Determinants of seed production in Geranium maculatum

Summary We experimentally examined factors limiting seed production in two populations of the perennial woodland herb Geranium maculatum in central Illinois, USA. To test the pollinator-limitation hypothesis, we compared the seed production of plants whose flowers were supplementarily pollinated with outcross pollen to that of control plants receiving natural pollination only. To test if fruit production by early flowers suppresses fruit and seed formation by late flowers, a third group of plants was prevented from producing seed from the first 50% of the flowers to open (stigmas were excised at flower opening). Finally, to test if seed maturation and flower initiation are correlated with photosynthetic capacity, we performed a defoliation experiment in which either the stem leaves within the inflorescence, the stem leaves below the inflorescence, or the rosette leaves were removed during late flowering. Plants that reccived supplemental pollination produced 1.5–1.6 times more seeds than control plants. We found no difference between hand-pollinated plants and controls in mortality, flowering frequency or number of flowers produced in the year following the experiment. In both control and hand-pollinated plants, the fruit set and total seed production of early flowers were more than twice as high as those of late flowers. In one of the two populations, plants whose early flowers were prevented from setting seed produced significantly more seeds from their late flowers than did control plants. Seed predation was low and did not differ between early and late flowers. Leaf removal did not affect seed number or size in the year of defoliation, nor did it reduce survival or flower production in the subsequent year. This suggests that the plants were able to compensate for a partial defoliation by using stored resources or by increasing photosynthetic rates in the remaining leaves. Taken together, the results demonstrate that both pollinator activity and resource levels influence patterns of seed production in G. maculatum. While seed production was pollinatorlimited in both populations, a seasonal decline in resource availability was apparently responsible for the low seed production by late flowers.     

FLORAL BIOLOGY AND REGULATION OF SEED SET AND SEED SIZE IN THE LILY,CLINTONIA BOREALIS

We studied the reproductive ecology of Clintonia borealis, a clonal understory species, in eastern Ontario. Flowers are protogynous and require insect pollination for outcrossing and maximum seed set. Most pollination is done by nectar-gathering bumble bees. We recorded pollen deposition and flowering patterns in 1983. Flowering lasted for about 10 days in mid-June. Stigmatic pollen loads were lowest at the onset of blooming, reflecting the scarcity of male-phase flowers. Pollination increased significantly by peak bloom and subsequently remained high. However, since bees fly mostly between neighboring stems, much of the pollen transferred may have been geitonogamous. In 1983, seed set per flower was not increased by supplementing pollination or by reducing the number of fruits competing for resources per stem. Rather, flowers set more seeds (14%) only if both treatments were performed simultaneously. Seed size was increased by 25% when competitive fruits were removed and by 5% more when pollen was added under the removal treatment. Further work in 1984 showed that pollination effects may be related to changes in pollen source. Selfed flowers set fewer and smaller seeds than outcrossed ones when screened from insects and hand pollinated. Outcrossing distance had little effect on seed set or seed size. Thus, in C. borealis seed number and seed size are limited by a balance between maternal resource availability and the amount of outcrossing provided by pollinators.     

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.     

华北驼绒藜自然种群结实的花粉和资源限制

运用人工授粉、补充无机营养、去叶处理以及疏花处理研究了华北驼绒藜自然种群花粉和资源有效性对结实的影响,并进一步探讨了该植物的选择性败育现象和繁殖对策。结果表明:花粉来源而不是花粉数量对华北驼绒藜结实存在显著影响;同枝授粉结籽率和结实率均低于自然授粉,异株授粉则显著高于同枝授粉和自然授粉;异株授粉种子千粒重显著高于自然授粉和同株异枝授粉,极显著高于同枝授粉。华北驼绒藜结实存在资源限制,补充施肥提高了单枝开花数和结籽率。随摘除叶片数量的增加,结籽率明显降低,摘除叶片处理与对照之间均形成显著差异,摘除1/2叶、3/4叶以及去全叶处理使种子千粒重显著低于对照。人工疏花实验表明华北驼绒藜存在选择性败育现象。华北驼绒藜可根据花粉和资源的可利用性来调整性分配、授粉方式和结实以获得最大适合度,具有复杂的繁殖对策,表现出较好的环境适应性。     

The role of pollination level on the reproduction of females and hermaphrodites in the gynodioecious plant Gypsophila repens (Caryophyllaceae)

In gynodioecious plant species, females are expected to have more resources available for maturing seeds because pistillate flowers are smaller, do not produce pollen, and are thus less costly that perfect flowers. The potential female advantage arising from more abundant resources is, however, likely to vary depending on whether seed production is limited by resource or pollen availability. Here we experimentally investigated the influence of pollen and resource limitation on female advantage in a gynodioecious species using two levels of pollination. Total seed production of females was always greater than that of hermaphrodites: females produced more flowers and more fruits that contained similar numbers of seeds of similar mass. Under low pollination, female and hermaphrodite plants allocated resources to increased flower production rather than to increased seed size or quality. We did not detect any influence of pollen or resource limitation on female advantage, which remained similar under low (= abundant resources) and full pollination. Outcrossed fruits performed better than selfed fruits when the same plant received both selfed and outcrossed pollen on different flowers. These differences were not greater under high pollination, possibly because resources available for each fruit did not differ between our pollen intensity treatments.     

