Effects of coppicing on demographic structure, fruit and seed set in Orchis mascula

Opening of the forest canopy may result in higher fitness of understory plants due to increased light intensity on the forest soil and increased nutrient availability. Especially, tuberous orchids may profit from increased light as photosynthetic assimilation products and nutrients are accumulated in their belowground storage organs. We investigated the effects of coppicing on demographic structure and fruit and seed set in 15 populations of the tuberous, perennial orchid Orchis mascula, seven of which were located in undisturbed, shaded woodland and eight in recently coppiced woodland. Coppicing resulted in a massive increase in flowering and in increased fruit set. On average, 42.9% of all individuals consisted of flowering plants in coppiced woodland, whereas in undisturbed woodland only 20.8% of all individuals flowered. The percentage fruit set varied between 20% and 55% in coppiced woodland and was strongly related to population size (measured as the number of flowering individuals). Mean fruit weight and fruit size were also significantly higher in populations located in coppiced woodland than in undisturbed woodland, whereas the proportion of viable seeds was not related to woodland type. Given that O. mascula is dependent on seed set to fill space with new individuals, these results suggest that the long-term survival of this species strongly depends on frequent opening of the canopy.     

Causes of Low Fruit and Seed Set in Bird-Pollinated Anagyris foetida (Leguminosae): Pollen Limitation and Other Extrinsic Factors

We studied the extrinsic factors behind the low number of offspring in two populations of Anagyris foetida, a bird-pollinated leguminous shrub of the Mediterranean region. Fruit initiation was pollen limited, but fruit maturation was not. This limiting effect varied between flowering seasons and between populations, and also within a given phenological stage. In the first season, the early flowers had the lowest fruit set, while in the second season fruit set was highest in these early flowers. This was possibly related to the pollination environment. Seed initiation (fertilized ovules) increased following pollen supplementation, but this was not translated into a significant increase in either the number of seeds per fruit, or seed mass. This indicates that seed initiation is pollen limited but that other factors (e.g., resource availability) are involved in seed maturation. Abiotic factors such as excess humidity during flowering were responsible for the loss of inflorescences, especially in one of the two populations. In this population, the prevailing wind in autumn-winter was less effective in drying the flowers when there was excess humidity. Also, significantly fewer inflorescences were lost from solitary plants than from clustered plants, probably reflecting the beneficial action of the wind and the greater light levels during flowering. Of the biotic factors analyzed, sheep predation was the most important, being worse in drought years. This predation, by affecting population density, could modify the plant-pollinator interaction and severely reduce the plant’s breeding success because of its pollen limitation.     

Patterns of fruit and seed set within inflorescences of Pancratium maritimum (Amaryllidaceae): nonuniform pollination,resource limitation,or architectural effects?

We investigated patterns of fruit and seed production on inflorescences of a population of Pancratium maritimum in northwest Spain over a 2-yr period. Initial findings showed that the earliest opening flowers on an inflorescence are more likely to set fruit and produce more seeds than later opening flowers and that this pattern is maintained throughout the flowering season. Supplementary pollination and flower-removal experiments were performed to investigate whether the observed pattern is attributable (a) to variation in pollen receipt, (b) to sequestration of resources by the earliest flowers on an inflorescence, and/or (c) to "architectural" limitations on the fruit/seed production of later flowers. Supplementary pollination did not improve fruit or seed production by late flowers in either of the 2 yr of study. In flower-removal experiments, the remaining flowers showed improved fruit set and mean number of seeds per flower, by comparison with flowers in the same position on control inflorescences. When all flowers except the latest third were removed, these showed fruit set and seed production similar to those of early flowers on control inflorescences. These results strongly suggest that the observed within-inflorescence patterns of fruit and seed production in P. maritimum are mainly attributable to competition for resources (i.e., explanation b), though other adaptive explanations cannot be ruled out.     

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.     

