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.     

Responses of the clonal sedge, Carex bigelowii, to two seasons of simulated climate change

The effect of enhanced temperature was studied in open-top chambers on the arctic clonal sedge Carex bigelowii Torr. ex Schwein. at Latnjajaure Field Station, Swedish Lapland for two growing seasons. The flowering phenology of C. bigelowii accelerated during the first field season that the open-top chambers were erected. After the second season of perturbation the sexual reproductive investment was higher in the open-top chambers than in the controls and flowering phenology was still accelerated. Leaf phenology and the growth of vegetative tillers followed the same pattern in the open-top chambers as in the controls, but the leaves were longer in the open-top chambers from the end of June until the end of August. The results suggest that relatively late flowering clonal graminoids may be favoured by warmer climate, at least in the short term.     

Limitations to Reproductive Success in the Dioecious Tree Rhamnus davurica

The reproductive success of a female plant in a dioecious species may be affected by pollen limitation and resource limitation. This study presents evidence that the reproductive success of the dioecious understorey tree species, Rhamnus davurica, is affected by the distance to the nearest male. The sex ratios were female-biased, although showing fluctuations in the three years of conducting the study. The mortality rate of females was higher than that of males indicating a trade-off between reproduction and survival. Altogether 49 females, designated as “focal females”, were randomly selected for monitoring their reproductive status between April and October in 2010. But successful reproduction (meaning that the flowering female trees had fruit in the fruiting season) was observed only in 28 females in 2011 and 16 females in 2012. The method of path analysis was applied to determine the effect of topography, local competition and proximity to the nearest male on the fruit set of the females. In the three years of the study, elevation, competition and female size had no significant effect on the fruit set. The distance to the nearest male, however, had a significant effect on fruit set. Number of fruits and fruit set were decreased with increasing distance to the nearest male. It was possible to estimate maximum fruit set, based on the comparatively large dataset. The number of fruits and the fruit set are exponentially related to the distance to the nearest male and the relationships are described by an exponential model. The results of this study support the importance of pollen limitation on the reproductive success in Rhamnus davurica.     

Movement herkogamy in Kosteletzkya virginica: Effect on reproductive success and contribution to pollen receipt and reproductive assurance

Herkogamy may act as a mechanism to reduce interference between the reproductive functions of female and male organs, but too great a distance between stigmas and anthers may also decrease the outcross pollen transfer and male and female fitness. Our goal was to quantify the effect of movement herkogamy in Kosteletzkya virginica on reproductive success and its contribution to pollen receipt and reproductive assurance. Field investigations and hand-pollination experiments were performed to test: (1) the relationship between herkogamy before curvature of un-pollinated lobes and reproductive success under pollinator exclusion; (2) the contribution of movement herkogamy to pollen receipt; and (3) the relationship between pollen load and seed set. We noted that variation in herkogamy before curvature of un-pollinated lobes did not affect the outcome of autogamous self-pollination measured by fruit set and mean number of seeds per fruit. Under natural conditions, 39% of pollen grains deposited per flower were from autogamous self-pollination via movement herkogamy resulting in contact between anthers and stigmas. The relationship between the amount of pollen grains and the number of seeds per capsule was highly significant. Flowers producing four or five seeds received more pollen grains than those producing only one seed. Our data provides experimental evidence that movement herkogamy, which decreases the separation between anthers and stigmas and results in delayed self-pollination, contributes to pollen receipt and further enhances seed set when pollinators are scarce or absent.     

Movement herkogamy in Kosteletzkya virginica: Effect on reproductive success and contribution to pollen receipt and reproductive assurance

Herkogamy may act as a mechanism to reduce interference between the reproductive functions of female and male organs, but too great a distance between stigmas and anthers may also decrease the outcross pollen transfer and male and female fitness. Our goal was to quantify the effect of movement herkogamy in Kosteletzkya virginica on reproductive success and its contribution to pollen receipt and reproductive assurance. Field investigations and hand-pollination experiments were performed to test: (1) the relationship between herkogamy before curvature of un-pollinated lobes and reproductive success under pollinator exclusion; (2) the contribution of movement herkogamy to pollen receipt; and (3) the relationship between pollen load and seed set. We noted that variation in herkogamy before curvature of un-pollinated lobes did not affect the outcome of autogamous self-pollination measured by fruit set and mean number of seeds per fruit. Under natural conditions, 39% of pollen grains deposited per flower were from autogamous self-pollination via movement herkogamy resulting in contact between anthers and stigmas. The relationship between the amount of pollen grains and the number of seeds per capsule was highly significant. Flowers producing four or five seeds received more pollen grains than those producing only one seed. Our data provides experimental evidence that movement herkogamy, which decreases the separation between anthers and stigmas and results in delayed self-pollination, contributes to pollen receipt and further enhances seed set when pollinators are scarce or absent.     

