 部分自交亲和。套袋和人工授粉实验发现, 半边莲不存在无融合生殖与自发自交, 但自交亲和性高。雄蕊合生(尤其是花药的合生)能把花药中的花粉聚拢在一起在传粉者的一次访问中就被同时带出, 与同样具有较低花粉胚珠比的花粉聚联(Pollen aggregation)传粉过程近似。半边莲的雄蕊合生结构(花药合生成筒、花丝上部也合生)可能与一些特定的花部特征, 如花被合生成未完全封闭的筒、雌雄异熟以及低花粉胚珠比等联系在一起, 形成了适应小型传粉者的“花部综合征”     

3种雌雄异熟樟科植物开花动态的比较研究

为了解樟科植物雌雄异熟的繁育系统特点,对3种樟科植物阴香(Cinnamomum burmannii)、紫楠(Phoebe sheareri)和浙江楠(Phoebe chekiangensis)雌雄异熟花的开花动态进行了比较研究。结果表明,3种植物雌性功能期的开始时间、雌性功能期和雄性功能期的时间分配有差异。3种植物的主要传粉者均为中华蜜蜂,在雌性功能期和雄性功能期均有访花;在雌性功能期第四轮退化雄蕊蜜腺和雄性功能期第三轮花丝基部的蜜腺分泌花蜜,不同开花阶段的花蜜蜜量和含糖量没有显著差异;柱头可授性和花粉活力在雌性和雄性功能期均较高;雌性功能期和雄性功能期均有自然花粉落置;座果率均是异交补粉自然套袋,表明都存在一定的传粉限制。因此,樟科植物雌、雄性阶段时间分配的不同,可能与退化雄蕊蜜腺和第三轮雄蕊附属蜜腺的分泌节律有关。     

海南凤仙花不同海拔种群的传粉生物学

凤仙花属(Impatiens)植物具有极为广泛的多样性和类型各异的特化传粉者, 被誉为“双子叶的兰花”, 受到众多传粉生物学家的关注。本文以海南特有种海南凤仙花(Impatiens hainanensis)为研究对象, 对3个不同海拔梯度的种群进行了开花生物学特性和开花物候、花器官结构、花粉活力和柱头活性、传粉者种类和访花行为及繁育系统的比较研究。结果表明: 单花花期4.10 ± 0.46 d, 雄性期和雌性期分别约为3.15 ± 0.24 d和0.95 ± 0.36 d; 种群开花峰期在8月初, 高海拔种群的花期高峰相对滞后。低、中海拔种群花粉活力呈现先升高后下降的趋势, 以开花第2 d花粉活力最高; 高海拔种群花粉活力随开花时间推移逐渐下降; 柱头活性随开花时间的推移而增强, 高海拔种群开花各天次均较低、中海拔种群低。主要传粉昆虫为黄黑无垫蜂(Amegilla leptocoma)和绿条无垫蜂(A. zonata), 低、中海拔种群以黄黑无垫蜂为主, 高海拔种群以绿条无垫蜂为主。未观察到自动自花授粉和无融合生殖现象, 人工授粉能明显增加坐果率(75-90%), 自然坐果率在高海拔种群相对较低(40-60%), 说明存在较强的传粉者限制。海南凤仙花的保护需要同时关注其有效传粉者的保护, 促进有效传粉昆虫在不同海拔种群之间的往来, 保证种群间的花粉流与种子流, 维持海南凤仙花的种群遗传多样性与有效种群大小。     

锦带花的花粉活力、柱头可授性及传粉者的观察

在野外实地观测锦带花(Weigela florida(Bunge)A．DC．)的开花动态和访花者,在显微镜下观测柱头和花粉的形态及花粉-胚珠比,用TTC法测定锦带花花粉的活力,用联苯胺-过氧化氢法测定柱头可授性。结果表明,锦带花的花粉活力在开花时达70％以上,且能维持3d,之后明显下降。锦带花的柱头在开花3h后多数呈白色、二裂、具可授性,柱头的可授期持续2～3d。柱头与花药之间有5．3士0．9mm的空间距离,自然条件下难于自花授粉．开花当天7．00～10．00之间访花者最丰富,开花第2天访花者减少,开花第3天时访花者更少。在锦带花上记录到的访花者隶属于膜翅目、鳞翅目和双翅目,其中,熊蜂和蜜蜂是主要传粉者。     

