Pollination, Pollen Viability and Pistil Receptivity in Cucurbita pepo

Cucurbita pepo carries male and female flowers on the same plant,and is pollinated by nectar-collecting bees. The nectaries aredimorphic in the two sexes and pollen is loaded and unloadedas the bees gain access to the nectar. Both types of flowerare open for only 6 h (from 0600 h to 1200 h); male flowersopen and close half an hour earlier than female flowers. Thelatter produce more nectar and are visited more often by thebees than the male flowers. Pollen viability determined by fluoresceindiacetate (fluorochromatic reaction) decreases by 20% duringanthesis and more rapidly after the flower closes. This decreaseis due to dehydration of the grain, especially around the porewhere the intine is exposed. An unusual feature of this speciesis that the grains do not dehydrate before anther dehiscence.Female receptivity has two aspects, that of the stigma lasting4 d, and that of the ovules lasting 2 d. The receptivity ofthe two sexes and the short period of anthesis are discussedin the light of the reproductive ecology of the species.Copyright1993, 1999 Academic Press Entomophilous pollination, anthesis, nectars, pollen viability, female receptivity, bees, pollinator efficiency, courgette Cucurbita pepo, Cucurbitaceae     

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.     

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。     

Dichogamy,gender variation and bet-hedging inPseudowintera colorata

Summary Temporal patterns of variability in the longevity of the male and female phases of individual flowers and in the gender expression of plants of a dichogamous New Zealand tree,Pseudowintera colorata (Winteraceae), were documented in field studies. Two measures for the duration of phases in a dichogamous flower are distinguished; the nominal phases based on morphological features of the flower, and the effective phases reflecting the duration of their functions. Flower and phase longevity and phenotypic gender varied considerably throughout the season and among individuals. Temporal variability in phenotypic gender was loosely synchronized among the 12 plants sampled. Three effects of an environmental factor (temperature) were noted. First, increased temperatures shortened the duration of the female phase but had no effect on the duration of the male phase. Second, pollination frequency was positively correlated with temperature. These results indirectly suggest that increased pollination may shorten the duration of the female phase. Third, average population maleness, measured as the proportion of open flowers in the population on a given day which were in the male phase, was positively correlated with temperature. It is postulated that temperature indirectly influences temporal patterns of gender expression in the population through its differential effects on the longevity of the male and female phases in individual flowers. A theoretical model of bet-hedging shows that, if the direction of an environmental effect on the proportions of the sexual phases is irreversible, selection favours asynchronous dichogamy and reduces the temporal variability as much as possible. If the direction of the response is reversible, heterodichogamy is favoured.     

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。     

峨眉拟单性木兰的开花生物学特性与繁育系统

为阐明峨眉拟单性木兰(Parakmeria omeiensis)在自然条件结实率低的原因和确定最佳人工授粉时期,该文通过观察峨眉拟单性木兰的开花动态,采用杂交指数估算、花粉胚珠比、花粉活力及柱头活性检测、人工授粉试验等方法对其繁育系统进行了研究。结果表明:(1)植物园保育的峨眉拟单性木兰花期在4月底到5月中下旬,持续17～23 d,雄株始花期比雌株早3～4 d,但两者花期可遇。(2)两性花经检测雄蕊败育,实为功能上的雌性,部分雄株个体的雄花上残留1～2个心皮,其性别分化是通过雌、雄蕊选择性败育形成的,为隐性雌雄异株(cryptic dioecy)。(3)雄花、两性花开放经历佛焰苞开裂、花被片开裂、展开、闭合、二次开放、凋落6个阶段,历时4 d。(4)雄花初次展开时花粉活力最高,达92.8%,开花2 d后活力显著下降;两性花柱头在花被片展开期可授性最强,盛开后柱头部分可授。(5)杂交指数为5,P/O比为2.14×10~4。(6)套袋试验表明,峨眉拟单性木兰不能进行自花传粉,人工异花授粉的结实率和出种数显著高于自然授粉,且不存在无融合生殖。这说明峨眉拟单性木兰繁育系统为专性异交,传粉过程需要传粉媒介,自然条件下结实率低,主要是受传粉昆虫和柱头可授期短的限制。     

The dynamics of pollen removal and deposition,and its effects on sexual phases in a protandrous plant: Glechoma longituba (Lamiaceae)

