THE INFLUENCE OF THE MATING SYSTEM ON SEED SIZE VARIATION IN CRINUM ERUBESCENS (AMARYLLIDACEAE)

We present evidence that extreme seed size variation in fruits of Crinum erubescens (range: 0.1 to 66.5 g per seed) occurs when mating pairs are inbred, either from selfing or biparental inbreeding. Several relatively uniform seeds of intermediate size are produced when pollen from several pollen donors is applied simultaneously to a flower. Selfed fruits and some fruits pollinated with a single pollen donor produce both large and small seeds, although selfed fruits produce fewer seeds than outcrossed fruit. These results are contrary to the hypothesis that variation in seed size is attributable to either pollen competition or differential allocation of maternal resource to seeds of different genotypes.     

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。     

毛翠雀花花序内的性分配和繁殖成功

两性花植物花序内不同位置的性分配和繁殖成功一般存在差异,通常认为资源竞争、结构效应和交配环境是形成这种差异的主要原因。为了研究雄性和雌性繁殖资源在花序内不同位置间的最优分配问题,该文以青藏高原高寒草甸典型高山植物毛翠雀花为材料,通过比较花序内不同位置的花部特征和种子性状,对其花序内的性分配模式和雌性繁殖成功进行研究,并通过观察传粉者运动特点以及人工去花和补授花粉实验,探讨花序内资源竞争和交配环境对繁殖资源分配的影响。结果表明:(1)不同位置间的雄蕊数、雄蕊鲜重/雌蕊鲜重、花粉数及花粉胚珠比从花序基部到上部显著增加,而雌蕊鲜重和胚珠数逐渐减少,表现出上部花偏雄的性分配;上部花的结籽率显著低于基部花和中部花,不同位置间的发育种子数/果实和发育种子重/果实随着花位置的升高而显著降低,说明基部花具有更佳的雌性繁殖成效。(2)去花处理后,剩余果实的单个种子重/果实显著增加,但发育种子数/果实没有显著增加;而给上部花人工补授异花花粉后,位置间结籽率的差异消失,说明传粉限制而非资源竞争导致了花序内位置依赖的种子生产模式。(3)毛翠雀花雄性先熟的开花特征,以及传粉者苏氏熊蜂从花序基部到上部的定向访花行为,导致了花序内交配环境的变化。综上结果表明,毛翠雀花花序内的性分配和繁殖成功差异是对交配环境适应的结果,对其在高山环境中实现雌雄适合度最优化具有重要意义。     

Fruit abortion in the Chihuahuan-Desert endemic cactus <Emphasis Type="Italic">Opuntia microdasys</Emphasis>

Resource and pollen limitation, as well as pollen/ovule incompatibility, have been proposed as causes to explain fruit abortion. To assess whether abortion in Opuntia microdasys was due to resource and/or pollen limitation and could therefore be reversed fruit set and seed set were studied using controlled pollination experiments on 60 plants that had been randomly assigned a combination of watering and fertilization treatments. On the other hand, to test whether fruit abortion was irreversible, due to pollen/ovule incompatibility, we examined the reproductive biology of the species. This included observations on floral phenology, nectar production, flower visitors, numbers of pollen grains and ovules, and self-pollination experiments. Results showed that O. microdasys is a fully self-incompatible species and its floral biology and the activity of the main pollinator allow constant deposition of incompatible pollen onto stigmas, which may contribute to fruit abortion. Reproductive success was limited by nutrients and pollen, but the fruit set increased only by 58%, compared to 47% of the control, after the experimental addition of pollen, nutrients and water. The magnitude of pollen and resource limitation suggests that similar levels of abortion will be present in good as well as in bad years. Selfing as well as incompatibility between ramets from the same clone and between closely related plants seem plausible candidates to explain the large proportion of fruit abortion, and experimental cross pollination between genotypes identified through molecular markers are necessary to fully understand the considerable abortion rate that remains unexplained after pollen and resource addition. Interestingly, the possible reason why the abortion of energetically expensive fruits has not been eliminated by natural selection is that the aborted fruits are propagules able to root and produce new plants with the same genotype of the mother. Abortion would have a dramatic effect on cross-fertilized genotypes because they result in zero fitness, but it would have a positive effect on the fitness of the maternal genotype because a clonal offspring is produced. Evidently, the exact fitness consequences to the maternal plant will depend on the differences in survival and reproduction of these different offspring types.     

