Pattern of Flower and Fruit Production in Stryphnodendron adstringens, an Andromonoecious Legume Tree of Central Brazil

Abstract: Patterns of flower and fruit production in racemes of Stryphnodendron adstringens, an andromonoecious Brazilian savanna tree species, were studied in two natural areas near Uberlândia-MG. Racemes were divided in three parts: apex, centre, and base. Number of flowers, gender, and nectar and pollen production were analyzed for each section. Frequency of visitors to each part of the inflorescence was also quantified. Hand self- and cross-pollinations were performed in complete racemes and fruit set used to determine breeding system. The racemes produced a mean of 329 flowers, more densely packed in the central portion. Hermaphrodite and male flowers occur along the inflorescence but hermaphrodite flowers are more common in the centre. Fruit set was markedly low but does not seem to be limited by pollination service, since free open-pollinated racemes and hand cross-pollinated ones do not differ in fruit production rates. Fruits resulted mostly from cross-pollinated flowers and fruit production was biased to the central portion of the raceme. Nectar yield was higher in the central portion of the raceme and visitors arrived more commonly on this portion of the inflorescence. However, most flowers did not produce nectar. The pattern of fruit production seems to be a consequence of the hermaphrodite flower distribution in the raceme and it is not constrained by pollen flow or flower opening sequence.     

Apical pattern of fruit production in the racemes of Ceratonia siliqua (Leguminosae: Caesalpinioideae): role of pollinators

Fruit production and arrangement within the raceme were studied in two dioecious populations of Ceratonia siliqua (Leguminosae: Caesalpinioideae), an arboreal species that produces caulogenous racemes (emerging only from the old branches) with numerous flowers. Fruit production per raceme was low and similar between years and populations and even between individuals. During flowering, there were considerable flower losses from predation and lack of pollination. A mean of nine flowers per raceme began the transformation into fruits, of which 77% aborted. The final fruit production per raceme increased significantly following hand pollination, but was always very much lower than the availability of flowers in the raceme. The results suggest that fruit production of each raceme is limited by both availability of resources and a deficient pollination. In racemes setting fruit arrangement follows a definite pattern that remains constant between years and populations: fruit production was significantly higher in the apical zone of the raceme and lower in the basal zone. The pollinators of C. siliqua (flies and wasps) showed a clear preference for beginning their visits at the apex of a raceme. As a result, the pollen load deposited on the stigmas decreased from apex to base of the raceme. In most of the flowers situated in the central and basal zone of the raceme, the number of pollen grains deposited on their stigmas was lower than the number of their ovules. The high number of seeds in developed fruits suggests that the plant selectively aborts flowers that receive a smaller pollen load. The results indicate that the final pattern of fruit arrangement within the raceme is a direct result of pollinator activity.     

Gender variation of sequential inflorescences in a monoecious plant Sagittaria trifolia (Alismataceae)

In protogynous plants, female flowers of early blooming plants are at a reproductive disadvantage because they cannot set fruit due to the lack of available pollen. To study this phenomenon, gender expression of the monoecious herb Sagittaria trifolia was investigated over the entire flowering season in two field and two cultivated populations in Hubei and Hunan Provinces, China. In racemes of S. trifolia, flowers open sequentially from bottom to top, with female flowers opening first followed by male flowers. This creates a temporal separation of sexes in the species. Under field conditions small plants are often male, with production of both male and female flowers increasing with plant size. Femaleness increased among sequential inflorescences since female flower production increased whereas male flower production did not. Seed production was greater in large inflorescences because they contain more female flowers, and the number of ovules increased in female flowers at basal positions within the raceme. A consistent pattern of high seed set was observed in flowers from both field and cultivated populations. About 1 % of unfertilized ovules resulted from no pollination and 2 % of the seeds produced were only partly developed due to resource limitation. In the first inflorescence of the six experimental populations, 6.7-40.0 % of individuals produced only male flowers, and female flowers of 1.9-6.5 % individuals were aborted. The occurrence of male flowers in early blooming inflorescences could be an adaptive strategy to conserve resources and enhance pollination of female flowers in protogynous S. trifolia.     

