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Background and Aims: Abiotic and biotic components of the environment both limit plant reproduction, but how they interact with one another in combination is less understood. Understanding these interactions is especially relevant because abiotic and biotic environmental components respond differently to various drivers of global change. Here, we aim to understand whether the effects of pollination (biotic component) on plant reproduction depend on soil moisture (abiotic component), two factors known to affect plant reproduction and that are changing with global change., Methods: We conducted pollen supplementation experiments for two plant species, Delphinium nuttallianum and Hydrophyllum fendleri, in subalpine meadows in the Western USA across 4 years that varied in soil moisture. In a separate 1-year field experiment, we crossed water addition with pollen supplementation factorially. We measured the proportion of fruit set, seeds per fruit and seeds per plant, in addition to stomatal conductance, to determine whether plant physiology responded to watering., Key Results: In the 4-year study, only H. fendleri reproduction was pollen limited, and this occurred independently of soil moisture. Experimental water addition significantly increased soil moisture and stomatal conductance for both species. The effect of pollen addition on reproduction depended on the watering treatment only for H. fendleri fruit production. Reproduction in D. nuttallianum was not significantly affected by pollen addition or water addition, but it did respond to interannual variation in soil moisture., Conclusions: Although we found some evidence for the effect of a biotic interaction depending on abiotic conditions, it was only for one aspect of reproduction in one species, and it was in an unexpected direction. Our work highlights interactions between the abiotic and biotic components of the environment as an area of further research for improving our understanding of how plant reproduction responds to global change., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

Herbivores decrease plant fitness by consuming reproductive tissues, limiting resources, and/or affecting mutualisms. Although these mechanisms were extensively tested in chewing herbivores, the impact of other functional groups (e.g., sap-feeders) remains poorly understood. We investigated whether aphids affect plant reproduction via direct resource limitation on seed production and/or pollination interference. We compared plant traits and the seed set of naturally aphid-free vs. aphid-infested plants and then manipulated aphid presence and pollen receipt. We used path models to examine the links between variables. Nectar volume and seed set of aphid-infested plants was 54% and 42% lower than that of aphid-free plants. 72 h after removing aphids, nectar volume was restored to the level of aphid-free plants. When pollinators were excluded, the seed set of aphid-infested and aphid-free plants did not differ, suggesting that direct resource limitation on seed production was not the cause of reduced plant fitness. Manual addition of pollen restored the seed set of aphid-infested plants to the level of aphid-free plants, evidencing that plants were pollen limited. The path analysis showed a negative link between aphids and the seed set via nectar volume, supporting that nectar shortage caused by aphids may interfere with pollination and reduce plant fitness. Since aphids are crop pests and feed on a large number of animal-pollinated plants, the potential of these insects to influence pollination and plant fitness is high. This study emphasizes the ecological importance of aphids and the need to better understand the links between sap-feeding herbivory, pollination, and plant fitness.

Background: A large proportion of the flowers and ovules of plants do not develop into fruits and seeds. Plant reproduction may be limited because of pollen limitation and resource limitation. Medicago sativa L. is an ecologically important species in northwest China. We conducted a pollen supplementation experiment to determine the degree of pollen limitation in this species and detect the possible effects of resource allocation on pollen supplementation. We crossed two factors, pollen level (natural condition and hand pollinated) and resource level (control, water added, and fertilizer added), to estimate the effects of pollen addition and resource limitation on the opening of flowers and seed set. We also analyzed the floral characters, visitation frequency of pollinators and pollinator activity to estimate the effect of pollinators on the reproduction of M. sativa., Results: Our results indicated that addition of pollen to some flowers did not divert resources from other flowers and that the addition of pollen boosted the seed set per flower, with no effect on flower number. The primary effect of resource limitation was on the number of flowers produced; however, there was no significant effect on seed set per flower. These findings showed that pollen limitation was an important limiting factor for seed set. In addition, Andrena lebedevi Popov was identified as the most effective pollinator, and pollinator visiting and activity affected reproduction success in M. sativa., Conclusions: We found outcrossing was dominant in the breeding system and insect pollination played an important role in outcrossing. These findings have identified the dominant factor influencing seed set of M. sativa. This study aspires to contribute to a better understanding of pollen limitation, resource limitation and reproductive success.

Premise of the Study: Understanding resource allocation to reproduction, a key factor in life history tradeoffs, has long intrigued plant ecologists. Despite the recognized importance of understanding the movement of resources among flowers following variable pollination, the patterns of resource reallocation to plant reproductive organs have not been thoroughly addressed. In this study, we aimed to empirically explore how resources redistribute within inflorescences in response to differential pollination intensities., Methods: Using a common herb, Sagittaria trifolia, we conducted supplemental and controlled pollination for single, some, or all flowers in simple and complex inflorescences, and compared their resulting fruiting probabilities, seed production, and average seed masses., Key Results: Pollen supplementation of a single flower significantly increased its fruiting probability; however, the same manipulation of an inflorescence did not increase its overall reproduction. Single pollen-supplemented flowers had a higher percentage fruit set than inflorescences receiving supplemental pollination. In complex inflorescences, supplemental pollination had no effect on the reproductive success of flowers on the lateral or main branches., Conclusions: We provided evidence of resource reallocation from controlled to pollen-supplemented flowers in simple inflorescences; however, resources were unlikely to be reallocated between the main and lateral branches in the complex inflorescences, suggesting that flowering branches represent integrated physiological units in S. trifolia. The results also demonstrated that single-flower supplemental pollination would exaggerate pollen limitation and lead to a biased understanding of a plant's reproductive status., (© 2018 Botanical Society of America.)

