Zhi-Qiang Zhang

Autonomous Selfing Provides Reproductive Assurance in an Alpine Ginger Roscoea schneideriana (Zingiberaceae)

BACKGROUND AND AIMS Reproductive assurance, the ability to produce seeds when pollinators or mates are scarce, is thought to be the major advantage of selfing in flowering plants. However, few studies have performed a direct cost-benefit analysis of the selective advantage of selfing, particularly given a long-term perspective among populations or across several flowering seasons within population. This study examined the fertility consequences of autonomous selfing in Roscoea schneideriana (Zingiberaceae), a small perennial Himalayan ginger typically found in habitats at around 3000 m a.s.l. METHODS The floral biology of R. schneideriana was studied in natural populations; the capacity for autonomous selfing was estimated using pollinator exclusion experiments; the timing of selfing was quantified by anther removal at different times during flowering; whether autonomous selfing increases seed production was tested by emasculating flowers; and the magnitude of inbreeding depression was estimated by comparing relative performance of progeny from self- and cross-pollinations. Pollinator observations were also conducted in the natural populations. KEY RESULTS The hooked stigmas of most flowers curl towards the anther and can contact pollen grains at an early stage of anthesis. Flowers with potential pollinators excluded set of as many seeds per fruit as hand-selfed and opened flowers. Autonomous selfing mostly occurs within 2 d of anthesis and can increase seed production by an average of 84 % in four populations during the flowering seasons of 2005-2007. Visits by effective pollinators were extremely rare. The cumulative inbreeding depression of R. schneideriana was 0.226. CONCLUSIONS Autonomous selfing in R. schneideriana is achieved by stigmas curling towards the anthers early in flowering. It is suggested that under the poor pollination conditions, autonomous selfing has been selected for in this alpine ginger because it provides substantial reproductive assurance with very low costs.

Multifunctional bracts enhance plant fitness during flowering and seed development in Rheum nobile (Polygonaceae), a giant herb endemic to the high Himalayas

Specialized bracts are thought to be important for the successful reproduction of some plants and are regarded as adaptations to diverse driving forces. However, few empirical studies have quantified the adaptive significance of bracts within a cost–benefit framework. We explored the adaptive significance of large and showy bracts for reproduction in Rheum nobile, a giant herb endemic to the high Himalayas. We examined whether the bracts enhance reproductive success during flowering and seed development. Bracts increased flower and fruit temperature on sunny days, greatly decreased the intensity of ultraviolet-B (UV-B) radiation reaching flowers and fruits, and prevented pollen grains being washed away by rain. Experiments indicated that high temperature could promote pollen germination, while pollen grains exposed to rain and UV-B radiation at ambient levels were seriously damaged. Furthermore, bract removal decreased the number of pollinators visiting flowers. When bracts were removed before or after flowering, fecundity and progeny quality were adversely affected, but seed predation by larvae of pollinators decreased. A cost–benefit analysis demonstrated that the cost of bracts, i.e., increased seed predation, is modest. Our results suggest that the bracts of R. nobile promote pollen germination, protect pollen grains from rain and intense UV-B radiation, enhance pollinator visitation during flowering, and facilitate the development of fertilized ovules during seed development. We conclude that multifunctional bracts of R. nobile are an effective adaptive strategy in alpine environments and might have been selected for because of abiotic environmental conditions as well as for enhancing pollination success.

Floral closure induced by pollination in gynodioecious Cyananthus delavayi (Campanulaceae): effects of pollen load and type, floral morph and fitness consequences

BACKGROUND AND AIMS Pollination-induced floral changes, which have been widely documented in flowering plants, have been assumed to enhance the plants reproductive success. However, our understanding of the causes and consequences of these changes is still limited. Using an alpine gynodioecious species, Cyananthus delavayi, we investigated the factors affecting floral closure and estimated the fitness consequences of floral closure. METHODS The timings of floral closure and fertilization were determined. The effects of pollen load, pollen type (cross- or self-pollen) and floral morph (female or perfect flower) on the occurrence of floral closure were examined. Ovule fertilization and seed production were examined to investigate the causes and consequences of floral closure. Flowers were manipulated to prevent closing to detect potential benefits for female fitness. KEY RESULTS Floral closure, which could be induced by a very low pollen load, occurred within 4-7 h after pollination, immediately following fertilization. The proportion of closed flowers was influenced by pollen load and floral morph, but not by pollen type. Floral closure was more likely to occur in flowers with a higher proportion of fertilized ovules, but there was no significant difference in seed production between closed and open flowers. Those flowers in which closure was induced by natural pollination had low fruit set and seed production. Additionally, seed production was not influenced by closing-prevented manipulation when sufficient pollen deposition was received. CONCLUSIONS The occurrence of floral closure may be determined by the proportion of fertilized ovules, but this response can be too sensitive to ensure sufficient pollen deposition and can, to some extent, lead to a cost in female fitness. These results implied that the control of floral receptivity by the recipient flowers does not lead to an optimal fitness gain in C. delavayi.