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。     

Effects of number of flowers per raceme and number of racemes per plant on bumblebee visits and female reproductive success in <Emphasis Type="Italic">Salvia nipponica</Emphasis> (Labiatae)

We studied the adaptive arrangement of raceme flowers in dense and sparse habitats of Salvia nipponica. We recorded bumblebee visits and collected mature seeds to estimate outcrossing rates from allozyme genotypes. To examine the resource investment in flowering stalks, we measured their length and dry mass. We found that a greater number of open flowers in a raceme enhanced bumblebee visits and successive probings in both density plots. However, greater height of a raceme did not enhance these in plots of either density. In the high-density plot, both a greater mean number of open flowers in a raceme and a greater number of flowering racemes on a plant enhanced bumblebee visits, successive probings, and also enhanced outcrossing rate in spite of an increase in successive probings. Although the number of flowers per raceme had a greater positive effect on seed-to-ovule ratios than did the number of racemes, the number of flowers per raceme may be constrained because a longer flowering stalk needs greater buckling strength. Hence, it may be advantageous for plants in high-density plots to increase both number of flowers per raceme and number of racemes. In the low-density plot, a greater number of flowering racemes on a plant enhanced pollinator visits, but a greater mean number of open flowers in a raceme did not. Hence, it may be advantageous for plants to increase the number of racemes while keeping the numbers of flowers per raceme small. Thus, the adaptive number of flowers per raceme and number of racemes per plant may differ between high- and low-density plots.     

Manipulation of bee behavior by inflorescence architecture and its consequences for plant mating

Angiosperms display flowers in many three-dimensional arrangements, but the functional significance of this diversity is largely unknown. We examined influences of inflorescence architecture on pollination and mating by quantifying the responses of bumblebees to three architectures and then using these observations as the basis of a model that simulated pollen dispersal. On artificial panicles, racemes, and umbels, each with 12 identical flowers, bees visited one more flower, on average, on umbels than on panicles (with racemes being intermediate). In contrast to this weak response, the consistency of foraging paths among flowers differed strongly among architectures (raceme > panicle > umbel). The simulation model revealed limited differences in self-pollination and pollen export among the three inflorescence designs when all flowers presented and received pollen, because mating differences depended on only the number of flowers visited. In contrast, in simulations of inflorescences on which pollen receipt and presentation were segregated so as to minimize interference among flowers, the consistency of movement paths governed mating. In this case, racemes self-pollinated much less than umbels (with panicles being intermediate), and racemes exported much more pollen than umbels and panicles. These effects have diverse consequences for the evolution of inflorescence architecture, flower design, and sexual segregation.     

Factors affecting regulation of maternal investment in an indeterminate flowering plant (Cercidium microphyllum; Fabaceae)

On palo verde trees, nearly 80% of potential offspring were lost during flower bud development. Flower buds at the base of racemes developed earliest and were more likely to survive depredations of gelechiid larvae or abortion from putative limited resources. Herbivory accounted for greater cumulative losses than putative resources; however, an herbivore exclusion experiment suggested that buds damaged by herbivores would have aborted anyway without damage. In a natural experiment, plants that received water runoff had significantly higher densities of racemes, with only slight increases in numbers of buds, flowers, or pods per raceme. Similarly, thinning racemes experimentally to assess limited resources among potentially competing racemes increased the number of viable flower buds per raceme only slightly, with negligible herbivore damage. The production of more racemes rather than more reproductive structures per raceme may be a mechanism to allocate limited resources more efficiently. Alternatively, in studies with animals, similar patterns in the production of broods are thought to be mechanisms to avoid nest predation. Thus, while putative limited resources and resource allocation patterns reduced the proximate effects of larvae, herbivory must be considered as a possible ultimate factor in the patterns observed here because limited resources may be allocated in ways to reduce herbivory.     