喜树开花特性及繁育系统的研究

定位观测了喜树(Camptotheca acuminata Decne)花部构造、开花特性,用杂交指数(OCI)、花粉/胚珠比(P/O)、去雄、套袋、人工授粉等方法测定了喜树的繁育系统.结果表明:喜树花序为由4～6个头状花序组成的聚伞花序,聚伞花序上部的头状花序由两性花组成,下部的花序由雄花组成,两性花雌雄蕊发育正常,雄蕊10枚,雌蕊1枚,下位子房,一室,倒生胚珠;雄花雄蕊发育正常,雌蕊不发育.自然条件下,同一头状花序散粉后2 d,花粉活力最高;两性花在柱头外翻后3～4 d,柱头可受性最强,第6天失去可受性.喜树两性花为半同步雌雄蕊异熟类型,每个头状花序的所有小花同步开放,雄蕊先成熟,表现为雄性时期,雄蕊脱落后,雌蕊成熟,表现为雌性时期,上一级头状花序雌蕊成熟期与下一级头状花序雄蕊成熟期重合.繁育系统检测结果为部分自交和异交,需要传粉者活动才能完成授粉过程.     

宁夏枸杞的花粉呈现式样及交配系统研究北大核心CSCD

被子植物的花回馈、雌雄蕊时空分离特征和花粉呈现式样等花部特征及传粉者效率会影响雄性适合度和有性繁殖过程。宁夏枸杞(Lycium barbarum)是中国宁夏、新疆、内蒙等干旱和半干旱地区分布的特有种。该研究对新疆喀什地区宁夏枸杞自然种群的花部综合征、花粉呈现式样、花回馈、传粉者行为与交配方式的关系进行观察及统计分析,以探讨其花粉逐步呈现的适应性及其在提高雌雄繁殖过程中的意义。结果表明:(1)宁夏枸杞的单花寿命为(4.07±0.15) d,而雄性持续时间(0.07±0.01 d)比雌性持续时间短(4±0.01 d);花寿命内雌蕊长度比雄蕊长,属柱头探出式异位类型;花粉呈现式样为不完全逐步呈现。(2)花寿命不同阶段花回馈间存在显著差异。(3)意大利蜜蜂(Apis mellifera)、熊蜂(Bombus sp.)、食蚜蝇(Syrphidae sp.)是宁夏枸杞在自然居群的主要访花者,其中熊蜂和食蚜蝇是主要传粉者,但传粉者效率低,属于高移出低沉积类型。(4)宁夏枸杞的花粉胚珠比(2 448.11±448.32)及授粉实验结果均表现出兼性异交特征;自发自交及人工自花授粉花的座果率及结籽率很低,属于自交不亲和类型;自然传粉花的座果率及结籽率比人工去雄异花低,存在其较高的花粉限制(40.71%)。研究发现,宁夏枸杞花部综合征表现出雌雄异位和花粉不完全逐步呈现式样,这是避免雌雄功能及雄蕊各花药间的干扰、减少花粉同步移出及保障其雄性适合度的有效途径,但自然居群的传粉者种类限制、传粉效率低以及自交不亲性是导致宁夏枸杞花粉限制及降低雌性繁殖成功的主要原因。     

黑果枸杞两种花型的花部综合征与传粉特性

黑果枸杞(Lycium ruthenicum)是中国西北地区极端环境中分布的国家二级保护植物, 该物种在新疆南部的自然种群中出现了同型花柱类型(同位花)和柱头探出式雌雄异位类型(异位花)个体, 并且遭遇沙尘暴频繁的种群中异位花个体出现频率减少。该研究对喀什市自然种群中黑果枸杞两种不同花型植株的花部综合征和传粉特性进行比较研究, 以期探讨该物种不同花型植株在南疆早春极端环境中的花部特征的可塑性及其适应性机制。结果表明: 同位花雌雄蕊高度间无显著差异, 而异位花雌蕊高度显著高于雄蕊; 同位花花冠直径、花冠筒长、胚珠数均高于异位花, 而异位花雌雄蕊空间距离、花粉数及花粉胚珠比均比同位花高。黑果枸杞同位花个体比例(68%)高于异位花个体(32%), 种群水平及个体水平同位花花期((117.00 ± 2.25) d, (101.65 ± 1.98) d)比异位花((26.00 ± 1.00) d, (18.75 ± 1.00) d)长, 而单花水平上异位花单花寿命((4.50 ± 0.14) d)比同位花((3.13 ± 0.11) d)长。两种类型花在花早期(紫色)分泌的花蜜量均高于花后期(白色)。在紫色花阶段(花开放早期), 同位花上的主要传粉者意大利蜜蜂、熊蜂和食蚜蝇的访花频率和停留时间均高于异位花; 而白色花阶段(花开放后期)意大利蜜蜂、熊蜂在异位花上的访花频率比同位花高。在不同花色阶段, 同位花柱头花粉落置数、花粉移出率、花粉传递效率均比异位花高, 并且同位花自然坐果率及结籽率均比异位花高。在新疆南部的沙尘暴极端环境下, 同位花通过较高的自交亲和性保障繁殖, 而异交为主的异位花提高了异交率。异位花与同位花在花部综合征和花报酬上的差异, 是影响其繁殖成功的主要因素。     