Identifying the resources that limit bee populations is essential for both bee conservation and pollination management in agroecosystems. Land‐use change typically leads to decreased habitat availability for wild pollinators, including loss of nesting habitat, which is an essential but often‐overlooked resource for wild bees. Cavity‐nesting bees, such as many Osmia spp. (Hymenoptera: Megachilidae), occupy holes in wood or reeds to build their nests; due to their nesting habits, they are frequently scarce in agricultural settings. Nevertheless, under the right circumstances, these bees can be ideal pollinators of apple and other orchard crops. Artificial nesting structures ('bee hotels' or 'trap‐nests') are often used to study cavity‐nesting bees and have been proposed as tools for bee conservation.To evaluate the effects of additional nesting habitat on the local abundance of pollinators, we selected 24 sites in apple orchards in eastern Canada in 2021 and 2022. Each site comprised two study plots: one in which we installed artificial nesting structures for cavity‐nesting bees, and one without added nesting habitat (control).Pollinator surveys were conducted in both plot types to measure pollinator visits to apple blossoms and, after apple bloom, to flowers in the undergrowth and/or in shrubs. Numbers and size of developing fruit were also recorded. Both Osmia bees and overall pollinator numbers were significantly higher in the treatment with trap‐nests (respectively, 33% and 22% higher during the 2021 growing season, and 113% and 30% higher during apple bloom across the 2 years of the study); however, there was no consistent difference in apple size or fruit set between the two treatments. Thus, trap‐nests locally increased the numbers of cavity‐nesting bees as well as total pollinator numbers, but they had little effect on apple yield, likely because apple production was not pollinator limited in the years of this study.Synthesis and applications. These findings suggest that bee populations in apple orchards can be limited by nesting resources such that addition of constructed nesting habitat could be an effective means of increasing pollinator numbers in orchards, and potentially, supporting bee conservation. [ABSTRACT FROM AUTHOR]

Premise: Understanding the factors that limit reproductive success is a key component of plant biology. Carnivorous plants rely on insects as both nutrient sources and pollinators, providing a unique system for studying the effects of both resource and pollen limitation on plant reproduction. Methods: We conducted a field experiment using wild‐growing Dionaea muscipula J. Ellis (Droseraceae) in which we manipulated prey and pollen in a factorial design and measured flower production, number of fruits, and number of seeds. Because understanding reproduction requires knowledge of a plant species' reproductive and pollination biology, we also examined the pollination system, per‐visit pollinator effectiveness, and pollen‐ovule (P/O) ratio of D. muscipula. Results: Plants that received supplemental prey produced more flowers than control plants. They also had a higher overall fitness estimate (number of flowers × fruit set (total fruits/total flowers) × seeds per fruit), although this benefit was significant only when prey supplementation occurred in the previous growing season. Neither pollen supplementation nor the interaction between pollen and prey supplementation significantly affected overall plant fitness. Conclusions: This study reinforces the reliance of D. muscipula on adequate prey capture for flower, fruit, and seed production and a mobile pollen vector for reproduction, indicating the importance of considering insects as part of an effective conservation management plan for this species. [ABSTRACT FROM AUTHOR]

Understanding whether and how resource limitation alters phenotypic selection on floral traits is key to predict the evolution of plant–pollinator interactions under climate change. Two important resources predicted to decline with our changing climate are pollinators and water in the form of increased droughts. Most work, however, has studied these selective agents separately and in the case of water deficit, studies are rare. Here, we use the common morning glory (Ipomoea purpurea) to investigate the effects of experimental reduction in pollinator access and water availability on floral signals and nectar rewards and their effects on phenotypic selection on these traits. We conducted a manipulative experiment in a common garden, where we grew plants in three treatments: (1) pollinator restriction, (2) water reduction and (3) unmanipulated control. Plants in pollinator restriction and control treatments were well-watered compared to water deficit. We found that in contrast to pollinator restriction, water deficit had strong effects altering floral signals and nectar rewards but also differed in the direction and strength of selection on these traits compared to control plants. Water deficit increased the opportunity for selection, and selection in this treatment favoured lower nectar volumes and larger floral sizes, which might further alter pollinator visitation. In addition, well-watered plants, both in control and pollinator deficit, showed similar patterns of selection to increase nectar volume suggesting non-pollinator-mediated selection on nectar. Our study shows that floral traits may evolve in response to reduction in water access faster than to declines in pollinators and reinforces that abiotic factors can be important agents of selection for floral traits. Although only few experimental selection studies have manipulated access to biotic and abiotic resources, our results suggest that this approach is key for understanding how pollination systems may evolve under climate change. [ABSTRACT FROM AUTHOR]