Functional implications of the staminal lever mechanism in Salvia cyclostegia (Lamiaceae)

BACKGROUND AND AIMS Flower morphology and inflorescence architecture affect pollinator foraging behaviour and thereby influence the process of pollination and the reproductive success of plants. This study explored possible ecological functions of the lever-like stamens and the floral design in Salvia cyclostegia. METHODS Flower construction was experimentally manipulated by removing either the lower lever arms or the upper fertile thecae of the two stamens from a flower. The two types of manipulated individuals were intermixed with the control ones and randomly distributed in the population. KEY RESULTS Removing the sterile lower lever arms significantly reduced handling time per flower of the main pollinator, Bombus personatus. Interestingly, this manipulation did not increase the number of flowers probed per plant visit, but instead reduced it, i.e. shortened the visit sequence of the bumble-bees. Both loss of staminal lever function by removing lower lever arms and exclusion of self pollen by removing upper fertile thecae significantly reduced seed set per flower and seed set per plant. Both the manipulations interacted significantly with inflorescence size for the effect on female reproductive output. CONCLUSIONS Though the intact flowers demand a long handling time for pollinators, the reversible staminal lever is of advantage by promoting dispersal of pollen and thus the male function. The particular floral design in S. cyclostegia contributes to the floral constancy of B. personatus bumble-bees, with the lower lever arms acting as an optical cue for foraging cognition.

Habitat-specific responses of seed germination and seedling establishment to soil water condition in two Rheum species in the high Sino-Himalayas

Knowledge of how germination and seedling establishment respond to soil water condition is crucial for plant conservation under global warming and land-use changes. We tested the flooding and drought tolerance of two plant species with different occurrences along a soil water gradient by assessing seed germination, seedling survival, seedling growth, and root characteristics. In the high Sino-Himalayas, Rheum alexandrae typically occurs in wetlands, R. nobile in scree or open slope with well-drained soils. Seeds and seedlings of the two species were subjected to different soil water conditions in controlled greenhouse experiments. Seed germination in both species was inhibited by high soil water content; however, seeds of R. alexandrae were more tolerant to flooding, especially to submergence. Seedling survival, biomass accumulation, root diameter, and root porosity of R. alexandrae increased significantly with increased soil water content, but submergence was lethal for seedlings. Seedling survival, biomass accumulation, and root length of R. nobile increased significantly in response to reduced soil water content. These results indicate that in the two species, seed germination and seedling establishment in response to different soil water condition are habitat-specific. Because both species are susceptible to moderate changes in soil water condition, their species-specific requirements with respect to this factor should be a consideration when planning their conservation.

Plant sexual systems correlated with morphological traits: Reflecting reproductive strategies of alpine plants

Plant sexual systems can be affected by various environmental factors, and thus show a range of morphological differences. The associations between plant sexual systems and morphology are unique, for example, the subnival belt of the Hengduan Mountains has harsh climatic conditions, but hosts remarkable biodiversity. However, these associations have received surprisingly little attention. We analyzed the associations in this region, and compared the pattern of sexual systems with that of other floras, and we also reviewed the breeding system of 36 alpine species. Information on plant sexual systems and these traits were mainly based on published information, herbarium collections, and field observations. The results showed that there was a high proportion of hermaphroditic species and a low proportion of dioecious and monoecious species. Hermaphroditism was strongly associated with showy floral displays, specialist entomophily, and perennial forbs. Dioecy was strongly associated with inconspicuous, pale‐colored flowers, anemophily, and shrubs, whereas monoecy was strongly associated with inconspicuous, pale‐colored flowers, generalist entomophily, and perennial grasses. Although no association was found between sexual systems and fruit type, plants with dry fruits (98.1%) were predominant in our study region. With respect to the plant breeding systems in the alpine region of southwest China, we predicted that pollination service might be particularly low; most of the plants lacked enough outcrossing pollinators. Our data revealed that 97.1% of the hermaphroditic species were self‐compatible, and 88.2% of them showed autonomous or facilitated selfing. The plant sexual systems identified might serve to optimize the probability of pollination and thus ensure reproductive success.