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。     

光果甘草生殖生物学特性的初步研究

对光果甘草的花部特征、花粉活力、柱头可授性和花粉-胚珠比(P/O)进行了初步研究.结果表明:(1)光果甘草雄蕊异型,雌雄蕊异位,有利于避免自花授粉;(2)开花当天花粉活力在上午12:00最高,为91.8%,14:00迅速下降,16:00到达最低,16:00以后逐渐上升;(3)柱头可授性可持续4 d,以开花当天可授性最强;(4)花粉-胚珠比(P/O)为1 751.1±217.002,表明其繁育系统类型为兼性异交.     

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。     

片断化生境中濒危植物黄梅秤锤树的开花生物学、繁育系统与生殖成功的因素

 研究处于片断化生境中濒危植物的繁育系统、传粉过程及生殖成功状况, 可为评估物种的受胁迫程度及制订相应的保护和管理策略提 供重要依据。通过野外观察和人工授粉实验等方法,对片断化生境中濒危植物黄梅秤锤树(Sinojackia huangmeiensis)的开花生物学和繁育系统 进行了初步研究, 并探讨了影响其生殖成功的因素。黄梅秤锤树的总状花序上通常有两性花2～6朵, 单花花期约5～7 d, 单花花药不同步开裂 使整朵花花粉活力持续时间延长。虽然花开放过程中柱头始终高于花药, 在单花水平上可在一定程度上避免同花自交的发生, 但由于花粉活力 和柱头可授期之间有一定的重叠期, 在自然情况下, 由于外界媒介动物和风媒的作用, 不可避免地会产生同株同花和同株异花传粉。实验结果 表明, 该种不存在无融合生殖现象, 繁育类型以异交为主, 部分自交亲和且需要传粉者。主要的传粉昆虫为黑带食蚜蝇(Epistrophe balteata) 、中华蜜蜂 (Apis cerana) 和中华回条蜂(Habropoda sinensis)。黄梅秤锤树自然状态下的结实率较低(10.33%), 平均每果种子数较少(0.387 个), 这主要与传粉昆虫少、花粉传递效率低, 以及柱头上自花及同株异花花粉的落置和自交亲和性低有关。但是单株较大的开花量保证了一定 的果实和种子产量, 使居群能自然更新。部分自交亲和性可能为该物种适应生境片断化提供了一定的生殖保障及进化潜力。对该残存居群的保 护, 应着眼于加强对传粉昆虫赖以生存的自然生态系统的恢复。     

Pollination and Breeding System of Putoria calabrica (Rubiaceae), a Mediterranean Dwarf Shrub

Abstract: The pollinator spectrum and the reproductive system of Putoria calabrica, a Mediterranean chamaephyte, were studied. Its flowers are markedly protandrous and the situation of the style with respect to the stamens can be described as approach herkogamy. In the male phase, the 4 lobes of the corolla are completely extended, but in the female, they are curved back, significantly decreasing the size of the corolla. The volume and sugar content of the nectar were similar between phases, with the concentration being slightly higher in the female phase (41.9 % versus 34.5 %). Putoria calabrica is totally self-compatible; the pollen, both the plant's own and from outside, emitted pollen tubes that reached the ovules within 48 h, and the percentages of fruit set from both types of pollination were high and similar. Diurnal and nocturnal pollinators originated 48.95 % and 15.2 % of fruit set, respectively, while fruit set from free pollination was similar to the sum of the former two. Diptera (mainly Bombyliidae) and Lepidoptera were the most important pollinators, visiting a high number of flowers per plant. The role of protandry and morphological changes of the corolla of Putoria calabrica in its reproductive success are discussed.     