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     

Fruit-set and seed weight variation inAnthyllis vulneraria subsp.vulgaris (Fabaceae)

Fruit-set and seed weight variation was studied in a population ofAnthyllis vulneraria subsp.vulgaris (Fabaceae) in northwestern Spain. The plants produce several shoots, each bearing two to four inflorescences that open one at a time from bottom to top. Fruit-set, seed-pod weight and seed weight were found to be significantly higher in proximal inflorescences than in distal inflorescences of the same shoot; mean seed weight was up to three times higher in proximal than in distal inflorescences. By contrast, none of the three variables varied significantly among plants or among shoots of the same plant. Similarly, none of the three variables differed significantly between early- and late-flowering plants, or between plants monitored in 1993 and in 1994. These results are compatible with the view that shoots function as semiautonomous units as regards resource allocation, and that within the shoot resources are preferentially allocated to proximal (= early-opening) inflorescences. In the plants studied, the ratio of seed-pod weight to seed weight was fairly constant, suggesting that the pod is important for seed success.     

Factors regulating fruit and seed production in the desert annual Lesquerella gordonii

Summary This study examines the pattern of reproduction and factors that limit the maternal reproductive output in the self-incompatible annual, Lesquerella gordonii. Greenhouse experiments showed that fruits do not abscise and they mature if they contain at least one fertilized (outcrossed) ovule. Field studies showed that flowering increased to a peak and then fell off rapidly, while seed production/fruit decreased through the season. Pollen addition did not increase the number of fruit or seeds per fruit, suggesting that fruit and seed production are not pollinator limited. Addition of water increased fruit and seed production by extending the life of the plant, by increasing the number of stalks which flowered, and by increasing the number of seeds per fruit. This indicates that fruit and seed production are resource limited. Fruits are relatively inexpensive as compared to seeds as over 93% of the water in fruits is contained in the seeds. Fifty-three percent of the plants were lost to herbivory as were 64.6% of the fruits on the remaining plants, indicating that herbivory also limited fruit and seed production. It is hypothesized that high fruit set coupled with variable seed set in L. gordonii is an adaptive response to unpredictable, variable resource levels, and high herbivory risk.     

Consequences of differences in flowering date on seed production in Heloniopsis orientalis (Liliaceae)

We examined the consequences of differences in flowering date on seed production in the self-compatible herb Heloniopsis orientalis. The number of selfed seeds per fruit, as determined by microsatellite markers, did not depend on when the plant flowered, whereas the number of outcrossed seeds per fruit increased with later flowering dates. Consequently, the selfing rate decreased with later flowering dates. The number of seeds (including both selfed and outcrossed ones) per fruit and the seed?:?ovule ratio increased with later flowering dates. We also examined the effects of pollinators and plant size on seed production. The visitation rate of Diptera did not depend on the flowering season, whereas that of Hymenoptera markedly increased as the flowering season progressed. Diptera stayed longer than Hymenoptera on each plant and flower. Seed production per fruit did not depend on plant size. Thus, the change in selfing rate associated with later flowering dates resulted from the seasonal change in pollinators rather than plant size.     

Variation in fruit- and seed set among and within inflorescences of Melampyrum roseum var. japonicum (Scrophulariaceae)

Abstract Variations in fruit set and seed set among and within inflorescences of the annual herb, Melampyrum roseum var. japonicum, were studied. Under natural conditions, although the mean fruit set was slightly different among inflorescences, the mean seedset was not significantly different among inflorescences within the plants. In constrast, within the inflorescences, the flowers located at a lower position of the inflorescence and which opened earlier showed higher fruit set than those at a higher position and which opened later. However, the seed set of matured capsules were not significantly different from each other, regardless of the position of flowers within the inflorescences. Patterns of the fruit- and seed set under open pollination indicated that variation in seed reproduction of M. roseum is due to variation in fruit production. The results of clipping experiments of flowers revealed that there was no functional limitation in seed production among flowers located at various positions within the inflorescence. It seemed that the variation in the fruit set within the inflorescences of M. roseum was not attributable to 'architectural effects'. Reduction of the number of flowers within the inflorescences resulted in an increase of fruit set and seed weight, indicating that the flowers in an inflorescence compete for resources. This phenomenon supports the 'resource competition hypothesis', and variation in fruit set within the inflorescence is attributable to competition among flowers within the inflorescence for limited resources. Consequently, it was concluded that, under natural conditions, the early blooming flowers located at lower positions of the inflorescences obtain more resources and produce more fruits than the late blooming flowers located at higher positions in M. roseum .     