FERTILIZATION DYNAMICS AND PARENTAL EFFECTS UPON FRUIT DEVELOPMENT IN RAPHANUS RAPHANISTRUM: CONSEQUENCES FOR SEED SIZE VARIATION

Seed weight varies significantly within and among fruits of wild radish (Raphanus raphanistrum). To determine sources of this variation, we studied fertilization and seed development following controlled pollinations. Within fruits, central ovules were fertilized prior to distal ovules and attained greater seed size. Ninety-seven percent of the variation in mean seed wt per fruit was explained by an analysis of variance incorporating parental effects, pollination date, and the number of seeds per fruit. We document strong maternal effects on the number of ovules per ovary, the number of fertilized ovules per ovary, the number of seeds per fruit, and mean individual seed wt per fruit. Across females, pollen donor had a slight but significant effect on seed wt; no paternal effects on fertilization rate, zygote number, or seed number per fruit were detected. Within females, with one exception, pollen donor had no significant effect on these components of seed development. Stronger maternal main effects may be due to donor x recipient interactions, cytoplasmic factors, the genetic inequity within triploid endosperm, and/or strict maternal control over resource allocation. The large maternal effects relative to paternal effects may limit the rate at which natural selection acts on paternal traits expressed prior to seed maturation.     

Maternal provisioning, sibling rivalry and seed mass variability in the dioecious shrub Rhamnus alpinus

In this paper, we study the sources of variability in seed provisioning in Rhamnus alpinus, a dioecious shrub producing three-seeded fleshy fruits (range 1–4). Discarding position effects in access to resources as a source of variation, since pyrenes are radially oriented inside the fruit, we conducted a pollination experiment, in which different pollen donors were used to make single and mixed pollinations on different female plants. We found strong maternal effects on brood size and seed mass. Although we found no effects of pollen load or pollen source on fruit set, brood size or seed mass, all variables showed enhanced values after multiple pollination treatments. Furthermore, we found an increase of intra-brood variance in seed mass after multiple pollination, suggesting that multiple siring really occurs when multiple donors were involved in pollination, and that sibling rivalry could be a mechanism implicated in seed provisioning, discarding maternal choice as the alternative scenario. Our results on resource provisioning fits with a hierarchical model, since disparities in shares increased down the dominance rank, being the middle sized seed the least sensitive to resource partitioning, and the disparities were reduced when more resources were available in terms of fruit mass. Maternal effects, number of pollen donors, and sibling competition are therefore accounting for seed mass variability in R. alpinus.     

Effects of inbreeding and pollen donor provenance and diversity on offspring performance under environmental stress in the rare plant Cochlearia bavarica

Habitat degradation and loss can reduce size and genetic variability of natural populations, increasing individual homozygosity and average relatedness between individuals. While the resulting inbreeding depression may be reduced by natural selection under prevailing environmental conditions, it may increase again under environmental stress. To investigate the effect of environmental stress on offspring performance and the expression of inbreeding depression, we hand-pollinated maternal plants in small (< 100, n=5) and large populations (> 400 flowering plants, n=5) of the rare plant Cochlearia bavarica (Brassicaceae) and raised the offspring under experimentally manipulated water and light regimes (normal or reduced supply). In addition to considering natural variation in inbreeding levels due to population size, we manipulated pollen donor provenance and diversity. Maternal plants were pollinated with nine donors from a different population or with one or nine donors from the same population. One further inflorescence of each maternal plant was exposed to free pollination. Offspring growth and survival were monitored over 300 days. Offspring performance varied significantly among populations and maternal plants. Environmental stress interacted significantly with these factors. However, there was no general indication that offspring from small populations were more negatively affected. In seven out of 10 populations, offspring derived from between-population pollination performed better than offspring derived from within-population pollination. Also, in five out of 10 populations, average offspring size was higher after within-population pollination with nine than after pollination with one pollen donor. These results suggest low genetic diversity within C. bavarica populations, both smaller and larger ones. Interactions between environmental stress and pollination treatment indicated that using pollen donors from outside a population or increasing the number of pollen donors can reduce inbreeding depression, but that this beneficial effect is impaired under stressful conditions.     