Tibouchina pulchra (Melastomataceae): reproductive biology of a tree species at two sites of an elevational gradient in the Atlantic rainforest in Brazil

Low-temperature environments interfere with plant reproduction by reducing the frequency of pollinators, and this may favour reproductive strategies such as self-pollination and apomixis. Tibouchina pulchra is a common tree species that occurs at high and low sites of the Brazilian Atlantic rainforest. This study focussed on the pollination biology and breeding system of this species, describing the pollinators and the reproductive success at the two sites of an elevational gradient. Observations were made to determine extent of flowering and fruiting, to identify the richness and abundance of pollinators, and to record data on the floral and reproductive biology at these two sites. Despite more dense flowering at the high site, five visits of bees (two species) were recorded during the observation time (60?h), whereas at the low site there were 948 visits (seven species) during the same period. In contrast with the low site, the flowers of the high site released and received few pollen grains on the stigma. At the high site less fruit was set with fewer seeds as a result of open pollination than at the low site; at that site, however, more seeds were obtained from cross-pollination than at the low site. Tibouchina pulchra is self-compatible; however it is not apomictic and needs pollinators for seed set at both sites. Life-history traits other than the breeding system, for example more dense flowering, advantage of greater fertility in cross-pollination, and multiple reproductive events during the lifetime of the tree may reduce inbreeding depression, increase the hybrid vigour, and balance the lack of pollinators at the high site.     

Phenology and phenotypic natural selection on the flowering time of a deceit-pollinated tropical orchid, Myrmecophila christinae

BACKGROUND AND AIMS: Flowering phenology is described and the effect of flowering time on pollination success is evaluated in the deceit-pollinated tropical orchid, Myrmecophila christinae. It was expected that, due to this species' deceit pollination strategy and low observed pollinator visit rate, there would be a higher probability of natural selection events favouring individuals flowering away from the population flowering peak. METHODS: The study covers two consecutive years and four populations of M. christinae located along the north coast of the Yucatán Peninsula. For phenological and pollination success data, a total of 110 individuals were monitored weekly in 1998, and 83 individuals in 1999, during all the flowering and fruiting season. KEY RESULTS: The results showed significant differences in the probability of donating and receiving pollen throughout the flowering season. The probability of receiving or donating pollen increased the further an individual flowering was from the flowering peak. Regression analysis showed directional and disruptive phenotypic natural selection gradients, suggesting the presence of selection events unfavourable to flowering during flowering peak, for both male success (pollen removal) and female success (fruit production). However, the intensity and significance of the natural selection events varied between populations from year to year. The variation between seasons and populations was apparently due to variations in the density of reproductive individuals in each population and each season. CONCLUSIONS: As in other deceit-pollinated orchids, natural selection in M. christinae favours individuals flowering early or late in relation to population peak flowering. However, results also suggested a fluctuating regime of selective events act on flowering time of M. christinae.     

Floral and reproductive biology of Drosophyllum lusitanicum (L.) Link (Droseraceae)

We studied the floral and reproductive biology of Drosophyllym lusitanicum (Droseraceae), a species endemic to the western Iberian Peninsula and northwest Morocco. Flowering lasted from March to August, peaking in April. The species is clearly homogamous, with pollen germination and stigma receptivity occurring even in preanthesis. This was reflected in the high rates of fruit set and seed set in bagged inflorescences. Fructification did not differ significantly between different pollination treatments, although we did find differences between some treatments in the numbers of viable seeds per flower and fruit. However, seed weight did not differ significantly. The importance of self-pollination over cross-pollination was also supported by the low percentages of fructification (24.67%) after emasculation and open pollination. Metameric deviations were seen in flowers, although this abnormality was not translated as any reproductive benefit or disadvantage (fructification, number or weight of seeds) in comparison with normal flowers.     