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 biology of Ziziphus mauritiana (Rhamnaceae)

Floral development of the synchronous dichogamous species Ziziphus mauritiana, as followed by light and scanning electron microscopy (SEM), was divided into 11 stages using a series of landmark events. Main cellular events happen synchronously in the female and the male structures, such as meiosis in micro- and macrosporocyte cells, tetrad microspore formation and appearance of the functional megaspore cell, and onset of embryo sac differentiation coinciding with mitosis in the microspores. The last stage was characterized by anthesis and continued development of the flower, beginning with anther dehiscence (male phase) and proceeding to the female phase, which was characterized by style elongation. Flowers exhibit synchronous protandrous dichogamy; anthesis takes place in the morning (group A, e.g., clone Q-29) and afternoon (group B, e.g., clone B5/4). Stigma receptivity started after the male phase and occurred synchronously and complementarily with pollen dispersal in the two clones. Pollen viability and production were similar in the two clones, but the pollen diameter of Q-29 was significantly larger than that of B5/4. This study provides the basis for understanding the biological mechanisms regulating floral development, thus expanding the prospects for Z. mauritiana breeding programs and for further molecular and genetic studies of this species.     

Pollinator limitaton in a deceptive orchid, Pogonia japonica, on a floating peat mat

Deceptive orchid species that offer no floral rewards to pollinators often experience pollinator limitation because they. The breeding system and fruit set were investigated in order to examine pollinator limitation for a population of a deceptive orchid, Pogonia japonica , on a floating peat mat in Mizorogaike Pond, western Japan. Fruit sets for outcross-pollinated, self-pollinated and pollinator-excluded flowers were 75%, 80%, and 0%, respectively. Thus, this species was self-compatible but neither autogamous nor apogamous. The fruit set for open-pollinated flowers was 22.9% and 17.4% in 1994 and 1995, suggesting that pollinator limitation did occur in the field. Larger flowers were more likely to develop into fruit than smaller ones in open pollination (36% vs 8%). Thus, flower size seems to affect fruiting of this species. No effective visits by major pollinators such as bees, which could carry pollinia, were observed during 16.5 h of observation during daytime. Instead, thrips were often found moving on gynostemium or pollinia and circumstantial evidence that suggests thrips may partly transfer pollen in pieces out of a granular pollinium of P. japonica was obtained. Pogonia japonica should come to use thrips as a pollinator to supplement low fruit set under limited flower visitation by major pollinators.     

Effects of floral sexual investment and dichogamy on floral longevity

Aims Floral longevity, the duration that a flower remains open and functional, varies greatly among species. Variation in floral longevity has been considered to be optimal strategy for resource allocation under different ecological conditions, mainly determined by the rates of pollination and cost of flower maintenance. However, it is unclear whether an intrinsic factor, floral sexual investment, constrains evolution of floral longevity. The theoretical model also predicts that dichogamy favors long-lived flowers, but empirical studies to test this prediction remain unexplored.Methods To examine the effect of floral sexual investment on floral longevity, we measured flower size together with pollen and ovule production in 37 sympatric flowering plants in a natural community. The duration of the female and male phase in 21 protandrous species and floral longevity of the other 16 adichogamous species were documented in the field.Important findings Floral longevity varied from 1 day to 15 days, while pollen number per flower varied from 643 to 710880 and ovule number per flower from 1 to 426 in the 37 species. Flower size was correlated with pollen production as well as ovule production. Floral longevity was positively related to pollen production but not to ovule production. Consistent with the prediction that dichogamy favors long-lived flowers, we found the floral longevity of protandrous species was significantly longer than that of adichogamous species. In the protandrous species, pollen production per flower was observed to be positively related to male duration, while ovule production was not related to female duration. Our analyses of variation in floral longevity and sexual investment among different species suggest that the floral sexual investment could be an intrinsic factor contributing to the selected floral longevity, particularly the male phase, and that high pollen production could potentially increase pollen removal, i.e. male productive success.     

THE POLLINATION BIOLOGY OF TILIA

An anthecological study of Tilia in the Great Plains and New England was undertaken for both native and introduced species. The floral bracts are postulated as being at least as important in pollinator attraction as they are in fruit dispersal. A characteristic sweet odor is always present, but becomes stronger at dusk. Flowers are protandrous with anthers opening for the first time late in the afternoon and releasing pollen abundantly for about 24 hr. Initial nectar production coincides with stigma receptivity which begins late in the afternoon, but on the second day a flower is open. Sixty-six species of insects in 29 families were collected. Bees and flies are the most common diurnal visitors and moths are the primary nocturnal visitors. Samples of pollen taken from the insects indicate a relatively high constancy. Experimental tests show that Tilia is not apomictic or self compatible. Anemophily plays a secondary role to entomophily in pollination. Nocturnal pollinators are slighly less effective than diurnal pollinators in effecting fruit set. Although nocturnal pollinators are favored by the syndrome of floral characteristics, it is concluded that Tilia is a generalist in regard to pollination. It is proposed that the lack of clear morphological differences between Tilia species leads to pollinator inconstancy and thus hybridization.     

