Yang Niu

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.

Phylogenetic clustering and overdispersion for alpine plants along elevational gradient in the Hengduan Mountains Region, southwest China

To better understand the elevational pattern of phylogenetic structure shown by alpine taxa and the underlying causes, we analyzed the phylogenetic structure of each elevational belt of alpine plants in the Hengduan Mountains Region, measured by net related index (NRI) and net nearest taxon index (NTI). We found both the indices of phylogenetic diversity indicated that alpine plants tended to show phylogenetic overdispersion at low elevational belts, implying that the distribution of alpine plants in these belts was mainly determined by interspecific competition. Alpine plants at higher elevational belts tended to phylogenetic clustering indicated by NRI, and NTI revealed phylogenetic clustering at the belts between 4300 m and 5500 m, which presumably suggested environment filtering and rapid speciation. Above 5500 m, NTI indicated that the phylogenetic structure became random again, perhaps due to the low intensity of filtering and the large distances between plants at the top of the scree slopes. We concluded that phylogenetic structure was, indeed, influenced by the environmental filter, interspecies interaction, rapid speciation during the uplift of the Qinghai–Tibet Plateau, and distance between plants.

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.

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.

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.

Systematic significance of cytology in Cyananthus (Campanulaceae) endemic to the Sino‐Himalayan region

Chromosome numbers and the morphology of members of 24 populations representing 15 species of the Sino‐Himalayan endemic genus Cyananthus (Campanulaceae) were studied using karyological and numerical taxonomic techniques. The chromosome numbers of these taxa are 2n = 10, 12, 14, 24, 28, and the basic chromosome numbers for the genus are x = 5, 6, 7. All species except C. inflatus Hook. f. & Thoms. and C. microphyllus Edgew. were examined for the first time and a new ploidy level (tetraploidy) is reported in the genus. Chromosome measurement data were analyzed using cluster analysis. The relationships between three sections within Cyananthus and evolutionary trends within the genus are discussed in the light of karyological data. The cytological data suggest that Cyananthus is a relatively primitive genus in Campanulaceae and there is significant division within the genus, and polyploidization may have played an important role in the chromosome evolution and speciation of Cyananthus in the Himalayan–Hengduan Mountains.

Sexual allocation in the gynodioecious species Cyananthus macrocalyx (Campanulaceae) at high elevations in the Sino-Himalaya Mountains

Sexual allocation pattern is considered highly relevant for the maintenance of females in the process of evolution from hermaphrodites to dioecy. Theoretical and empirical studies predict that gynodioecious plants should invest more resources in male function under harsher environments and/or when female frequency is higher; and that there are trade-offs between male and female function. We studied sexual allocation pattern in the gynodioecious species Cyananthus macrocalyx in two populations in SW China. The results showed that although the total flower mass in hermaphrodites was significantly higher than that in females, females and hermaphrodites allocated similar biomass to female function (pistil biomass). As a consequence, females allocated relatively more resource to female function, while hermaphrodites allocated relatively more resource to pollinator attractiveness. There was no difference in total flower mass, pistil mass, and pollen production in hermaphrodites between the two populations. These results suggest that the females compensate for the disadvantage of lacking male function by allocating relatively more resource to female function, while hermaphrodites gain fitness through both male and female functions. This study supports the idea that if females are to be maintained in a gynodioecious population, they must allocate more resource to female functions than is necessary for hermaphrodites. However, our results were in contrast to those of previous studies, in that trade-offs between male and female functions and male-biased allocation under high female frequency were not observed. This study adds to the body of research on plant sexual allocation, and for that it is, at least at some aspects, contrary to previous statements, it could be helpful for advancing the theoretical predictions in the future.