Factors affecting fruit and seed production of Paeonia ostii “Feng Dan”, an economically important oil tree

Many factors may affect reproduction of animal-pollinated species. In this study, the effects of pollen limitation, attractive traits (flower number, plant height and flower width) and flowering phenological traits (flowering onset, duration and synchrony) on female reproduction, as well as the patterns of variation in fruit and seed production within plants, were investigated in Paeonia ostii “Feng Dan” over two flowering seasons (2018 and 2019). Fruit set was very high (90%), and pollen supplementation did not increase fruit and seed production in either year, indicating no pollen limitation. Fruit set, ovule number per fruit and mean individual seed weight per fruit were not affected by any of the six attractive and phenological traits in either year, whereas seed number per fruit was related to the three attractive traits in one or both years. Seed number per plant was positively affected by the three attractive traits and best explained by flower number in both years, but the effect of each of the three phenological traits on seed number per plant differed between years. Within plants, the fruit set, ovule number, seed set and seed number per fruit declined from early- to late-opening flowers, presumably because of resource preemption, but the mean individual seed weight did not vary across the flowering sequence. Our study shows that attractive traits of Paeonia ostii “Feng Dan” are more important than flowering phenological traits in the prediction of total seed production per plant.     

Sequential variation in the components of reproductive success in the distylousJasminum fruticans (Oleaceae)

A 2-year study of three natural populations of the distylousJasminum fruticans showed that mean fruit and seed production were significantly greater in shortstyled plants (thrums) than in long-styled plants (pins). In this study, we investigated the role of four sequential factors which may differentially influence fruit and seed set in the two floral morphs: (1) differences in flowering phenology, (2) a limitation of pollen transfer towards pins, (3) a differential capacity of the two morphs to act through famale and male function and (4) differential fruit abortion in the two morphs. Fruit set was significantly influenced by differences in flowering phenology although there were no differences in flowering time between the two morphs. supplementary pollinations in a natural population significantly increased fruit set and reduced the difference in fruit set between the two morphs in relation to controls, indicating a limitation on pollen transfer which was most severe towards pin stigmas. In reciprocal crosses, seed set was significantly dependent on the paternal and maternal identity of the pin parent. There was no significant variation among thrums for their performance as male or female parent. Furthermore, individual pin plants with relatively high percentage seed set as female parents gave poor seed set as male parents and vice-versa. Whereas fruit removal had no effect on seed number in thrum plants, a greater proportion of viable seeds were produced on pin plants which were left to naturally mature their fruits than on pins which had fruits artificially removed, suggesting the occurrence of selective fruit abortion in pins but not in thrums. The initially greater maternal fitness of thrums due to their greater success as pollen recipients may thus be opposed by increased viable seed set in the pins due to factors acting after the pollination stage. The relative reproductive success of floral morphs in the distylousJ. fruticans is thus differentially influenced by ecological factors occurring at different stages of the reproductive process.     

Pollination efficiency and reproductive patterns in relation to local plant density,population size,and floral display in the rewarding Listera ovata (Orchidaceae)

Pollination efficiency and reproductive success vary strongly among populations of most animal‐pollinated plant species, depending on their size and local density, whereas individual plants within populations experience varying levels of reproductive output as a result of differences in floral display. Although most orchid species have been shown to be severely pollination limited, few studies have investigated the impact of the above‐mentioned factors on pollination success and reproduction, especially in rewarding species. In this study, the impact of population size, local density of flowering plants, and floral display on the rates of pollinia export and fruit production was investigated in 13 natural populations of the rewarding terrestrial orchid Listera ovata. In addition, an emasculation experiment was set up to examine how floral display and local density of flowering plants affected the relative importance of cross‐ vs. geitonogamous pollination in determining fruit set. In the studied populations, pollination efficiency, pollen removal, and fruit set increased with increasing population size until a threshold value of 30–40 flowering plants was reached, above which pollination efficiency and reproductive output decreased again. On average, plants with large floral displays showed higher proportional pollinia removal and fruit set compared with smaller plants. Fruit production was also significantly and positively related to local plant density, whereas emasculation did not affect the relationship between local plant density and fruit set, suggesting that geitonogamous pollination did not affect the outcome of female function. The results of this study are discussed in the light of the flowering mechanism of the species and its generalized pollination system. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157 , 713–721.     

Defense mechanisms against grazing: a study of trypsin inhibitor responses to simulated grazing in the sedge Carex bigelowii

Trypsin inhibitors have been suggested to constitute an inducible defense in the sedge Carex bigelowii , and some former studies suggest that this might be a cause for the cyclic population dynamics in many alpine and arctic small mammals, for example lemmings ( Lemmus lemmus ). We investigated this further by using a method of simulated grazing (clipping) at different intensities, in three different habitats with varying resource availability, with different harvest times (hours after clipping), and two different stages of ramets (reproductive/vegetative) in a study from the Swedish mountain range. Our results do not indicate that C. bigelowii has an inducible defense constituted by an increase in trypsin inhibitor activity (TIA), but rather that the amount of soluble plant proteins (SPP) is lowered in wounded plants. The responses were somewhat different in the three habitats, with ramets growing in the marsh showing the highest ratio of TIA to SPP, due to low amounts of SPP. We did not find any significant effects of harvest time, or of the stage of the ramet that could support the hypothesis of an inducible defense. To conclude, we could not find any evidence for an inducible defense consisting of trypsin inhibitors in Carex bigelowii ramets, but we did find variations in the amount of SPP that may have nutritional consequences for herbivores.     