In habitats where resource availability declines during the growing season, selection may favor early-flowering individuals. Under such ephemerally favorable conditions, late-blooming species (and individuals) may be particularly vulnerable to resource limitation of seed production. In California, a region prone to seasonal drought, members of the annual genus Clarkia are among the last to flower in the spring. We compared pollen limitation (PL) of seed set and outcrossing rates between early- and late-flowering individuals in two mixed-mating Clarkia taxa to detect whether flowering time is associated with changes in seed set due to resource depletion, PL, or increased selfing. In 2008-2010, we hand-pollinated one flower on a total of 1855 individual plants either Early (near the onset of flowering) or Late (near the end of flowering) in the flowering season and compared seed set to adjacent, open-pollinated flowers on the same stem. To assess the contribution of pollen quality to reproduction, we first (2008) used allozymes to estimate outcrossing rates of seeds produced by Early and Late open-pollinated flowers. Second (2009), we conducted an anther-removal experiment to estimate self-pollen deposition. Seed set in Clarkia unguiculata was not pollen-limited. Clarkia xantiana ssp. xantiana was pollen-limited in 2008 and 2010, but not 2009. PL did not differ between Early and Late treatments. In both taxa, seed set of Early flowers was greater than Late flowers, but not due to PL in the latter. Reproduction was generally pollinator-dependent. Most pollen deposition was xenogamous, and outcrossing rates were >0.7 - and similar between Early and Late periods. These results suggest that pollen receipt and pollen quality remain seasonally consistent. By contrast, the resources necessary to provision seeds decline, reducing the fitness benefits associated with resource allocation to ovules.

The transition from outcrossing to selfing is a common evolutionary trend in flowering plants, and floral traits change significantly with the evolution of selfing. Whether or not plant traits are subjected to selection remains an open question in species with mixed mating systems. We examined phenotypic selection in two populations of Halenia elliptica with different selfing rates. We found that the pollen–ovule ratio, seed size, plant height, spur length, and pollinator visitation rate in the population with the higher selfing rate were lower than those in the population with the lower selfing rate. Selfing provides reproductive assurance for populations when pollinator service is low, and the floral traits that are associated with selfing syndrome are evident in populations with a higher selfing rate but are subjected to weak selection in each of the two populations with different selfing rates. Directional selection for an early flowering time indicated that late blooming flowers could experience a risk of seed development in alpine environments, and for large plants, selection indicated that seed production could be limited by the available resources. The floral traits that are associated with pollinator attraction and specialization could be subjected to weak selection at the plant level as selfing evolves, and the selective pressures that are independent of pollinators might not change significantly; highlighting the selective biotic and abiotic pressures that shape the morphological traits of plant species and their independence from the mating system. [ABSTRACT FROM AUTHOR]

Seed production can be affected by water availability and also depend on the amount (pollen intensity) and quality of pollen deposited. The way pollen receipt on the stigma translates into seeds produced follows that of a saturating dose–response. Not only can water availability and pollen intensity each influence seed production, these factors could interact in their effects on seed production. Changes to the relationship between seed production and pollen intensity can in turn influence pollinator effectiveness and pollinator-mediated selection. We asked how water availability affected indices of plant fitness (seed set, fruit set and seed mass) and the relationship between pollen intensity and seed production in Phacelia parryi. We conducted a greenhouse experiment where we manipulated water availability (either high- or low-water) to pollen recipient plants and hand-pollinated each plant with a range of pollen intensities. We conducted 703 hand-pollinations on 30 plants. For each hand-pollinated flower we measured pollen deposited, seed production and seed mass. We then generated a piecewise regression of the relationship between pollen intensity and seed production, and determined average effects of water on plant fitness measures. This experiment was paired with a field observational study aimed to document natural variation in pollen deposition. Average seed production per fruit was 21 % higher in the high-watered plants. The relationship between pollen intensity and seed production differed between the two water treatments. Plants under high-water exhibited a wider range in which pollen deposition increased seed production. Average natural pollen intensities fell within different regions of the piecewise regression for low- and high-water plants. Water availability can alter the efficiency by which pollen received is translated into seeds produced. Our greenhouse data suggest that only under certain pollen intensity environments will water availability affect how pollen received is translated into seeds produced. [ABSTRACT FROM AUTHOR]

Limitations on pollen and resources may significantly affect plant reproduction in fragmented habitats. In this study, phenology and pollinator frequency and activity were investigated to estimate the role of pollinators in Zygophyllum xanthoxylum reproduction, and this species is ecologically important in northwest China. In addition, the relative impact of restrictive amounts of pollen and resources on the seed set per flower was evaluated. It was found that adding pollen boosted the size of the seed set per flower, but had no significant effect on the number of flowers. By contrast, the addition of resources increased flower numbers as well as had a slight impact on the seed set per flower. These results indicate the amount of available pollen is a limiting factor for reproductive success. Moreover, Apis mellifera was identified as the most effective pollinator of Z. xanthoxylum, and there were more overall pollinators and visitations in the control than in the fragmented habitats. Furthermore, the limitations in pollen were more restrictive in the fragmented area than in the control. This was due to increased pollinator visitations in the control that could ameliorate the effects of lower pollen levels. When there is a limited availability of suitable pollinators, self-pollination is critical in fragmented habitats. Z. xanthoxylum has reproductive strategies that aid in adapting to harsh environments, including protogyny and delayed selfing. [ABSTRACT FROM AUTHOR]

Plant height is an important trait for plant reproductive success. Plant height is often under pollinator-mediated selection, and has been shown to be correlated with various other traits. However, few studies have examined the evolutionary trajectory of plant height under selection and the pleiotropic effects of plant height evolution. We conducted a bi-directional artificial selection experiment on plant height with fast cycling Brassica rapa plants to estimate its heritability and genetic correlations, and to reveal evolutionary responses to artificial selection on height and various correlated traits. With the divergent lines obtained through artificial selection, we subsequently conducted pollinator-choice assays and investigated resource limitation of fruit production. We found that plant height variation is strongly genetically controlled (with a realized heritability of 41-59%). Thus, plant height can evolve rapidly under phenotypic selection. In addition, we found remarkable pleiotropic effects in phenology, morphology, floral scent, color, nectar and leaf glucosinolates. Most traits were increased in tall-line plants, but flower size, UV reflection and glucosinolates were decreased, indicating potential trade-offs. Pollinators preferred plants of the tall selection lines over the short selection lines in both greenhouse experiments with bumblebees and field experiment with natural pollinators. We did not detect any differences in resource limitation between plants of the different selection lines. Overall, our study predicts that increased height should evolve under positive pollinator-mediated directional selection with potential trade-offs in floral signals and herbivore defense. [ABSTRACT FROM AUTHOR]