Responses to selection on male-phase duration in Chamerion angustifolium

Protandry (when male function precedes female) can enhance fitness by reducing selfing and increasing pollen export and outcrossed siring success. However, responses to selection on protandry may be constrained by genetic variation and correlations among floral traits. We examined these potential constraints in protandrous Chamerion angustifolium (Onagraceae) by estimating genetic variation in male-phase duration and associated floral traits using a paternal half-sib design and selection experiment. Narrow-sense heritability of male-phase duration was estimated as 0.23 (SE +/- 0.04) and was positively correlated with floral display. The selection experiment shortened male-phase duration 0.8 SD from the parental average of 17.0 h and lengthened it by 2.0 SD. Furthermore, fixed floral longevity caused a negative association between male- and female-phase durations. These results suggest that selection on male-phase duration is not limited by genetic variation. However, changes in male-phase duration may influence pollinators through correlated changes in floral display and reduced opportunities for pollen receipt during female phase.     

Flower biology of the cactus Coryphantha elephantidens in the tropical dry forest of Central Mexico

Floral traits and sexual systems in angiosperms are strategies that enhance outcrossing within hermaphrodite flowers and among individuals in a population. Sexual systems with unisexual flowers have also evolved among angiosperms, resulting in sex specialization. Furthermore, the interaction of floral traits and floral visitors determines successful plant reproduction. Globose cacti are bee pollinated, and variation in the diversity of their pollinator assemblages is strongly associated with floral phenotype. Our objective was to describe the floral biology of the cactus Coryphantha elephantidens and to determine its relationship with pollinators. Floral traits were studied by direct observations in live and fixed flowers. The breeding system was determined using two estimators based on floral morphology: pollen grains to ovules per flower (P/O) ratio and outcrossing index. Pollination treatments were conducted to determine the mating system. Floral visitors were recorded using direct observation. Flowers of C. elephantidens are variable in color, protandric, herkogamous and nectarless. Estimators of the breeding system indicated xenogamy, which is consistent with the obligate outcrossing revealed by the pollination experiment. Thirty-seven percent of the plants have female flowers that do not produce pollen, making this population functionally gynodioecious. Both fruit and seed set were high compared to other globose cacti. Pollinators included eight species of native bees, a more diverse pollinator assemblage than other globose cacti. Given the high pressure on pollen due to functional gynodioecy, nectarless flowers, an outcrossing mating system, and the necessity of pollinators to set seeds, we concluded that native bees are highly efficient pollinators that play a crucial role in the sexual reproduction of C. elephantidens.     

Gender specialization and factors affecting fruit set of the wind-pollinated heterodichogamous Juglans regia

Gender expression, flowering phenology, reproductive performance and factors affecting fruit set (i.e., flowering synchrony, size and distance to the nearest pollen donor) were investigated in a cultivated population of a wind-pollinated self-compatible heterodichogamous Juglans regia (Juglandaceae). Four flowering morphs, (i.e., protandrous, protogynous, male and female) were observed. The sexual functions of the protandrous and protogynous morphs were almost synchronous; however, they were not reciprocal, and the separation of male and female flowering within most monoecious individuals was not complete. Thus, within-morph mating and geitonogamous pollination may be common. The ratio of protandrous versus protogynous morphs was biased towards the protandrous morph, but the fruit set did not differ between the morphs, suggesting that the fruit set of the protandrous morph could be partly compensated by within-morph pollination. The ratio of the female flower number or fruit number to the total male catkin length was higher in the protogynous morph than in the protandrous morph and did not vary with plant size, suggesting that gender variation was not size dependent and that the sexual function of protandrous morphs was more male biased. Fruit set depended on plant size only for protandrous morphs. The fruit set of individual plants decreased with increasing distance to the nearest pollen donor regardless of morph, possibly because of pollen limitation. The fruit set of individual plants increased with flowering synchrony, indicating that flowering synchrony could affect reproductive success.