Floral sex allocation and reproductive success within inflorescences of Hosta ventricosa,a pseudogamous apomict

Aims Within inflorescences of sexually reproducing hermaphrodites, the production of ovules, fruits and seeds commonly declines from basal (early-opening) to distal (late-opening) flowers, while pollen production remains constant or only changes slightly, with the result that distal (late-opening) flowers become functionally male. However, few empirical studies have specifically examined whether or not changes in allocation to pollen production actually lead to changes in the number of seeds sired, a more direct measure of male fitness. In pseudogamous apomicts, fitness depends on the number of seeds produced; thus, a contrasting pattern of variation in the pollen-to-ovule (P/O) ratio within inflorescences might be expected.Methods We investigated floral sex allocation and reproductive success within racemes of Hosta ventricosa, a pseudogamous apomictic hermaphrodite possessing flowers that open acropetally. We quantified variations in pollen number, ovule number, the P/O ratio and fruit and seed production, from 2007 to 2011, among flowers within racemes of four populations of H. ventricosa in southwest China. Ecological causes for fruit and seed production were evaluated by observing patterns of pollen deposition, flower removal and supplemental pollination.Important findings Pollen number, ovule number and the P/O ratio declined from basal-to-distal positions in all sampled populations (years). Fruit and seed production decreased distally in most populations (years). Low fruit and seed set of distal flowers was not due to pollen limitation because pollen deposition never declined distally and the low fruit and seed set of distal flowers remained even after supplementary pollination was provided. The flower-removal experiment indicated that inter-fruit competition for resources among flowers was common. The low P/O ratio of distal flowers in H. ventricosa might be favored because they were unable to obtain fitness by donating pollen and siring seeds on other plants. Our study may help to understand the adaptive significance of sex allocation among flowers within inflorescences of sexually reproducing hermaphrodites.     

药物植物桃儿七不同种群种子产量初步研究

对药用植物桃儿七5个不同种群种子产量的研究表明：各种群中每花平均胚珠数、每果实平均种子数及种群平均结实率与种群所在的海拔高度呈正相关;不同种群个体平均胚珠数的变异范围为58.97-87.97,平均种子数变异范围为40.02-80.58,平均结籽率变异范围为61.29%-91.60%;果料内单粒种子平均干重与种群分布海拔呈明显负相关,与果实内种子数呈弱负相关;同一种群在不同年份间平均各上子产量差异不大;种子产量除主要受个体营养状况影响外,还受海拔及其它因素的影响;果实内种子数与果实大小及重量呈正相关,但单粒种子平均重量与果实大小及果实内种子总重量无显著相关。     

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     

High temperature stress of Brassica napus during flowering reduces micro- and megagametophyte fertility, induces fruit abortion, and disrupts seed production

High temperature stress (HTS), during flowering, decreases seed production in many plants. To determine the effect of a moderate HTS on flowering, fruit and seed set in Brassica napus, plants were exposed to a HTS (8/16 h dark/light, 18 degrees C night, ramped at 2 degrees C h-1, over 6 h, to 35 degrees C for 4 h, ramped at 2 degrees C h-1 back to 23 degrees C for 6 h) for 1 or 2 weeks after the initiation of flowering. Although flowering on the HTS-treated plants, during both the 1 week and 2 week HTS treatments, was equal to that of control-grown plants, fruit and seed development, as well as seed weight, were significantly reduced. Under HTS, flowers either developed into seedless, parthenocarpic fruit or aborted on the stem. At the cessation of the HTS, plants compensated for the lack of fruit and seed production by increasing the number of lateral inflorescences produced. During the HTS, pollen viability and germinability were slightly reduced. In vitro pollen tube growth at 35 degrees C, from both control pollen and pollen developed under a HTS, appeared abnormal, however, in vivo tube growth to the micropyle appeared normal. Reciprocal pollination of HTS or control pistils with HTS or control pollen indicated that the combined effects of HTS on both micro- and megagametophytes was required to knock out fruit and seed development. Expression profiles for a subset of HEAT SHOCK PROTEINs (HSP101, HSP70, HSP17.6) showed that both micro- and megagametophytes were thermosensitive despite HTS-induced expression from these genes.     