Female reproductive success is predicted to be simultaneously limited by the availability of pollen and resources. Selection on floral traits results from both factors, but their relative importance and interaction is poorly understood., We increased nutrient and pollen availability of the orchid Dactylorhiza lapponica in a factorial experiment to quantify resource- and pollinator-mediated selection on floral traits. Hand-pollination increased female fitness (number of fruits × mean fruit mass) by 74% in both nutrient treatments, whereas nutrient addition did not significantly affect female fitness., There was selection for more flowers and longer spurs, and selection on spur length was significantly pollinator-mediated and of similar strength across nutrient treatments (Δβpoll = 0·54 and Δβpoll_ NPK = 0·59). There was no statistically significant resource-mediated selection. Nutrient addition increased flower size the following year, but did not affect flower or fruit production, or selection on any trait., The results demonstrate that D. lapponica does not increase flower production in response to nutrient addition, that the increase in female fitness in response to hand-pollination is not resource limited, and suggest that natural resource variation does not influence selection on floral traits. The study illustrates that crossed manipulations of pollen and resources can clarify their relative importance for selection on floral traits., A is available for this article. [ABSTRACT FROM AUTHOR]

Masting, the synchronized production of variable seed crops, is widespread among woody plants, but there is no consensus about the underlying proximate mechanisms. To understand this population-level behaviour, it is necessary to dissect the behaviour of individual trees as well as the interactions that synchronize them., Here, we test a model of masting in which variability in seed set is driven by resource limitation within trees and synchrony is driven by pollen limitation due to phenological asynchrony in some years., We used a 35-year seed set data set and a 12-year phenological data set to analyse seed production of 84 valley oaks ( Quercus lobata) in central coastal California. Individual trees varied tremendously in their seed production patterns; trees with high levels of seed production were less variable over time, but showed stronger negative autocorrelation between years, suggesting that they are more resource-limited than unproductive trees. In years of more asynchronous flowering, Q. lobata produced fewer seeds, consistent with the importance of phenological synchrony., We parametrized a model with these results to investigate how individual resource limitation and population-wide pollen limitation - a consequence of asynchronous flowering during cold spring temperatures - interact to shape annual variation in seed production. The model illustrates that this proximate abiotic driver can synchronize the behaviour of individuals, resulting in population-wide seed production patterns that closely resemble the field data., Synthesis. Our findings support the hypothesis that an interaction between two proximate mechanisms, individual resource limitation and environmental variation affecting population-wide pollen availability, drives masting in this population of Quercus lobata. This combination of internal and external proximate drivers may underlie masting behaviour in many wind-pollinated plants. [ABSTRACT FROM AUTHOR]

Talbotiella gentiiis a critically endangered caesalpiniaceous legume endemic to Ghana, restricted in distribution to the margins of the forest zone where there has been extensive fire damage in recent decades. It occurs as isolated, near-monospecific populations, some of which show little seedling regeneration. We studied its reproduction to determine limitations that might inform efforts to prevent its extinction. We report on flowering phenology, pollen dispersal and germination, fruit and seed set, seed size and germination, and seedling survival, using controlled and open pollination.Talbotiella gentiiflowers profusely in most years. Although the flowers are coloured and scented, no animal pollinators were observed but pollen can be dispersed by wind. In open pollination, the number of pollen grains on stigmas and their germination was sufficient to fertilise the three ovules, despite competition from heavy stigmatic infestation by fungal spores and hyphae. Fruit set and seed set were low due to high flower losses and fruit abortion during development but ripe seeds showed high germination. Controlled pollination experiments (self, supplemented self, and intra- and inter-population crosses) showed that, while the species is self-compatible, selfing was the least successful treatment in all of the above steps of reproduction, whereas cross-pollination enhanced most steps, the more so for inter-population crosses. Seed size was notably greater in inter-population crosses and was positively correlated with genetic distance of the cross. The larger seeds associated with cross-pollination showed higher germination and seedling survival in the nursery. We conclude that limitations to natural reproduction inT. gentiiare principally due to processes between pollen germination and fruit and seed set, including pollen quality (low genetic diversity and inbreeding), fertilisation rates, flower and fruit abortion, and resource limitation. Low seed size in open and self pollination may have implications for seedling establishment. Species fitness might be improved by introducing other genotypes to existing populations, butin situprotection of the populations is essential in the medium term. [ABSTRACT FROM PUBLISHER]