The duration of sexual phases in dichogamous plants are affected by many factors. Using both experimental and observational studies, we investigated natural patterns of pollen removal and deposition, visiting frequency of pollinators, patterns of nectar secretion, and effects of pollen removal and stigmatic pollen deposition on the duration of sexual phases in a protandrous plant, Glechoma longituba. We found that visiting frequency of pollinators correlated with the nectar secretion pattern. The nectar volume during the male phase was higher than during the female phase. In the morning, the main pollinator, the bee Anthophora plumipes, mainly foraged for nectar and showed no preference for flowers in male or female phase, despite male phase flowers producing higher amounts of nectar. However, in the afternoon, they changed their behavior and foraged mainly for pollen, and then showed a preference for flowers in male phase. Furthermore, the rates of pollen removal and stigmatic pollen deposition can affect the starting time and the duration of the female phase. When pollen removal and pollination rates are low due to scarcity of pollinator services, the sexual phase can be prolonged, leading to an overlap, and thereby enhance the chance for sexual reproduction through pollinator‐facilitated self‐pollination. We consider the variation of sexual phases in Glechoma longituba an adaptive mechanism prepared for both cross‐pollination enhancement and reproductive assurance depending on the available pollination services.     

Pollen:ovule ratio and its relationship with other floral traits in Papilionoideae (Leguminosae): an evaluation with Argentine species

The pollen:ovule ratio (P/O) has traditionally been used as a rough estimator of plant breeding systems. It has been shown that plant breeding systems are associated with particular floral traits. In this study, we determined the P/O in 21 Leguminosae species from Argentina and explored relationships between P/O and taxonomic position, flower size, floral rewards, pollen presentation and pollination mechanisms. According to the results, 15 out of the 21 species classified were obligate xenogamous, although some of them have been recorded as facultative xenogamous in previous studies. There was a significant effect of taxonomic position (genus), reward type and pollination mechanism on P/O. Species offering only nectar as a floral reward (which were species with a brush mechanism) had a significantly lower P/O than species offering pollen or pollen and nectar. Species with the brush pollination mechanism had the lowest P/O, while species with valvular and pump mechanism had the highest P/O. However, pollen presentation (primary and secondary) and flower size did not have a significant effect on P/O. Our results demonstrate that P/O variability is determined by taxonomic position and pollination mechanism in this plant group.     

Floral reward presentation favored the expression of male function in the pollen‐only flower Melastoma malabathricum

Abstract Animal‐pollinated plant species modulate the presentation of pollinator rewards to maximize reproductive success. In plants providing pollen as the only reward for pollinators, it is usually difficult to unravel the dual roles of reward presentation and the realization of male and female functions (pollen removal and deposition). Exploiting the two types of anther in the androecia of Melastoma malabathricum L., we examined whether the removal of pollen for reward is regulated primarily to favor male function or female function. Pollen removal by carpenter bees from the feeding and pollination anthers, as well as pollen deposition on the stigmas, were quantified during anthesis of M. malabathricum. There was no significant difference in pollen removal rates from the feeding and pollination anthers of M. malabathricum between the onset of anthesis and flower wilting. The stigmatic pollen loads exceeded the ovule number after three sonication bouts, and female function was satisfied earlier than male function. The results support the hypothesis that the presentation of pollination reward in this species is regulated primarily to favor the expression of male function, rather than female function, in agreement with the pollen‐donation hypothesis. A cooperative relationship between the feeding and pollination anthers was demonstrated in heterantherous flowers, which optimizes the balance in investments between pollinator rewards and “functional pollen” for gene transfer.     

Reproductive barriers in Annona cherimola (Mill.) outside of its native area

Cherimoya (Annona cherimola) is a subtropical tree crop of Andean origin whose fruit set results extremely low in farming areas outside of its natural occurrence. The lack of efficient pollinators and dichogamy are often argued to be the main constraints resulting in this low reproductive success. Herein, we describe the reproductive barriers exhibited by this crop and whether wind and insects play a role in cherimoya pollination in Spain, the main region of cultivation. A.?cherimola exhibits marked protogynous dichogamy with large differences in the duration of female (around 28?h) and male (<8?h) phases. Stigma receptivity and pollen release do not fully coincide with the morphological changes of the petals defining the female and male phases. Synchronization of sexual phases among different flowers from different trees of the same genotype was high during the whole blooming season. Effective herkogamy of approach type also limits pollen deposition within the same flower. Wind does not play any role in cherimoya pollination. Insect visitors to cherimoya flowers in Spain were found to be inefficient in transferring pollen grains. Cherimoya flowers do not reject self-pollen to achieve fertilization. A.?cherimola shows preferential allogamy based on efficient dichogamy reinforced by elevated synchrony among flowers in their sexual phases. Herkogamy hampers autogamy, although pollen deposition by gravity in cherimoya pendulous flowers explains the reduced reproductive success observed in isolated flowers.     