Asymmetrical disassortative pollination in a distylous primrose: the complementary roles of bumblebee nectar robbers and syrphid flies

Heterostyly is a floral polymorphism characterized by reciprocal herkogamy maintained through high levels of mating between morphs, serviced by appropriate pollinators. We studied how differential efficiency and abundance of distinct pollinators affect plant female reproduction in self- and intra-morph incompatible distylous Primula secundiflora. Bumblebees and syrphid flies were found to be the most abundant floral visitors. Bumblebees frequently exhibited nectar-robbing behavior. Because the robbing holes were always situated between the high- and low-level organs on both morphs, nectar-robbing bumblebees only pollinated S-styled flowers. L-styled flowers set four times as many seeds as did S-styled flowers after being visited by pollen-collecting syrphid flies. The natural female fecundity and the magnitude of pollen limitation varied between the morphs within populations because of the mosaic distribution of nectar-robbing bumblebees and syrphid flies. L-styled flowers and S-styled flowers set the same number of seeds after supplemental hand pollination, indicating equivalent female reproductive potential. We suggest that bumblebee nectar robbers and syrphid flies play an important role in sustaining the floral dimorphism of heterostyly in P. secundiflora because of their complementary roles in the pollination system.

A nodding capitulum enhances the reproductive success of Cremanthodium campanulatum (Asteraceae) at high elevations in the Sino–Himalayan Mountains

Background: Flower orientation is considered an evolutionary response to pollinators and abiotic factors. Several members of the genus Cremanthodium (Asteraceae) growing in alpine habitats in the Sino–Himalayas have conspicuously nodding capitula, the function of which is not understood. Aims: We investigated the influences of nodding capitula on floral thermal conditions, pollinator visitation rate, pollen viability and achene production in Cremanthodium campanulatum. Results: (1) the nodding capitulum did not modify internal thermal conditions, but the elongated involucral bracts could shelter pollen from UV-B radiation; (2) water and UV-B radiation seriously reduced the viability of pollen grains; (3) capitula artificially held erect set significantly fewer achenes than nodding ones; and (4) no pollinator preference was observed between capitula that were artificially held erect and natural nodding ones. Conclusions: This study indicated that the nodding capitulum of C. campanulatum may protect pollen from being damaged by UV-B radiation, being washed away from the anther and stigma, and enable it to avoid rain damage during the wet monsoonal period. Our results suggest that nodding capitula in C. campanulatum possibly confer a selective advantage and may be driven by non-biological agents rather than pollinator attractors.

Morphological and molecular evidence for natural hybridization in sympatric population of Roscoea humeana and R. cautleoides (Zingiberaceae)

The sympatric occurrence of some species in Roscoea is very common, but little information is available on natural hybridization. However, some intermediate individuals were found on the sympatric population of Roscoeahumeana and R. cautleoides at Ganhaizi population in northwestern Yunnan Province, China. We suspected that these intermediate individuals were the hybrids of R. humeana and R. cautleoides from the previous evidence, but could not confirm them. In this study, morphometric analysis was followed by examination of HAT-RAPD polymorphisms to determine the occurrence of natural hybridization between sympatric R. humeana and R. cautleoides. The results showed that most morphological characters of the putative hybrids were found to be intermediate between those of R. humeana and R. cautleoides. Meanwhile, molecular analysis confirmed that the morphological intermediates were derived from hybridization between the two species. From the analysis of the NewHybrids, the hybridization individuals were mainly F1s. These results indicated that interspecific hybridization between R. humeana and R. cautleoides indeed occurred in sympatric population.

A Sexually Dimorphic Corolla Appendage Affects Pollen Removal and Floral Longevity in Gynodioecious Cyananthus delavayi (Campanulaceae)

The floral traits of bisexual flowers may evolve in response to selection on both male and female functions, but the relative importance of selection associated with each of these two aspects is poorly resolved. Sexually dimorphic traits in plants with unisexual flowers may reflect gender-specific selection, providing opportunities for gaining an increased understanding of the evolution of specific floral traits. We examined sexually dimorphic patterns of floral traits in perfect and female flowers of the gynodioecious species Cyananthus delavayi. A special corolla appendage, the throat hair, was investigated experimentally to examine its influences on male and female function. We found that perfect flowers have larger corollas and much longer throat hairs than female flowers, while female ones have much exerted stigmas. The presence of throat hairs prolonged the duration of pollen presentation by restricting the amount of pollen removed by pollen-collecting bees during each visit. Floral longevity was negatively related to the rate of pollen removal. When pollen removal rate was limited in perfect flowers, the duration of the female phases diminished with the increased male phase duration. There was a weak negative correlation between throat hair length and seed number per fruit in female flowers, but this correlation was not significant in perfect flowers. These results suggest that throat hairs may enhance male function in terms of prolonged pollen presentation. However, throat hairs have no obvious effect on female function in terms of seed number per fruit. The marked sexual dimorphism of this corolla appendage in C. delavayi is likely to have evolved and been maintained by gender-specific selection.