Variation in resource availability can select for traits that reduce the negative impacts of this variability on mean fitness. Such selection may be particularly potent for seed production in flowering plants, as they often experience variation in pollen receipt among individuals and among flowers within individuals. Using analytically tractable models, we examine the optimal allocations for producing ovules, attracting pollen, and maturing seeds in deterministic and stochastic pollen environments. In deterministic environments, the optimal strategy attracts sufficient pollen to fertilize every ovule and mature every zygote into a seed. Stochastic environments select for allocations proportional to the risk of seed production being limited by zygotes or seed maturation. When producing an ovule is cheap and maturing a seed is expensive, among-plant variation selects for attracting more pollen at the expense of producing fewer ovules and having fewer resources for seed maturation. Despite this increased allocation, such populations are likely to be pollen limited. In contrast, within-plant variation generally selects for an overproduction of ovules and, to a lesser extent, pollen attraction. Such populations are likely to be resource limited and exhibit low seed-to-ovule ratios. These results highlight the importance of multiscale variation in the evolution and ecology of resource allocations. [ABSTRACT FROM AUTHOR]

Abstract: Flowers of many plant species are visited by both birds and insects, making it necessary to establish their relative contributions to seed set. In Protea, available evidence points to an overwhelming preponderance of bird-pollination systems in the genus, but the scented flowers of several dwarf grassland “sugarbush” species suggest that some Protea species may be adapted for insect pollination. In this study, we used both selective exclusion of vertebrates and complete exclusion of all visitors to investigate whether the insects that visit the scented flowerheads of three Protea species (Protea dracomontana, Protea simplex and Protea welwitschii) in KwaZulu-Natal, South Africa contribute to seed production. We also performed supplemental hand pollinations to test for pollen limitation. Seed set was generally higher in inflorescences subjected to vertebrate exclusion than in those from which all visitors were excluded, suggesting that fertile cross-pollen was deposited by insects, but these differences were slight because of high levels of self-fertilization in the study species. Pollen deposition and pollen tube growth were similar for vertebrate-excluded and open-pollinated inflorescences. Supplemental hand-pollination treatments revealed that seed set in P. simplex and P. welwitschii was not pollen-limited. Overall seed set was low, typical of the family Proteaceae, and infructescences were highly predated by lepidopteran larvae. We conclude that insects are likely to contribute to seed set of the study species, but further studies using molecular markers are required to establish the actual level of insect-mediated outcrossing. [Copyright &y& Elsevier]

Background and Aims Plants are sessile organisms that face selection by both herbivores and pollinators. Herbivores and pollinators may select on the same traits and/or mediate each others' effects. Erysimum capitatum (Brassicaceae) is a widespread and variable plant species with generalized pollination that is attacked by a number of herbivores. The following questions were addressed. (a) Are pollinators and herbivores attracted by similar plant traits? (b) Does herbivory affect pollinator preferences? (c) Do pollinators and/or herbivores affect fitness and select on plant traits? (d) Do plant compensatory responses affect the outcome of interactions among plants, pollinators and herbivores? (e) Do interactions among E. capitatum and its pollinators and herbivores differ among sites and years? Methods In 2005 and 2006, observational and experimental studies were combined in four populations at different elevations to examine selection by pollinators and herbivores on floral traits of E. capitatum. Key Results Pollinator and herbivore assemblages varied spatially and temporally, as did their effects on plant fitness and selection. Both pollinators and herbivores preferred plants with more flowers, and herbivory sometimes reduced pollinator visitation. Pollinators did not select on plant traits in any year or population and E. capitatum was not pollen limited; however, supplemental pollen resulted in altered plant resource allocation. Herbivores reduced fitness and selected for plant traits in some populations, and these effects were mediated by plant compensatory responses. Conclusions Individuals of Erysimum capitatum are visited by diverse groups of pollinators and herbivores that shift in abundance and importance in time and space. Compensatory reproductive mechanisms mediate interactions with both pollinators and herbivores and may allow E. capitatum to succeed in this complex selective environment. [ABSTRACT FROM PUBLISHER]

Ecology Letters (2010) 13: 321–329 The evolutionary stability of mutualisms is enhanced when partners possess mechanisms to prevent overexploitation by one another. In obligate pollination–seed consumption mutualisms, selective abortion of flowers containing excessive eggs represents one such mechanism, but empirical tests have long been limited to the yucca–yucca moth mutualism. We present evidence for selective abortion in the recently discovered mutualism between Glochidion trees and Epicephala moths. In Glochidion acuminatum, proportion of aborted flowers progressively increased both with higher egg load and increased ovule damage. Selective abortion resulted in a 16% seed production increase compared with expectations under random abortion, and moths suffered fitness losses as high as 62% when ovipositing into pre-infested flowers. Moth eggs were laid singly more often than expected under random oviposition, thus avoiding potential disadvantages from multiple infestations. As new pollination mutualisms are being discovered, selective abortion mechanisms may prove to be more widespread than previously thought. [ABSTRACT FROM AUTHOR]

Many nonnative plants require pollinators to produce the seed needed to establish beyond their native range. Local variation in pollination can cause variation in mating patterns, reproductive output, genetic diversity, and selection, which can influence invasion dynamics. We quantified pollinator visitation, pollen deposition, and pollen limitation of reproduction in the invasive shrub Lonicera maackii in adjacent forest edge and forest interior habitats. Flowers in edge habitats received as many or more pollinator visits than those in interior habitats, resulting in twofold greater pollen deposition and nearly twice as many seeds produced per flower. Supplemental pollination increased seed production over open pollination by 37%-145% in both edge and interior habitats. Edge flowers produced more seed than interior flowers. Despite greater pollen deposition in edge habitats, the magnitude of pollen limitation was similar to interior plants. Thus, different mechanisms of pollen limitation characterize these habitats. Pollen receipt and other resources limited reproduction of interior plants, which were located in a shady environment. Pollen quality likely limited reproduction of edge plants, because pollinators visited more flowers per plant and likely delivered more geitonogamous pollen in this habitat than in the interior. Therefore, local variation in abiotic conditions influenced reproduction in this invasive plant, both directly and indirectly via pollinators. [ABSTRACT FROM AUTHOR]