Asynchronous development of stigmatic receptivity in the pear (Pyrus communis; Rosaceae) flower

While stigma anatomy is well documented for a good number of species, little information is available on the acquisition and cessation of stigmatic receptivity. The aim of this work is to characterize the development of stigma receptivity, from anthesis to stigma degeneration, in the pentacarpellar pear (Pyrus communis) flower. Stigma development and stigmatic receptivity were monitored over two consecutive years, as the capacity of the stigmas to offer support for pollen germination and pollen tube growth. In an experiment where hand pollinations were delayed for specified times after anthesis, three different stigmatic developmental stages could be observed: (1) immature stigmas, which allow pollen adhesion but not hydration; (2) receptive stigmas, which allow proper pollen hydration and germination; and (3) degenerated stigmas, in which pollen hydrates and germinates properly, but pollen tube growth is impaired soon after germination. This developmental characterization showed that stigmas in different developmental stages coexist within a flower and that the acquisition and cessation of stigmatic receptivity by each carpel occur in a sequential manner. In this way, while the duration of stigmatic receptivity for each carpel is rather short, the flower has an expanded receptive period. This asynchronous period of receptivity for the different stigmas of a single flower is discussed as a strategy that could serve to maximize pollination resources under unreliable pollination conditions.     

北京地区蒿柳的开花物候期及单花序开放过程中花粉活力和柱头可授性变化

Flowering phenology and flowering dynamics of female and male inflorescences of Salix viminalis Linn. in Beijing were observed in 2015 and 2016, and changes in pollen activity and stigma receptivity were also investigated. The results show that early flowering stage of S. viminalis in Beijing is happened in early?March, flowering period is 11-18 d, and flowering pattern is the first opening type of male flower. Flowering period of single inflorescence in male and female plants is 10-15 d and 8-10 d, respectively, which can be divided into five stages:emerging color stage, initial flowering stage, half flowering stage, full flowering stage and withered stage. Flowering date of male plant is earlier about 3-4 d than that of female plant. However, the difference of full flowering stage between male and female plants is only 1 d. Pollen germination rate and stigma receptivity show a trend of firstly increasing then decreasing during blossom process. Pollen germination rate reaches the highest of 72% at full flowering stage, and stigma receptivity is more than 90% at half and full flowering stages. Therefore, it is suggested that pollen collection and pollination in mid?March can improve success rate of cross breeding of S. viminalis.     

Influence of pollen limitation and inbreeding depression in the maintenance of incomplete dichogamy in Salvia elegans

The widespread presence of incomplete dichogamy (i.e., partial separation in time between male and female phases) in flowering plants is a long‐standing question in floral evolution. In this study, we proposed four scenarios in which depending on the particular combination of pollen limitation and inbreeding depression, the presence of complete dichogamy, incomplete dichogamy, or adichogamy may be favored. Moreover, we evaluated the role of pollen limitation and inbreeding depression in a natural population of Salvia elegans to test the validity of our predicted scenarios. Our results indicate that S. elegans is partially protandrous as pollen viability and stigma receptivity overlap in the last days of life of the flower. Furthermore, through pollination treatments, we found no evidence of pollen limitation or inbreeding depression in any of the evaluated fitness components. As expected by one of the proposed scenarios, incomplete dichogamy seems to be favored in plants with absence of inbreeding depression and pollen limitation as a way to diminish interference between male and female functions.     

Floral longevity in Campanula americana (Campanulaceae): a comparison of morphological and functional gender phases

Plastic responses to pollination and/or pollen removal may shift a flower's realized longevity closer to an optimal longevity that maximizes reproductive output per unit resource input. In particular, conditional responses to pollen removal and pollen deposition are expected in flowers of protandrous species in which the lengths of the male and female phases may be adjusted independently. We investigated plasticity in floral longevity in Campanula americana, a protandrous, insect-pollinated herb. In greenhouse studies, we found that the longevity of the morphological male phase was shortened by pollen removal and that the longevity of the morphological female phase was shortened by pollen deposition. In a natural population, male and female sexual functions saturated within a few hours of morphological gender phase onset. In contrast to theory, morphological gender phases did not terminate immediately upon saturation of sexual function. These findings are discussed in the context of current floral longevity theory.     