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。     

Does Neighborhood Floral Display Matter? Fruit Set in Carpenter Bee‐pollinated Heterophragma quadriloculare and Beetle‐pollinated Lasiosiphon eriocephalus

Fruit set is pollen‐limited in the self‐incompatible tree Heterophragma quadriloculare (Bignoniaceae), pollinated by long‐distance flying carpenter bees, and in the self‐compatible shrub Lasiosiphon eriocephalus (Thymdeaceae), pollinated by weak‐flying, sedentary beetles. We studied a single H. quadriloculare population over high and low flowering years and found no difference in bee visitation rates between these years. For H. quadriloculare, neighborhood floral display did not make a significant contribution to reproductive success. We investigated dense and sparse L. eriocephalus populations in the same year. In the low density L. eriocephalus population, individual floral displays were higher than in the dense population, yet reproductive success was low, indicating that plant isolation was a major factor influencing fruit set. This result was due to the extremely low number of beetles per plant and per flower in this population. In the dense L. eriocephalus population, although the displays of individual neighbors were smaller and plants were closer, neighborhood floral display did not contribute significantly to reproductive success, whereas the effect of individual floral display was ambiguous. Species with self‐incompatible rather than self‐compatible breeding systems are expected to experience neighborhood effects on reproductive success; however, at the spatial scale and floral display levels of plants in this study, only individual floral display affected fruit set in H. quadriloculare, whereas neither individual nor neighborhood display influenced fruit set in L. eriocephalus. Therefore, pollinator type, pollinator behavior, and plant and population isolation, rather than breeding system alone, will determine if neighborhood floral display affects fruit set.     

Durian flowering, pollination and incompatibility studies

Seasonal fluctuations in weather conditions were found to influence the flowering and fruiting phenology and reproductive biology of durian grown around Darwin. Manually assisted cross-pollination resulted in higher fruit set of 31% in contrast to < 10% by selfing, producing higher yields and better quality fruit. Self-pollination resulted in more fruit abortion and produced fruits that were mis-shapen, curved with lower numbers of arils/locule, arils/fruit and lower numbers of seeds which were usually shrunken and disfunctional. Evidence from these studies also confirmed that the self-incompatibility system in durian is gametophytically controlled and that cultivars could be totally self-incompatible, partially self-incompatible or totally compatible. The durian pollen was found to exert metaxenia effects that influenced the development of the fruit characteristics and also xenia effect on the seed.     

Evidence for stabilising selection acting on flowering time in Arum maculatum (Araceae): the influence of phylogeny on adaptation

The relationship between flowering time and reproductive success was investigated in the fly-pollinated, monoecious perennial herb Arum maculatum L. (Araceae). This species temporarily traps its principle pollinator, a psychodid midge. Probability of fruit set was analysed in relation to early, peak and late periods of the flowering phenology of four British populations between 1992 and 1997. In three out of five cases, plants which flowered during early and late periods were significantly less likely to set fruit. In addition, one population showed a similar relationship for percentage fruit set of individual inflorescences, and seeds from peak-flowering plants were significantly heavier. There was no variation in number of female flowers per inflorescence over the flowering season. Probability of fruit set appears to be mediated by the likelihood of trapping psychodid midges that have previously been trapped and picked up pollen, an unlikely event during early and late flowering periods when few inflorescences are open. The majority of plants in all populations produce only one inflorescence which means that timing of flowering may be crucial to reproductive success. We interpret our findings as evidence that stabilising selection may be acting on some populations and/or during some years. The ultimate cause, however, can be related to the very short (12–18 h) female phase of each inflorescence, a phylogenetically conservative trait within the Araceae. Received: 19 August 1998 / 15 February 1999     

Flowering phenology, display size, and fruit set in an understory dioecious shrub, Aucuba japonica (Cornaceae)