• This study tests the Haig–Westoby model, which predicts that seed output will be limited simultaneously by pollen and resources when plants optimally distribute their reproductive investment. • The test was conducted over 2 yr using Stylidium armeria in a factorial design that fully crossed three pollination levels (small stigmatic loads, open pollination, and supplementation of natural loads) with three levels of resource availability (reduction through partial defoliation, unmanipulated resource conditions, and supplementation through nitrogen, phosphorus and potassium (NPK) addition). • There was no evidence of pollen limitation from supplemental pollination; however, pollen reductions (to about half the normal mean stigmatic loads) sharply reduced seed output. There was no evidence of resource limitation, in that NPK addition did not, by itself, significantly elevate seed output in either year of the study, while resource reduction by defoliation lowered seed output in the second year. Simultaneous addition of both pollen and resources strongly and significantly increased seed production. • These results match the direction of effects predicted by the Haig–Westoby model, and suggest that S. armeria plants at our site are at or near an equilibrium of joint limitation of seed production by pollen capture and resource availability. [ABSTRACT FROM AUTHOR]

Spatial and temporal variation in the seed production of a species and the various factors affecting seed production provide essential information for understanding biotic and abiotic controls on reproduction and for assessing population trends. We conducted a 4-year study combining field observations and pollination experiments to evaluate the major factors causing variation in seed production of Primula modesta at two sites (fen and grassy sites) on Mt Asama in central Japan. Primula modesta switched from a vegetative to reproductive life phase at a stage that depended on its accumulation of biomass, and the number of flowers produced by each plant was also highly correlated with plant size. Plants at the grassy site, where snow melts 2–3 weeks earlier than at the fen site, can accumulate more biomass and, thus, bear more flowers and ovules than those at the fen site. Although P. modesta is distylous with both self and intramorph incompatibility, pollen availability did not limit seed production at either site. Annual seed production was relatively constant at both sites over the four seasons except 2002, but it was influenced by many environmental factors, including stochastic events, and by antagonistic interactions with herbivores and pathogens. In 2002, many flowers failed to bloom at the fen site. This was likely caused by frost damage to flower buds because of an unusually early snowmelt in that year. Unusual climatic events may be a crucial factor causing spatial and temporal variation in seed production at our study sites. Longer-term observations are needed to evaluate the effect of unusual climatic events on reproduction. [ABSTRACT FROM AUTHOR]

For successful reproduction animal pollinated plants must provide resources for both pollinator attraction and offspring production, and theory suggests that resources and pollen delivery limit reproduction simultaneously. We conducted a series of experiments involving supplemental pollination, flower removal, fertilizer addition and foliage removal to investigate the interaction of resources and pollen on fruit-set of Aciphylla squarrosa, a long-lived, dioecious, masting herb in Wellington, New Zealand. Reducing floral display decreased open-pollinated fruit-set, suggesting that display size is a reflection of an optimal investment between attraction and fecundity. In combination with supplemental pollination, resource reduction and fertilization addition did not alter fruit-set, suggesting that changes in resource availability did not limit reproduction in the current year. In addition, supplemental pollination of non-manipulated treatments did not increase fruit-set, demonstrating that plants were not naturally pollen limited. While we found that simultaneous pollen and resource limitation did not occur within a season, this is possibly mitigated by life history patterns including mast flowering and a storage taproot. Multiple year studies are required to further examine simultaneous resource and pollen limitation. [ABSTRACT FROM AUTHOR]

Although several factors can limit female fertility in perennial plants, rarely have they been jointly studied in a single species over several years. In this study we experimentally manipulate seed production and simultaneously analyse the potential contribution of pollen limitation, costs of reproduction and plant size to variation in seed output over a 3-year period in the perennial herb Paeonia officinalis, in southern France. Since this rare species is threatened by forest closure in many sites we also examine the causes of female fertility variation in relation to habitat closure (open habitat vs. woodland). P. officinalis has a partial self-incompatibility system and only very low ability for autonomous self pollination in the absence of pollinators. However, supplementary pollination of individual plants in three consecutive years did not significantly increase seed production above natural levels. Forest closure was associated with a decline in ovule and seed production, which again was not due to pollen limitation since supplementary pollination had no significant effect on seed set in the woodland habitat. Comparison of the maternal fertility of plants which were previously excluded from reproduction with those which were hand pollinated to maximise seed set in two previous years produced no evidence that seed production in year three is limited by costs associated with prior reproduction. Likewise, flowering probability was not related to prior seed production but was however positively related to plant size. The absence of any influence of pollen limitation or prior reproduction on seed production suggests that sub-maximal seed production in long-lived perennial herbs may be part of a size-dependent strategy that maximises life-time seed production and fitness without compromising survival. [ABSTRACT FROM AUTHOR]

The standard method of measuring pollen limitation is to add pollen to a number of flowers, preferably to a whole plant, and to compare fruit and seed set with that of naturally pollinated flowers on other plants. In 25 yr of research, this method has yielded valuable data, but it is difficult to use in large plants. This has caused a bias in the available data towards smaller, herbaceous plants with relatively few flowers. I argue that, in order to widen our knowledge of how pollen limitation affects plants, we should go beyond whole-plant pollen addition and change our concept of how a flowering plant functions. The traditional method does not take into account the variation in and dynamics of resource allocation and pollen availability. The concept of integrated physiological units (IPUs) does, but, although it has been applied to pollination biology, it has not received the attention it deserves. I use this article to present its merits again, to propose a step-by-step methodology for studying pollen limitation, and to examine factors influencing possible plant strategies. [ABSTRACT FROM AUTHOR]