钝叶柃不同性别花的花部形态与传粉特征比较

据有关资料记载,柃属(Eurya Thunb.)植物属于雌雄异株植物,目前仅在柃木(Eurya japonica Thunb.)中有过两性花的报道。近年的调查发现,钝叶柃(Eurya obtusifolia H.T.Chang)也有性别变异,存在两性花。在对其性别特征及功能研究的基础上,进一步跟踪了钝叶柃的开花和传粉过程,对不同性别花的花部形态和传粉特征进行了比较分析。在重庆市北碚区选取了4个样地,测定了花部形态、花朵朝向、花粉活力、柱头可授性、花粉对水的耐受性、单花花粉量和单花泌蜜体积等形态和传粉相关指标,并对其开花动态、传粉昆虫种类和昆虫传粉行为进行了仔细观察,还进行了套袋试验。结果发现钝叶柃不同性别花的花部形态和传粉特征既有一些共同点,又存在着明显差异,性别变异株则呈现出雌雄植株间的过渡特征。钝叶柃是一种花期短,花小而多,开花同步性高的植物,雌花和雄花色味相同。该植物的主要传粉方式为虫媒,主要传粉者为两种蜜蜂科昆虫,但风媒也在其传粉中起着一定作用。该植物不同性别花的主要差异有:1)雄花呈灯笼状,花瓣不反卷,雌花辐射状,花瓣反卷;2)雄花较雌花大;3)雄花倾向于垂直朝下,雌花倾向于斜向下朝向枝条末端;4)雌花寿命较雄花长。钝叶柃在开花和传粉上的许多特征都在一定程度上体现了其对雌雄异株性系统的适应。相应地,传粉者在不同性别植株上的行为也存在一定差异。钝叶柃不同性别花在形态上的差异是其周围各种生物因素及非生物因素共同作用的结果,其中传粉昆虫和雨水的选择可能在这些差异的塑造中起着重要作用。     

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...     

Partial preference of insects for the male flowers of an annual herb

Summary The flowers of the annual herb Impatiens capensis have distinct male and female phases. The male phase lasts four times as long as the female phase, and male flowers contain about 50% more nectar than female flowers. This suggests that the bulk of allocation to the flower is designed to ensure the dispersal of pollen rather than the fertilization of ovules. Honeybees, wasps and bumble bees all land on male flowers more often than would be expected by chance, and, having landed, wasps and bumble bees are more likely to enter a male flower than a female flower. The frequency of male flowers in the diet therefore exceeds their frequency in the population. This preference, although strong and consistent, is only partial, since some female flowers are included in the diet. We propose two hypotheses to account for the observed partial preference, the first based on competition between bees for flowers, and the second asserting that the bees detect nectar levels directly without using floral gender as a cue. The results of an experiment in which the most obvious gender cue, the androecium, was removed are consistent with the second hypothesis.     

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。     

Female reproductive success in gynodioecious Thymus vulgaris: pollen versus nutrient limitation and pollinator foraging behaviour

Gynodioecy is a dimorphic breeding system in which female individuals coexist with hermaphroditic individuals in the same population. Females only contribute to the next generation via ovules, and many studies have shown that they are usually less attractive than hermaphrodites to pollinators. Several mechanisms have been proposed to explain how females manage to persist in populations despite these disadvantages. The ‘resource reallocation hypothesis’ (RRH) states that females channel resources not invested in pollen production and floral advertisement towards the production of more and/or larger seeds. We investigated pollination patterns and tested the RRH in a population of Thymus vulgaris. We measured flower display, flower size, nectar production, visitation rates, pollinator constancy and flower lifespan in the two morphs. In addition, we measured experimentally the effects of pollen and resource addition on female reproductive success (fruit set, seed set, seed weight) of the two morphs. Despite lower investment in floral advertisement, female individuals were no less attractive to pollinators than hermaphrodites on a per flower basis. Other measures of pollinator behaviour (number of flowers visited per plant, morph preference and morph constancy) also showed that pollinators did not discriminate against female flowers. In addition, stigma receptivity was longer in female flowers. Accordingly, and contrary to most studies on gynodioecious species, reproductive success of females was not pollen limited. Instead, seed production was pollen limited in hermaphrodites, suggesting low levels of cross‐pollination in hermaphrodites. Seed production was resource limited in hermaphrodites, but not in females, thus providing support for the RRH. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 395–408.     

Relationship between floral longevity and sex allocation among flowers within inflorescences in Aquilegia buergeriana var. oxysepala (Ranunculaceae)

Understanding the fitness of plants with inflorescences requires examining variation in sex allocation among flowers within inflorescences. We examined whether differences in the duration of the male and female phases of flowering lead to variation in sex allocation and reproductive success among flowers within inflorescences. In 2002 and 2003, we quantified floral longevity, floral sex allocation, and reproductive success between the first and the second flowers within inflorescences in a protandrous species, Aquilegia buergeriana var. oxysepala. Floral longevity was greater in the first flowers than in the second ones in both years. The male phase lasted longer, and the initial number of pollen grains and the number of pollen grains removed were greater in the first flowers than in the second ones in both years. Within first flowers, the number of pollen grains removed was greater in flowers that had longer male phases, thus duration of the male phase may positively affect male reproductive success in the first flowers. The female phase lasted longer and the number of ovules was greater in the first flowers than in the second only in 2002. However, seed production per flower and female phase duration in both years were not significantly related. The variation in the number of pollen grains among flowers in this species may be caused by the variation in male phase duration.