Seed germination patterns of Salvia mellifera in fire-prone environments

Summary I examined the effects of pollen loads containing pollen from one, three and five donors on fruit production and fruit quality in Campsis radicans. Number of pollen donors had no significant effect on % fruit production, seed number, seed weight or seed germination. In singledonor pollinations the identity of the donor did have a strong effect on the above parameters. Furthermore, the best single donor sired fruits with more seeds and heavier seeds than any mixture containing this donor. This pattern indicates interference of pollens or preemption of some ovules by the inferior pollen. In Campsis, therefore, the number of pollen donors contributing to a pollen load is less important than the identity of these donors in determining fruit production and fruit quality. Seeds from fruits resulting from mixed pollination were slightly more variable than seeds from fruits resulting from single-donor pollinations.     

Floral trait variation in fourCyclamen (Primulaceae) species

The purpose of this study is to examine floral trait variation between fourCyclamen species that show variation in their ability to reproduce in the absence of pollinators and in levels of inbreeding. Pollen and ovule production, pollen/ovule ratio, pollen volume, petal length and width, diameter of the corolla mouth, pistil and stamen length, and stigma-anther separation varied significantly between the four study species. Flower, pollen and ovule production, pollen volume and corolla size were generally highest inC. hederifolium, the species with the lowest level of inbreeding (mean F_is = 0.329), intermediate in both species with relatively high levels of pollinator-mediated inbreeding,C. repandum (mean F_is = 0.658) andC. creticum (mean F_is = 0.748), and lowest in the highly inbredC. balearicum (mean F_is = 0.930). The two species with the most different inbreeding coefficients,C. hederifolium andC. balearicum, had lower pollen-ovule ratios and shorter longevities of stigma receptivity and pollen viability thanC. creticum andC. repandum. These patterns of variation in floral traits are discussed in relation to the relatedness, pollination ecology and levels of inbreeding of the four species.     

Effects of pre-dispersal selection on offspring growth and survival in Erythronium grandiflorum

Effects of pre-seed-dispersal processes on offspring vigor were examined in Erythronium grandiflorum using manipulations of the number of pollen donors contributing to the pollen pool and comparisons of means and variances in offspring growth measurements. There were no effects of the number of donors on measures of pollen-tube growth, ovule abortion, seed set, mean seed weight, or seedling germination. Seeds from pollinations with only one donor produced corms that averaged 5% lighter after one season of growth and had lower overall survival after three years compared to corms from pollinations with either three or ten donors. Patterns of within- and among-family variance estimates for the different treatments were consistent with the hypothesis that less-vigorous offspring were eliminated prior to seed dispersal in the multiple-donor treatments. The difference in the growth of offspring from different treatments was apparently not due to pollen competition because pre-zygotic attrition of pollen tubes led to incomplete fertilization of ovules. Results from this study suggest that post-fertilization abortion of less-vigorous progeny, perhaps as a consequence of early-acting inbreeding depression, is responsible for the increase in the average vigor of offspring from multiply-sired fruits.     