Pollen limitation and resource limitation were invoked to account for the pattern that flowering plants produce more flowers and ovules than fruits and seeds. This study aimed to determine their relative importance in Veratrum nigrum, a self-compatible, perennial, andromonoecious herb. In order to determine whether female production was limited by pollen grains on stigmas or by available resources, we performed supplemental hand pollination in three populations, male-flower-bud removal in three other populations, and emasculation of hermaphroditic flowers in still another population, resulting in a total of seven populations experimentally manipulated. Across the three populations, supplemental hand pollination did not significantly increase fruit set, seed number per fruit, and total seed production per individual, nor did emasculation of hermaphroditic flowers. Taken together, our results suggest that pollen grains deposited on stigmas were abundant enough to fertilize all the ovules. Male-flower-bud removal significantly increased the mean size of hermaphroditic flowers in all three populations. Female reproductive success was increased in one population, but not in the other two populations possibly due to heavy flower/seed predation. We concluded that the female reproductive success of V. nigrum was not limited by pollen grains but by available resources, which is consistent with Bateman's principle. Furthermore, the female reproduction increase of male-flower-bud removal individuals might suggest a trade-off between male and female sexual functions. (Managing editor: Wei Wang) [ABSTRACT FROM AUTHOR]

Pollen limitation, resource limitation, fruit abortion, and predation have all been proposed as factors explaining low fruit set in hermaphroditic plants. We conducted a 5-year study combining field observations and pollination experiments to determine the causes of the low fruit set in Aristolochia paucinervis, a Mediterranean species with a specialized pollination system in two populations in SW Spain. Fruit initiation was markedly low, and between 28.6 and 75.0% of the flowering stems did not initiate any fruit. In most flowers, the number of germinated pollen grains was less than the number of ovules, and supplemental pollination significantly increased fruiting, indicating deficient pollination. In A. paucinervis, autonomous self-pollination seems to be a decisive factor in fruit production because the number of germinated pollen and the fruit set from flowers bagged before anthesis were similar to those in free-pollinated flowers. Only in 2005 did flowers that were successfully pollinated outnumber ripened fruits, suggesting that other factors limit fruiting. We found a significant positive correlation between tuber mass and fruit set. Deficient pollination and lack of resources could explain the low fruit set, but the relative consequences seem to vary spatially and temporarily. [ABSTRACT FROM AUTHOR]

Abstract: We assessed natural rates of floral abortion in four common mangrove species from northern Australia and subsequently manipulated pollination experimentally. Sonneratia alba J. Smith exhibited the highest rate of fruit set of the four species (23%), indicating this mangrove was best able to utilise the natural pollination opportunities provided. Fruit set in S. alba appeared, however, to be pollinator limited, as large increases in fruit set occurred after manual cross-pollination of flowers. Avicennia marina (Forsk.) Vierh. had the highest rate of natural pollination, but fruit set was lower (15%) and appeared to be impeded by resource limitations. Although a range of insects visited Ceriops australis (C.T. White) Ballment, T.J. Sm & Stoddart, the rate of fruit set was low (3%) and the capacity for flower fertilisation limited, despite evidence of autogamy in this species. There was an indication of both resource and pollinator limitation in C. australis. Rhizophora stylosa Griff. exhibited limited fruit set (0.5%), possibly due to limiting maternal resources and the lack of adaptation of flowers to either animal or wind pollination. Large increases in fruit set were recorded after manual cross-pollination of R. stylosa flowers. R. stylosa and C. australis, characterised by resource rich propagules with long periods of development, both aborted a large proportion of propagules during the fruit maturation process. [Copyright &y& Elsevier]

Quantifying the extent to which seed production is limited by the availability of pollen has been an area of intensive empirical study over the past few decades. Whereas theory predicts that pollen augmentation should not increase seed production, numerous empirical studies report significant and strong pollen limitation. Here, we use a variety of approaches to examine the correlates of pollen limitation in an effort to understand its occurrence and importance in plant evolutionary ecology. In particular, we examine the role of recent ecological perturbations in influencing pollen limitation and discuss the relation between pollen limitation and plant traits. We find that the magnitude of pollen limitation observed in natural populations depends on both historical constraints and contemporary ecological factors. [ABSTRACT FROM AUTHOR]

Differences between plant sex morphs in pollen or resource availability may affect their relative fitness and thereby the sex ratio of dimorphic species. In gynodioecious species, in which hermaphroditic and female plants coexist, a variety of factors (e.g.. hermaphrodite self-fertility or rarity or pollinator discrimination against females) might be expected to lead to stronger pollen limitation in females than in hermaphrodites. On the other hand, females have been found to be superior compared to hermaphrodites in low-nutrient conditions. The effects of supplemental hand-pollination and resource addition on the reproductive output of the self-fertile gynodioecious perennial Geranium sylvaticum (Geraniaceae) were tested for several populations that differ in their female frequency 4.4–23.O%). Both pollen and resource availability limited fruit set and the number of seeds produced per plant; however, seed set (i.e., the number of seeds produced per fruit) was limited only by resources. Because pollen limitation in females did not correlate with female frequency, our results suggest that pollen limitation in females does not depend on the frequency of the pollen-producing hermaphrodites. Furthermore, because pollen and resource availability limited reproductive output of both sex morphs, these factors may not contribute significantly to maintenance and evolution of gynodioecy in G. sylvaticum. [ABSTRACT FROM AUTHOR]