I investigated the effects of display size and flowering phenology on fruit set in Aucuba japonica, an understory dioecious shrub pollinated by opportunistic insects. Natural variations in display size, flowering phenology, and fruit set were monitored in 1997. A hand-pollination experiment was also conducted to check whether pollen limitation was a factor in fruit set in the field. Increases in floral display size did not affect fruit set; the proportion of flowers that set fruit was almost constant irrespective of the total number of flowers per inflorescence, the total number of inflorescences per plant, and the total number of flowers per plant. The hand-pollination experiment showed that fruit set was not pollen limited despite the low mating probability that resulted from the combination of dioecism and the species' dependence on opportunistic pollinators. This was due, in part, to the fact that female flowers did not have a predetermined period of receptivity, but instead remained receptive until they received pollen. In contrast, flowering phenology did affect fruit set. Fruit set was most abundant when male and female flowering was most abundant. This suggests there was some degree of pollen limitation during the part of the flowering season when male flowers were scarce.     

Endogamy costs and reproductive biology of <Emphasis Type="Italic">Laelia autumnalis</Emphasis>, an endemic orchid of Mexico

It has been hypothesized that the fitness of selfed progeny must be lowered by inbreeding depression. Most research into the breeding systems of orchids shows a similar fruit set from outcrossing and self-pollination, but few studies have measured seed production and viability. In five populations of Laelia autumnalis, in central México, we studied the species reproductive system and the early costs of endogamy. We performed spontaneous self-pollination, apomixis, assisted self-pollination, cross-pollination, and exogamous pollination (using pollen from a different population) treatments of bagged flowers and measured fruit set, seed production, and germination. No fruits were produced in the apomixis and spontaneous self-pollination treatments. Reproductive success from self-pollination was lower than that from cross- and exogamous pollination and no difference was found between the latter two treatments. The ratio between self-pollination and cross-pollination success in different traits ranged from 0.80?±?0.18 (fruit set) to 0.22?±?0.13 (seeds with embryo). The latter value suggests a high endogamy costs in the production of viable seeds. The concatenated success of the different traits studied showed that the relative fitness of self-pollination was 63% lower than with cross-pollination. Laelia autumnalis is a self-compatible non-autogamous species in which the cost of endogamy occurs at seed production and cannot be compensated for at other early stages. This also occurs in other orchid species and is likely to be a general pattern.     

Autogamy in Boronia falcifolia: consequences for inbreeding and persistence in fragmented coastal heaths

Coastal heath is becoming increasingly fragmented, resulting in reduced pollinator abundance affecting species viability. The reproductive ecology and inbreeding of a common coastal heath species, Boronia falcifolia (Rutaceae), was investigated. Pollination observations and experiments were carried out at one site, and inbreeding was assessed using genetic markers at eight populations. Four pollination treatments were assessed: cross-pollination, self-pollination, autogamous pollination and open (insect) pollination. Boronia falcifolia was found to be highly self-compatible, frequently setting seeds from autogamous pollination. There were no significant differences between the autogamous, self and cross pollen treatments for any measures of reproductive success (pollen tube growth, fruit set, swollen locules and seed production). Insect activity resulted in fewer seeds per fruit than hand cross pollination, although pollinator visits were few. All populations studied were highly inbred. Autogamous self-pollination is widespread in B. falcifolia and this combined with its ability to vegetatively regenerate enhances its potential for survival when pollinators are few.     

Temporal and spatial variation in flower and fruit production in a food-deceptive orchid: a five-year study

Deceptive orchids are generally characterized by low levels of fruit set; however, there may be substantial variations in fruit set between sites and years. Within a single population, individual plants may also differ greatly in their reproductive output as a result of differences in inflorescence size or local density. In this study, we determined flower and fruit production over 5 years in two populations of the food-deceptive orchid, Orchis purpurea . All plants were monitored annually for survival and flowering at each site to determine whether flowering and fruiting induced costs. The number of flowers per inflorescence varied considerably from year to year (min: 36.6, max: 49.5). Average fruit set was low (7%) and varied considerably among years and populations. A considerable proportion of plants also failed to set any fruit. However, the probability of producing at least one fruit was not affected by inflorescence size or local density. The number of fruits was significantly related to inflorescence size, but proportional fruit set was not. Local density also did not affect the number of fruits, nor proportional fruit set. There was also no evidence that plants with large inflorescence size or high fruiting success had a larger probability of remaining vegetative the year after flowering than plants with small inflorescence size or low fruiting success. Our results suggest that pollinator-mediated selective forces on inflorescence size through female reproductive success alone are weak, most likely because of the low overall level of visitation and the resulting uncertainty of pollination at the individual level. Our results further demonstrate that investigation of patterns of fruit set over several years is needed to better understand the variability in female reproductive success that is typical of most plant–pollinator interactions.