Osmotic stress at seedling stage impairs reproductive development in buckwheat (Fagopyrum esculentum)

Common buckwheat ( Fagopyrum esculentum Moench) is highly sensitive to drought. Many studies have investigated the impact of water stresses, applied late during plant development, on seed production by this non-Poaceae cereal but information concerning the effects of stresses occurring early during plant's life is essentially lacking. The aim of the present work was to define which yield-related parameters are more affected by a 4-day water constraint, applied at three different developmental stages, by transferring buckwheat plants of the cultivar 'La Harpe' grown in a hydroponic system to a medium supplemented with polyethylene glycol. Stresses during the vegetative phase, when the shoot apical meristem (SAM) was still initiating leaves, or later, when its activity was stopped after production of approximately 10 inflorescences, were less detrimental than an osmotic constraint applied to plants recently engaged in the reproductive phase, with SAM having initiated six to seven inflorescences. The number of cymes and flowers per inflorescence and the number of pollen grains per anther were decreased. The production of seeds in the first inflorescence was also reduced but this reduction paralleled the decrease in cyme number, suggesting that seed set, per se, was not affected. Pollen viability, female fertility and weight of seeds reaching maturity were not altered by the water deficit. The present work clearly identified two processes that, in buckwheat, are particularly sensitive to an osmotic stress: (1) the activity of reproductive meristems involved in the production of cymes and flowers and (2) male sporogenesis.     

Reproductive patterns within racemes in protandrous Aconitum gymnandrum (Ranunculaceae): potential mechanism and among-family variation

The adaptive significance and mechanism of patterns in floral sex allocation and female success within inflorescences has attracted attention recently, whereas few studies have examined genetic variation of intra-inflorescence pattern. The purpose of this study is to investigate patterns of reproduction within racemes in protandrous Aconitum gymnandrum Maxim., and illuminate potential mechanisms and genetic variation of such patterns. Data from pot experiment on 40 maternal families were collected in field. Anther number, pollen:ovule ratio and seed germination rate increased from bottom to top flowers within racemes, but other traits, such as gynoecium mass, carpel number, sepal galea height and seed production decreased significantly with flowering sequence. Variation in floral sex allocation within racemes in A. gymnandrum fitted entirely the prediction of protandry, which was not a result of architectural effect. Such selected pattern may result from a variety of factors influencing the mating environment, such as pollinator directionality, display size and flower longevity. Decline of female success within racemes in A. gymnandrum also resulted from male-biased allocation selected by variation in the mating environment, not resource competition or pollen limitation. Moreover, there was genetic variation for most reproductive traits and the position effect, as evinced by significant variation among families.     

Plasticity of floral sex allocation within inflorescences of hermaphrodite Aconitum gymnandrum

Aims Sex allocation in plants is often plastic, enabling individuals to adjust to variable environments. However, the predicted male-biased sex allocation in response to low resource conditions has rarely been experimentally tested in hermaphroditic plants. In particular, it is unknown whether distal flowers in linear inflorescences show a larger shift to male allocation relative to basal flowers when resources are reduced. In this study, we measure position-dependent plasticity of floral sex allocation within racemes of Aconitum gymnandrum in response to reduced resource availability.Methods Using a defoliation treatment in the field applied to potted plants from a nested half-sibling design, we examined the effects of the treatment, flower position, family and their interactions.Important findings Allocation to male function increased with more distal flower position, while female allocation either did not change with position or declined at the most distal flowers. Defoliation significantly reduced the mass of both the androecium and gynoecium, but not anther number or carpel number. Gynoecial mass declined more strongly with defoliation than did androecial mass, resulting in a significant increase in the androecium/gynoecium ratio as predicted by sex allocation theory. Plastic responses of androecium mass and gynoecium mass were affected by flower position, with less mass lost in basal flowers, but similar plastic magnitude in both sexual traits across flower position lead to consistent variation in the androecium/gynoecium ratio along the inflorescence. A significant treatment*paternal family interaction for the androecium/gynoecium ratio is evidence for additive genetic variation for plastic floral sex allocation, which means that further evolution of allocation can occur.     