两种互逆资源梯度影响下刺叶锦鸡儿的种子选择性败育格局

种子的选择性败育在被子植物中普遍存在,开展结籽格局及其影响因素的研究有助于深入了解种子的形成机制及多样化的生殖对策。以刺叶锦鸡儿Caragana acanthophylla为材料,通过传粉过程、胚珠发育动态和资源分配状况的研究,以探讨种子的选择性败育格局及相关影响因素。结果显示：(1)刺叶锦鸡儿具高度自交不亲和性,为泛化的传粉系统,蜂类是主要的传粉者。自然状态的结实率为(86.00±4.96)%,不存在传粉限制,但受精具明显的时间效应。(2)刺叶锦鸡儿单花期4—5d,每朵花有(14.00±0.14)粒胚珠。胚珠的发育从荚果顶部开始,其中,开花后第3天荚果顶部胚珠开始膨大;第11天绝大多数胚珠出现了膨大,此时,在荚果基部的胚珠开始败育,随后荚果顶端受精后的胚珠也出现败育,最终仅荚果中部形成2—3粒种子。其结籽格局为成熟荚果中部胚珠,而败育荚果顶部和基部胚珠的选择性败育类型。(3)受微地形影响,居群内开花植株具斑块分布,对较少开花植株通过添加水肥进行资源调控后,结籽率显著提高,说明在种子形成过程中存在资源限制。综上所述,受精顺序和资源限制两个互逆的资源梯度决定了刺叶锦鸡儿的结籽格局。其中,...     

EFFECT OF CROSSING DISTANCE AND MALE PARENT ON IN VIVO POLLEN TUBE GROWTH IN CHAMAECRISTA FASCICULATA

Gametophytic competition among pollen grains has been proposed as an important mechanism of sexual selection in plants. The purpose of this paper is to examine the contribution of pollen source on in vivo pollen tube growth in Chamaecrista fasciculata. We addressed two questions: 1) Is pollen tube growth affected by the genetic relatedness between the pollen source and the pollen recipient? 2) Is there significant phenotypic variation among pollen donors for pollen tube growth? We compared pollen tube growth by measuring the number of pollen tubes which germinated, reached quarter length of style, and reached the ovary resulting from self- and outcross-pollinations. The outcross pollinations included three interplant distance classes: near (within genetic neighborhood, ca. 1 m), far (between far neighborhoods and within subpopulation, ca. 20 m), and distant (between neighborhoods and adjacent subpopulations, ca. 50–100 m). Our results show that pollen tube growth was not affected by genetic relatedness, by differences between self and outcross, nor by differences due to phenotypic variation among pollen donors. In contrast, maternal environment had a strong impact on pollen tube growth. These results suggest a lack of gametophytic competition and indicate little opportunity for sexual selection on pollen tube growth in C. fasciculata.     

EFFECTS OF POLLINATION INTENSITY IN CAMPSIS RADICANS

I examined effects of pollination intensity on fruit, seed and seedling characteristics in Campsis radicans, and joint effects of pollen donor and pollination intensity on fruit production. Large pollen loads were more likely to initiate fruit production than small pollen loads, and the former fruits contained more seeds and a greater total seed mass. No further increases in seed number or mass occurred for pollen loads above 4,000 grains. The weight of individual seeds was unaffected by pollen load. Effects of pollen donor were generally larger than effects of pollen load, and fruit production from small loads of pollen from one donor were sometimes equal to fruit production from larger pollen loads from another donor. The ratio of pollen grains deposited to resultant seeds increased with pollen load, and several explanations are proposed. Seeds from heavy pollinations emerged better than seeds from light pollinations, but did not differ in speed of germination or in the performance of seedlings up to 126 days. The emergence differences are probably due to differing intensities of pollen tube competition.     

Role of antioxidant defense system and biochemical adaptation on stress tolerance of high mountain and steppe plants

Plants growing in small fragmented populations under stressful environmental conditions may have reduced sexual reproduction. This can cause low gene flow between populations and eventually extinction. Here we report on a pollination experiment with Sorbus torminalis, a rare fleshy-fruited tree with a submediterranean distribution in Europe. At the northern limit of its range in SE-Denmark two relatively small and isolated populations were studied for effects of seven pollination treatments on fruit production and on the timing of fruit abortion. There was evidence that lack of pollination and spontaneous self-pollination caused particularly high fruit abortion, which indicates that apomixis is unlikely and spontaneous self-pollination not efficient. Fruit abortion was delayed after hand pollination, which suggests limitation by pollen quantity. Self-pollination caused earlier abortion than experimental cross-pollination within or between populations indicating inbreeding depression. There was no evidence for outbreeding depression as measured by fruit abortion. We conclude that generative reproduction of S. torminalis is reduced on its northern distribution limit and that it might be negatively affected by pollen limitation and inbreeding effects, which have not been compensated for by increased self-compatibility or apomixis.     