Comparison of floral properties and breeding system in dimorphic Buddleja delavayi (Scrophulariaceae)

In this study, floral color, scent composition and emission rate, nectar property, pollinators, and breeding system of dimorphic Buddleja delavayi Gagnep. were investigated. Flower color of B. delavayi was determined using a standard color chart and spectrophotometer, and two distinct color polymorphisms were observed having purple or white flowers. Floral scents of B. delavayi were collected using dynamic headspace adsorption and identified with coupled gas chromatography and mass spectrometry. In total, 28 compounds were identified from the flowers of B. delavayi. The identified scents were divided into three chemical classes based on their biosynthetic origin: terpenes, fatty acid derivatives, and benzenoids. The scent profiles in all individuals were dominated by a few components, such as lilac aldehyde and alcohol, 4‐oxoisophorone, benaldehyde, and oxoisophorone oxide. Floral scent composition (benzenoids and terpenes) showed a significant difference between white and purple flower morphs. Flower color–flower scent associations in B. delavayi were identified with two distinct scent profiles in the two color phenotypes. The studies of other floral characteristics (nectar, floral visitors, breeding system, and fruit set) indicated that floral scent emission rate, nectar volume, visitor visitation frequency, and natural fruit set were not significantly different between the two flower color morphs. Bagging experiments revealed that seed production of B. delavayi is dependent mainly on honeybee Apis cerana. Lastly, this study implies that dimorphic floral color in B. delavayi may have been maintained by floral visitors and nectar guide color.

Sex allocation in gynodioecious Cyananthus delavayi differs between gender morphs and soil quality

Key messageSex allocation in Cyananthus delavayi.AbstractGynodioecy, where females and hermaphrodites coexist in the same natural population, is particularly suitable for predicting the ecological pressures that drive the stability of gender polymorphism. Since females have a disadvantage in that they only contribute to the next generation via ovules, they should gain an advantage via other means, of which resource allocation is an important component. Thus, to study their sex allocation is very helpful to understand how the dimorphic sexual system is maintained in natural systems. We studied the sex allocation patterns and reproductive output of the gynodioecious Cyananthus delavayi in three populations with different soil qualities (organic matter, N, P and K). The hermaphroditic flowers and pistils were much larger than those of female individuals. Although both gender morphs invested similar biomass in the pistils, females allocated more of their resource pool to the seed production, while hermaphrodites allocated more to pollinator advertisement. The pollen production of hermaphrodites did not differ between populations, suggesting that pollen production by hermaphrodites was not limited by soil nutrients. Fruit set of females, but not hermaphrodites, decreased with declining soil quality, whereas seeds per fruit of both females and hermaphrodites were highest in poor soils. Overall, this study shows that females achieve greater reproductive success by allocating more of their resource pool to enhancing seed production, which should favor their presence in gynodioecious populations. The hermaphrodites achieve reproductive success from both pollen and seed production, and unnecessarily reduce their allocation to pollen production. Soil quality should explain, at least partially, the sexual allocation patterns. Furthermore, some of our findings contradict previous hypotheses, thus adding a new example to the body of research on plant sex allocation and the development of future theories.

Function of male and hermaphroditic flowers and size-dependent gender diphasy of Lloydia oxycarpa (Liliaceae) from Hengduan Mountains

Although hermaphroditism is common in flowering plants, unisexual flowers occur in many plant taxa, forming various sexual systems. However, the sexual system of some plants is difficult to determine morphologically, given that their sex expression may be influenced by both genetic and environmental factors. Specifically, androdioecy (the coexistence of both male and hermaphroditic individuals in the same population) has often been confused with the gender diphasy, a gender strategy in which plants change their sex expression between seasons. We studied the reproductive function of male and hermaphroditic flowers of Lloydia oxycarpa (Liliaceae), in order to investigate its sexual system and determine whether it is a gender-diphasic species. We found that although male flowers occur in a considerable number of plants, relative to hermaphrodites, they did not exhibit any significant reproductive advantage in terms of flower size, pollen quantity, attractiveness to visitors or siring success. In addition, this plant has spontaneous self-pollination and showed no inbreeding depression. These results render the maintenance of male individuals almost impossible. Furthermore, a considerable number of individuals changed their sex in successive years. The sex expression was found to be related to bulb size and dry weight, with larger individuals producing hermaphroditic flowers and smaller individuals producing male flowers. These results suggest that L. oxycarpa is not an androdioecious plant but represents a rare case of size-dependent gender diphasy.