Female reproductive output in a Mediterranean shrub: effects from inflorescence to population

Inflorescence effects have been poorly studied, in spite of the functional relevance of the inflorescence in fruit and seed ecology. The present study focused on the effects of inflorescence size and phenology, and flower position within the inflorescence, in relation to fruit and seed production of the Mediterranean shrub Ononis fruticosa. Variability in fruit and seed production, seed weight and germination were estimated and modelled. Results confirmed that the most important predictors in seed production were inflorescence flowering time and flower position within the inflorescence. Thus, the number of mature seeds per fruit was higher in earlier inflorescences and in basal positions. On the other hand, predation was higher in fruits in basal positions. In fact, seed predation seemed to be the most important factor controlling final seed production. Models at the plant level suggested a negative incidence of geitonogamous pollination and resource limitation, which were also observed at the fruit level. This study confirmed the relevance of inflorescence effects on the reproductive output of O. fruticosa. Although the underlying processes could not be identified, our results provide several hypotheses for future experimental studies.     

Flowering phenology and reproductive success of native shrub Berberis darwinii Hook. along different light environments

Studies evaluating flowering phenology and reproductive success are necessary when we want to direct a domestication project in a species with a potential productive value. We studied flowering phenology and reproductive success of Berberis darwinii growing under different light conditions in its native distribution area in the Andean Patagonian forests of Argentina. We test the hypothesis that plants grown under conditions of high-light availability exhibit advanced phenology and higher reproductive success than those grown under conditions of lower light availability. Phenology and reproductive success were determined in three contrasting light conditions at two forest sites, which were, canopy, gap and forest edge. Plants did not bloom under the forest canopy. Flowering and fruiting period lengths were similar in both sites and light conditions of gap and forest edge during spring and summer. Although gap plants had more racemes per shoot, racemes of edge plants had more flowers, fruits and a higher proportion of flowers producing ripe fruit. We show that B. darwinii reproduction studied in the Andean Patagonian forests is conditioned by the canopy openness. Regarding reproductive success, edge plants invest less resources in flower production than gap plants to have similar fruit production.     

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.     

Fertility of Vriesea gigantea Gaud. (Bromeliaceae) in southern Brazil

Plant fertility is a central subject of many questions in plant evolutionary and conservation biology. Pollen availability, abiotic resources, and flowering pattern can limit fruit and seed production. Open pollination and pollen supplementation studies are used to estimate any pollen limitation in natural populations. To study the impact of these factors on the reproductive success of Vriesea gigantea, an epiphytic bromeliad in the Atlantic Rainforest in Brazil, its fertility in four natural populations in Itapu? State Park was assessed by considering plant and inflorescence size, flower production, fruit and seed set, flower and fruit set pattern, and seed viability and germination rate. Supplemental pollination in adult plants was used to determine whether fruit production in V. gigantea is limited by reception of pollen. The results showed that V. gigantea has a high production of flowers, fruits, and seeds. Seeds are highly viable in all populations, presenting an average germination rate of 94% (SE ± 3.5). Plants of V. gigantea from Itapu? State Park are highly fertile. The high proportion of fruit and seed set after manual hand pollination indicates that the species is self-compatible. Pollination treatments showed evidence of pollinator limitation in the Itapu? State Park population.     

A Top-Down Hierarchy in Fruit Set on Inflorescences in Iris fulva (Iridaceae)

Abstract: In Iris fulva, the apical flower on an inflorescence opens first, and the flowering sequence then proceeds from the most basal flower upward to the apex (acropetally). Flowering order and flower position are thus partially decoupled, and this provides an opportunity to separately investigate the effects of these two factors upon fruit formation. We recorded natural patterns of fruit set and seed production, and found that fruit set patterns were determined by what appears to be a form of apical dominance. The first, apical flower had the highest fruit set and seed production. Fruit set decreased towards the base of the inflorescence, with later-opening flowers having a higher fruit set. This is contrary to the pattern usually observed in other acropetally flowering plants, which is a higher fruit set for basal, early flowers. By performing additional hand pollinations and counting pollen grains on naturally pollinated flowers, we found that pollen deposition was not a major factor limiting fruit set, and that it could not explain the large difference in fruit set between the apical and basal flowers. Removing the first flower after it had wilted increased fruit set in the remaining flowers, but mainly in the more apical flowers. Only by removing the two topmost flowers could we obtain an increase in fruit set for the basal flower. The fact that the basal flower rarely sets fruit, despite being closer to resources, suggests that the apical meristem is either a strong sink for resources or produces hormones that form a gradient along the inflorescence, which is comparable with apical dominance.     