Self‐incompatibility and post‐fertilization maternal regulation cause low fecundity in Aegle marmelos (Rutaceae)

Outbreeding confers an evolutionary advantage, and flowering plants have evolved a variety of contrivances for its maximization. However, neither fruit set nor seed set is realized to its fullest potential for a variety of reasons. The causes of low flower to fruit and seed to ovule ratios were investigated in naturally occurring bael trees (Aegle marmelos) at two sites for three seasons. The study established that the mass effect of synchronized flowering attracted a variety of insect pollinators to the generalist flowers; Apis dorsata was the most efficient pollinator. The seed to ovule ratio was low despite high natural pollination efficiency (c. 2400 pollen per stigma). Although pollination‐induced structural and histochemical changes in the style allowed many (9.5 ± 2.1) pollen tubes to grow, only cross‐pollen tubes could grow through the style. Gametophytic self‐incompatibility, manifested in the stylar zone, resulted in a significantly slower growth rate of self‐pollen tubes. The occurrence of obligate self‐incompatibility, coupled with increased self‐pollen deposition (geitonogamy), caused a significant number of flowers to abort. Fruit retention in the trees declined from 40% to 12% as a result of abortion of fruits at different stages of development. The number of mature fruits on a tree was negatively correlated (r = ?0.82) with their size. It is inferred that low natural fecundity in A. marmelos is primarily a result of obligate self‐incompatibility and strong post‐fertilization maternal regulation of allocation of resources to the developing fruits. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 572–585.     

Variation of pollen limitation in the early flowering Mediterranean geophyte Narcissus assoanus (Amaryllidaceae)

A recent literature review indicates that pollen limitation of female fertility is a common feature of flowering plants. Despite the ecological and evolutionary significance of pollen limitation, most studies have only examined fertility in a single population at one time. Here we investigate pollen limitation of fruit and seed set in five populations of Narcissus assoanus, a self- sterile, insect-pollinated geophyte, over 2–3 years in southern France. In common with many early spring flowering plants, pollinator visitation to N. assoanus is often infrequent. Supplemental hand-pollination of flowers with outcross pollen significantly increased overall fruit and seed set by 11% and 19%, respectively. Four of the five populations experienced some pollen limitation during the study. For a given year, there was significant variation in pollen limitation among populations. Two of the populations were pollen limited in one year but not in other years in which they were studied. Seed:ovule ratios for open- and hand-pollinated flowers averaged 0.29 and 0.33, respectively. While hand pollination significantly increased the seed:ovule ratio, the low value obtained indicates that the majority of ovules in flowers do not mature seeds despite hand pollination. The role of genetic and environmental factors governing low seed:ovule ratios in N. assoanus is discussed. Received: 28 December 1999 / Accepted: 6 April 2000     

ECOLOGICAL CONSEQUENCES AND PHENOTYPIC CORRELATES OF PETAL SIZE VARIATION IN WILD RADISH,RAPHANUS SATIVUS (BRASSICACEAE)

In this paper we examine some ecological consequences and phenotypic correlates of flower size variation in wild radish, Raphanus sativus. Mean corolla diameter varied significantly among individuals within natural populations of R. sativus in California. On the average, almost 40% of flower biomass was allocated to corolla tissue. In field experiments, pollinator visitation increased significantly with corolla size. Large flowers also accumulated more nectar when pollinators were excluded from plants. In three populations, corolla size was positively correlated with allocation to pollen per flower (either anther weight or pollen grain number), but there was usually no phenotypic relationship between corolla size and several measures of female allocation (ovule number per flower, proportion fruit set, and total seed mass per fruit). Plants growing in the field produced fewer large flowers per unit of stem, and stem biomass was negatively related to corolla size for plants grown under controlled greenhouse conditions. Male and female fitness may covary differently with allocation to attractive floral features in species such as R. sativus, where seed production is often limited by resources rather than by pollen.     