Patterns of Seed Size,Germination and Seed Viability of Tropical Tree Species in Southern India1

Seed weight, days to germination and seed viability were observed for 99 species growing in the Western Ghats of Karnataka, India. Seed size was strongly correlated with days to germination; smaller seeds germinated faster than larger seeds. Species which flowered during the rainy season had lighter seeds than species which flowered during the dry season. It was also found that seed size and viability of seeds were related to the season of fruiting. Species which fruit during the rainy season had heavier seeds and shorter viability than species which fruit during the dry season. These flowering and fruiting patterns and varying seed sizes are argued to be adaptations to the time of dispersal, time of moisture availability in the habitat and seedling survival.     

Sources of variation in pollinator contribution within a guild: the effects of plant and pollinator factors

Among plants visited by many pollinator species, the relative contribution of each pollinator to plant reproduction is determined by variation in both pollinator and plant traits. Here we evaluate how pollinator movement among plants, apparent pollen carryover, ovule number, resource limitation of seed set, and pollen output affect variation in contribution of individual pollinator species to seed set in Lithophragma parviflorum (Saxifragaceae), a species visited by a broad spectrum of visitors, including beeflies, bees and a moth species. A previous study demonstrated differences among visitor species in their single-visit pollination efficacy but did not evaluate how differences in visitation patterns and pollen carryover affect pollinator efficacy. Incorporation of differential visitation patterns and pollen carryover effects —commonly cited as potentially important in evaluating pollinator guilds — had minor effects (0–0.6% change) on the estimates of relative contribution based on visit frequency and single-visit efficacy alone. Beeflies visited significantly more flowers per inflorescence than the bees and the moth. Seed set remained virtually constant during the first three visited flowers for beeflies and larger bees, indicating that apparent pollen carryover did not reduce per-visit efficacy of these taxa. In contrast, Greya moth visits showed a decrease in seed set by 55.4% and the smaller bees by 45.4% from first to second flower. The larger carryover effects in smaller bees and Greya were diminished in importance by their small overall contribution to seed set. Three variable plant traits may affect seed set: ovule number, resource limitation on seed maturation, and pollen output. Ovule number per flower declined strongly with later position within inflorescences. Numbers were much higher in first-year greenhouse-grown plants than in field populations, and differences increased during 3 years of study. Mean pollen count by position varied 7-fold among flowers; it paralleled ovule number variation, resulting in a relatively stable pollen:ovule ratio. Resource limitation of seed set increased strongly with later flowering, with seed set in hand-pollinated flowers ranging from 66% in early flowers to 0% in the last two flowers of all plants. Variation in ovule number and resource limitation of seed maturation jointly had a strong effect on the number of seeds per flower. Visitation to early flowers had the potential to cause more seed set than visitation to later flowers. Overall, the most important sources of variation to seed production contribution were differences among pollinators in abundance and absolute efficacy (ovules fertilized on a single visit) and potentially differential phenology among visitor species. These effects are likely to vary among populations and years.     

PATTERNS OF FRUIT PRODUCTION IN A NEOTROPICAL ORCHID: POLLINATOR VS. RESOURCE LIMITATION

Patterns of growth, and of flower and fruit production, were monitored over two years in a population of the orchid Aspasia principissa Reichb. f. in central Panama. Observations and experimental manipulations were used to determine the relative importance of pollinator and resource limitation on fruit production. Within a season, fruit production was limited by pollinator availability. Fruit set for hand-pollinated flowers was over six times greater than that for naturally pollinated flowers. However, in plants that produced more than one fruit, fruit size declined in subsequent fruits, indicating that resources could limit seed production within a season. Plants producing fruits in 1986, on average, produced smaller shoots and inflorescences in 1987 relative to plants that flowered but produced no fruit in 1986. Thus, plants are likely to be resource limited over their lifetimes. Most individuals of reproductive size (82.5%) did not produce fruit over a three-year period. The reproductive dominance of a few individuals in this population of Aspasia principissa may have important implications for understanding the population structure of the species and the high species diversity of orchids.     