Fruit Abscission in Cowpea, Vigna unguiculata (L.) Walp: I. DISTRIBUTION OF 14C-ASSIMILATES IN THE INFLORESCENCE, AND COMPARATIVE GROWTH OF OVARIES FROM PERSISTING AND ABSCISING OPEN FLOWERS

One of the hypotheses proposed for the abscission of flowersand fruits from the cowpea was that the oldest fruits at raceme1 on the peduncle monopolize the mobile nutrients availableto the whole inflorescence. Consequently, the distal young fruitsand flowers starve, abort, and are shed. This hypothesis wastested through autoradiographic studies of the distributionof ¹⁴C-assimilates on the cowpea inflorescence and the comparativegrowth of ovaries from persisting and abscising open flowersand young fruits. ¹⁴C-asaimilates were distributed to the fruits at raceme 1 andto flower buds, open flowers, and young fruits distal to raceme1. Therefore, the hypothesis that the fruits at raceme 1 monopolizethe mobile assimilates cannot be maintained. At the time of opening the persisting flowers of raceme 1 hadheavier ovaries than the aborting flowers of raceme 3 althoughboth types of flowers were visually alike. During the initial12 h following opening, the difference in ovary size was maintainedand often accentuated. The ovary of the persisting flower increasedin both fresh and dry weights. The growth of the ovary of theaborting flower was variable; the fresh weight often decreased,although it also increased in some experiments; the dry weightremained unchanged or decreased or increased remarkably. The availability of nutrients was considered to be of secondaryimportance in the chain of events leading to fruit abscission.Other internal factors which may control vital processes relatedto embryo development appear to be of primary, underlying importance.     

Combined effects of inflorescence architecture, display size, plant density and empty flowers on bumble bee behaviour: experimental study with artificial inflorescences

Pollen dispersal by pollinators is governed by the extent to which diverse effects on pollinator behaviour act independently or augment or moderate each other. Using artificial inflorescences, we assessed the behavioural responses of bumble bees to inflorescence architecture (raceme, panicle, and umbel), inflorescence size (7 or 13 flowers), inter-inflorescence distance and the proportion of empty flowers per inflorescence. The advantage of large inflorescences in terms of attractiveness was larger for racemes and umbels than for panicles, whereas the effect of inter-inflorescence distance on the number of successive probes was smaller for racemes than for panicles and umbels. The number of flowers probed per visit increased almost proportionally with display size when fewer flowers were empty, whereas the number increased less when many flowers were empty. Our results suggest that display size and the spatial arrangement of flowers and nectar within inflorescences can contribute to efficient pollination by affecting pollinator behaviour interactively.     

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.     

The effect of flower position on male and female reproductive success in a deceptively pollinated tropical orchid

In many plants, including orchids, differential fruit set along the inflorescence has been attributed to pollinator behaviour. For instance, the pollinator, moving up the inflorescence, becomes satiated with the resources and leaves before visiting the upper flowers. Consequently, the pollinators do not visit flowers as frequently higher up the inflorescence. Alternatively, flower size may vary along the inflorescence, making pollination ineffective as flowers decrease in size. I tested for the presence of differential pollination along the inflorescence in a pollinator-limited tropical epiphyte, Lepanthes rupestris Stimson, and determined the likely cause of the observed pattern. As this species has inflorescences with sequential flowering, pollinator behaviour, moving up the inflorescence as in synchronous multiflowering inflorescences, can be discounted as an explanation for differential fruit set. Fruit set is shown to be more frequent at the base of the inflorescence, but male reproductive success through pollinarium removal is basically independent of flower position. Moreover, cross-pollination by hand at variable flower positions along the inflorescence results in equal fruit set, suggesting that resources are not limiting and cannot explain the cause of differential fruit production along the inflorescence in natural populations. Furthermore, flower size is shown to diminish along the inflorescence, suggesting that the pollinator(s) may be ineffective at depositing the pollinarium in the smaller higher flowers. Consequently, pollinator behaviour and its interaction with flower size, and not resource limitation, is likely to be the main cause of differential fruit set along the inflorescence in L. rupestris .  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 405–410.