AbstractPollinator limitation and resource limitation of seed production were examined in flowers of a natural population of Scotch broom, Cytisus scoparius, by taking advantage of the floral characteristics that flowers tripped open by an effective pollinator were easily distinguishable from unvisited flowers. In total, 40.26% of flowers were visited by effective pollinators, 28.91% were pollinated and 13.75% matured fruits. Therefore, 71.80% of visited flowers were pollinated and 47.58% of pollinated flowers matured fruits. Thus, the most limiting factor for fruit production was pollinator visitation rate (pollinator limitation), and secondarily the process from pollination to fruiting further constrained fruit production (resource limitation). The pollinator visitation rate was significantly higher for plants growing in a sunny habitat than in a shady one, and pollinator limitation was more severe in the shady habitat than the sunny one. The proportion of fruit produced to pollinated flowers was not significantly different between the sunny and shady habitats, but was lower on larger plants, indicating that resource limitation was more severe on larger plants producing many more flowers. [ABSTRACT FROM AUTHOR]

Destruction and fragmentation of natural habitats is the major reason for the decreasing biodiversity in the agricultural landscape. Loss of populations may negatively affect biotic interactions and ecosystem stability. Here we tested the hypothesis that habitat fragmentation affects bee populations and thereby disrupts plant-pollinator interactions. We experimentally established small ”habitat islands” of two self-incompatible, annual crucifers on eight calcareous grasslands and in the intensively managed agricultural landscape at increasing distances (up to 1000 m) from these species-rich grasslands to measure effects of isolation on both pollinator guilds and seed set, independently from patch size and density, resource availability and genetic erosion of plant populations. Each habitat island consisted of four pots each with one plant of mustard ( Sinapis arvensis) and radish ( Raphanus sativus). Increasing isolation of the small habitat islands resulted in both decreased abundance and species richness of flower-visiting bees (Hymenoptera: Apoidea). Mean body size of flower-visiting wild bees was larger on isolated than on nonisolated habitat islands emphasizing the positive correlation of body size and foraging distance. Abundance of flower-visiting honeybees depended on the distance from the nearest apiary. Abundance of other flower visitors such as hover flies did not change with increasing isolation. Number of seeds per fruit and per plant decreased significantly with increasing distance from the nearest grassland for both mustard and radish. Mean seed set per plant was halved at a distance of approximately 1000 m for mustard and at 250 m for radish. In accordance with expectations, seed set per plant was positively correlated with the number of flower-visiting bees. We found no evidence for resource limitation in the case of mustard and only marginal effects for radish. We conclude that habitat connectivity is essential to maintain not only abundant and diverse bee communities, but also plant-pollinator interactions in economically important crops and endangered wild plants. [ABSTRACT FROM AUTHOR]

1. An experimental approach, manually pollinating all the flowers of individual plants, was used to measure the effect of pollen limitation on female fecundity of the hummingbird-pollinated perennial shrub Echeveria gibbiflora in the ecological preserve of Pedregal de San Angel around México City, México. Eleven randomly selected plants were manually over-pollinated in all their flowers and another 11 were left to be freely visited by natural pollinators. 2. Manually pollinated plants produced significantly more fruit and seeds than control plants (1·38 and 1·74 times, respectively). There was no change in average mass of fruits. 3. Considering individual fruit production per plant sampled three times in one season, decreases in fruit mass and average seeds per fruit were observed within the same reproductive season for both treatments. For the manually pollinated plants, from the start to the end of the reproductive season, seed set decreased 55·9%; while in control plants seed set decreased 33·4% in the same period. For both treatments, average fruit mass decreased 26%. 4. Vegetative growth was not significantly different between control and experimental plants but hand-pollinated plants showed a smaller reproduction probability for the following year. 5. It is concluded that female fecundity in E. gibbiflora is limited by pollen early in the reproductive season and by resources in the middle and the end of the season. [ABSTRACT FROM AUTHOR]

The differentiation patterns of reproductive systems, including breeding and pollination systems, and perennation strategies of the genus Trillium were critically studied. The pedicellate- and sessile-flowered species with different floral morphologies and ecological distributions exhibited distinct modes of reproductive systems in relation to their habitat conditions. The pedicellate-flowered species occurring in stable mesic temperate deciduous forests reproduce exclusively by seeds, while sessile-flowered species growing in the alluvial flood plain habitats depend to a large extent on vegetative reproduction to compensate for the insufficient sexual reproduction under ecologically unstable conditions, e.g., frequent disturbance due to flooding. These sessile-flowered species are characterized by low ovule numbers, subsequent low seed outputs and low fecundity levels. In contrast to the sessile-flowered species, the pedicellate-erect-flowered species showed very high seed setting rates of over 50%. Breeding experiments for four Japanese species suggest that, although they substantially possess the capability of both inbreeding and outbreeding, the high fecundity levels are maintained by predominant inbreeding system. Furthermore, resource levels in the stock organs of these species obviously determine fecundity levels. As a consequence the amount of reserved food in the rhizomes transferred to the next season is determined, which gurantees a continuous reproductive activity of a given individual plant. [ABSTRACT FROM AUTHOR]