FLOWERING,SEX RATIOS,POLLEN-OVULE RATIOS,FRUIT SET,AND REPRODUCTIVE EFFORT OF A DIOECIOUS TREE,MYRISTICA INSIPIDA (MYRISTICACEAE), IN TWO DIFFERENT RAIN FOREST COMMUNITIES

The flowering of Myristica insipida R. Br. was studied in two rain forest communities in northern Queensland. This dioecious, subcanopy tree had a male-biased sex ratio at both study sites. In the lowland population the male-bias could be attributed to males (trees producing staminate flowers) starting to flower at a smaller average size than females (trees producing pistillate flowers). There were no intersexual differences in spacing or distribution within the study sites. Males trees flowered earlier, flowered longer, and produced over twice as many flowers as females during the study season. Although the onset of flowering was rather variable, 18–22 days following heavy rains, most trees had a synchronous period of maximum flowering. Pollination manipulations determined that there was no fruit development without pollination, and that increasing pollen loads resulted in increased fruit set with diminishing effect. Taking into account the sex-ratios and intersexual differences in flower production, the pollen-ovule ratio was calculated to be 16,000–19,000. Male trees were found to expend more energy on flowering than female trees. Open-pollination resulted in 1.0% of female flowers setting fruit. The much greater cost of fruit production resulted in females expending 421% more energy on reproduction than males. Fruit and seed production were judged to be pollination-limited. Nonetheless, this species exhibited several characteristics that are predicted if dioecy evolved by means of sexual selection.     

Reproduction and development of the endangered Sedum integrifolium ssp. leedyi (Crassulaceae)

Information on reproduction and life history is important for the conservation of endangered plants. We investigated rates of flowering, seed set, and germination in populations of the endangered perennial plant Sedum integrifolium ssp. leedyi. Germination and flowering rates differed significantly among populations, but seed set rate did not. We assayed 26 plant clusters (81 stems) from four of the five known populations for evidence of clonal reproduction using 28 randomly amplified polymorphic DNA (RAPD) markers. Of the 81 stems, 75 had unique genotypes and three pairs had identical genotypes, suggesting that clonal reproduction is infrequent. Flowering, seed set, and germination rates were correlated with our estimates of ratios of effective to actual population sizes (Ne/N), but not with Ne. The single formally protected population may be experiencing inbreeding depression. We grew plants from seed to maturity in a greenhouse, with a germination rate of 77% and survival of 98% of the germinants at 6 mo, suggesting that this will be a viable means of ex situ propagation. Plants flowered 4-6 mo after germination and produced mature fruits 1-2 mo later, suggesting that they have the potential to sexually reproduce in their first or second season of growth.     

露蕊乌头（Aconitum gymnandrum Maxim）花序内位置依赖的性分配

植物会采取什么样的繁殖对策来保障它的雌性和雄性繁殖成功,特别是那些生活在高寒地区的植物,已经引起了很多学者的关注.通过野外盆栽试验对青藏高原东部高寒草甸上的常见物种露蕊乌头(Aconitum gymnandrum Maxim)花序内的性分配模式进行了研究,发现同其它顶向式(从花序基部到顶部)开花的植物一样,随着花从基部到顶部的次序,露蕊乌头的心皮数、雌蕊重、种子数和种子重/果实减少;而花药数,花粉数/胚珠数以及雄蕊重/雌蕊重增大,顶部花表现出偏雄的性分配.结籽率在不同的位置间没有显著差异,不受花粉数量的限制;去花实验没有增加剩余部位花的种子数,只是增加了种子大小,这是花序内不可避免的位置效应导致了花序内位置依赖的性分配模式.