Resource limitation and pollen source (self and outcross) affecting seed production in two louseworts, Pedicularis siphonantha and P. longiflora (Orobanchaceae)

Two animal-pollinated hermaphrodite plants, Pedicularis siphonantha and P. longiflora , have been used to investigate factors limiting seed production in natural populations. To evaluate the potential seed abortion due to resources limitation, seed development has been observed and seed count conducted twice. Seed production per capsule has been compared when flowers have been removed and in a control group. Open pollination has been investigated and pollen supplementation undertaken to estimate the possibility of pollen limitation. Results show that seed abortion is frequent. Stigmatic pollen load is significantly higher than ovule number per ovary under open pollination for both species. Additional self and outcross pollen did not affect seed production. Flower removal significantly increases seed production per capsule, which indicates that seed production of the studied species is limited by available resources. To detect differences in seed production between flowers pollinated by self and outcross pollen, hand pollination of bagged flowers has also been conducted in natural populations of the two Pedicularis species. Compared with open pollination, hand-pollinating self-pollen decreases, while outcross pollen increases seed production per capsule. Such results suggest that inbreeding depression in the two self-compatible species may also result in partial seed abortion under open pollination if mixed pollen is deposited on the stigma. Our results also suggest that pollen interference plays an important role in low female fertility in the two Pedicularis species.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 83–89.     

Testing hypotheses for excess flower production and low fruit-to-flower ratios in a pollinating seed-consuming mutualism

Pollinator attraction, pollen limitation, resource limitation, pollen donation and selective fruit abortion have all been proposed as processes explaining why hermaphroditic plants commonly produce many more flowers than mature fruit. We conducted a series of experiments in Arizona to investigate low fruit-to-flower ratios in senita cacti, which rely exclusively on pollinating seed-consumers. Selective abortion of fruit based on seed predators is of particular interest in this case because plants relying on pollinating seed-consumers are predicted to have such a mechanism to minimize seed loss. Pollinator attraction and pollen dispersal increased with flower number, but fruit set did not, refuting the hypothesis that excess flowers increase fruit set by attracting more pollinators. Fruit set of natural- and hand-pollinated flowers were not different, supporting the resource, rather than pollen, limitation hypothesis. Senita did abort fruit, but not selectively based on pollen quantity, pollen donors, or seed predators. Collectively, these results are consistent with sex allocation theory in that resource allocation to excess flower production can increase pollen dispersal and the male fitness function of flowers, but consequently results in reduced resources available for fruit set. Inconsistent with sex allocation theory, however, fruit production and the female fitness function of flowers may actually increase with flower production. This is because excess flower production lowers pollinator-to-flower ratios and results in fruit abortion, both of which limit the abundance and hence oviposition rates, of pre-dispersal seed predators.     

MATERNAL INVESTMENT AND FRUIT ABORTION IN PHASEOLUS VULGARIS

Plants lose some of their parental investment through fruit abortion. Resource conservation theory predicts that the maternal plant should abort those fruits furthest from maturity in order to conserve scarce resources. Embryo-quality theory predicts that the maternal plant should favor offspring of particular genotypes. I recorded the patterns of maternal investment in floral cohorts of domesticated and wild beans, Phaseolus vulgaris L. In accord with the resource conservation theory, late pod cohorts have the highest fruit abortion rates. When domesticated beans are grown in smaller pots, late cohort survivorship declines. When domesticated beans are crossed with pollen donors of different degrees of relatedness, the fruit abortion rate is the same even though seeds from the outbred crosses weigh more. The challenge for ecologists is to model and test the combined and possibly conflicting effects of selection for resource conservation and high offspring quality.