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。     

毛翠雀花花序内的性分配和繁殖成功

两性花植物花序内不同位置的性分配和繁殖成功一般存在差异,通常认为资源竞争、结构效应和交配环境是形成这种差异的主要原因。为了研究雄性和雌性繁殖资源在花序内不同位置间的最优分配问题,该文以青藏高原高寒草甸典型高山植物毛翠雀花为材料,通过比较花序内不同位置的花部特征和种子性状,对其花序内的性分配模式和雌性繁殖成功进行研究,并通过观察传粉者运动特点以及人工去花和补授花粉实验,探讨花序内资源竞争和交配环境对繁殖资源分配的影响。结果表明:(1)不同位置间的雄蕊数、雄蕊鲜重/雌蕊鲜重、花粉数及花粉胚珠比从花序基部到上部显著增加,而雌蕊鲜重和胚珠数逐渐减少,表现出上部花偏雄的性分配;上部花的结籽率显著低于基部花和中部花,不同位置间的发育种子数/果实和发育种子重/果实随着花位置的升高而显著降低,说明基部花具有更佳的雌性繁殖成效。(2)去花处理后,剩余果实的单个种子重/果实显著增加,但发育种子数/果实没有显著增加;而给上部花人工补授异花花粉后,位置间结籽率的差异消失,说明传粉限制而非资源竞争导致了花序内位置依赖的种子生产模式。(3)毛翠雀花雄性先熟的开花特征,以及传粉者苏氏熊蜂从花序基部到上部的定向访花行为,导致了花序内交配环境的变化。综上结果表明,毛翠雀花花序内的性分配和繁殖成功差异是对交配环境适应的结果,对其在高山环境中实现雌雄适合度最优化具有重要意义。     

Effects of pollen and nitrogen availability on reproduction in a woodland herb,Lysimachia quadrifolia

Summary Relationships between pollen loads, resource availability, and fruit and seed production were determined for Lysimachia quadrifolia ramets in two adjacent sites (the scrub site and the open site) in 1982, 1983, and 1984. Pollen loads limited % fruit set and seed production in the open site in 1982 and 1983. Reproduction in the scrub site was resource limited in 1982, as shown by an increase in % fruit set when one-half of the flowers on a ramet were removed prior to fruit initiation. In the scrub site in 1983, pollination of one-half of the flowers on a ramet decreased the % fruit set of the remaining, unpollinated flowers. Fruit production in the same site was limited by pollen in 1984. Addition of nitrogen fertilizer to the scrub site in 1984 had no effect on fruit and seed production. There was more variation in fruit set between sites than between years. There was no trend to greater fruit set or number of seeds/fruit on early flowers compared to late flowers on the same ramet. Correlations between measures of reproduction were positive or insignificant. These results demonstrate year-and site-specific variation in the factors that limit plant reproduction.     

Pollination intensity and potential seed set in Passiflora vitifolia

Summary Initial seed set and fruit set were pollen-limited in a Costa Rican population of Passiflora vitifolia, a self-incompatible species with 200–350 ovules per flower. Pollination intensity was measured by counting the number of allogamous pollen grains on stigmas of the large one-day flowers. Hand-pollinations demonstrated that 25–50 pollen grains are required for fruit set, and >450 are needed for maximum seed set, with a pollen:seed ratio of about 1.6:1.0. Hummingbirds (Phaethornis superciliosus) delivered sufficient allogamous pollen for maximum seed set to only 28% of the flowers examined. Naturally pollinated flowers yielded fewer fruits and fewer seeds per fruit than those pollinated by hand. Most pollen transferred by humming-birds was self-incompatible; emasculated flowers yielded higher seed set than flowers with intact anthers. Visitation rates did not provide a good index of effective pollination.There were significant differences in ovule number, maximum seed set, and maximum per cent seed set among individual vines. More than half of an individual's flowers failed to set fruit, whether pollinated by birds or by hand. In this population, maximum reproductive potential may be limited by maternal resources for fruit development, but seed set varies with pollination intensity. Pollen-limited seed set may be a disadvantage of self-incompatibility, especially in species with many-seeded fruits.     

EFFECTS OF POLLINATION INTENSITY IN CAMPSIS RADICANS

I examined effects of pollination intensity on fruit, seed and seedling characteristics in Campsis radicans, and joint effects of pollen donor and pollination intensity on fruit production. Large pollen loads were more likely to initiate fruit production than small pollen loads, and the former fruits contained more seeds and a greater total seed mass. No further increases in seed number or mass occurred for pollen loads above 4,000 grains. The weight of individual seeds was unaffected by pollen load. Effects of pollen donor were generally larger than effects of pollen load, and fruit production from small loads of pollen from one donor were sometimes equal to fruit production from larger pollen loads from another donor. The ratio of pollen grains deposited to resultant seeds increased with pollen load, and several explanations are proposed. Seeds from heavy pollinations emerged better than seeds from light pollinations, but did not differ in speed of germination or in the performance of seedlings up to 126 days. The emergence differences are probably due to differing intensities of pollen tube competition.     

Pollen source and resource limitation to fruit production in the rare species Eremosparton songoricum (Fabaceae)

Eremosparton songoricum (Litv.) Vass. is a rare, central Asian desert species which shows lower fruit set and seed set (<16%) than most hermaphroditic species. We hypothesized that fruit production was limited by pollen and resources. To evaluate potential fruit abortion due to pollen limitation, supplemental hand‐pollination was undertaken, the mating system was investigated and the foraging behavior of pollinators was recorded. To investigate possibile resource limitation, flowers and young pods were artificially removed and fertilization were manipulated. The results showed that under natural pollination, the number of pollen deposited on the stigma greatly exceed the number of ovules per ovary. Mating system experiments showed that the species is self‐compatible, but depended on pollinators to set seeds. Supplemental outcross pollination increased fruit set significantly. The most frequent effective pollinator Megachile terminate Morawitz, was observed pollinating many flowers of the same individual plant (74.5±1.3%). These results suggested that fruit production is affected by insufficient outcross pollen rather than by pollen quantity. Removal of 2/3 of the flowers and young pods led to significantly higher fruit set, as did addition of fertilizers (N–P–K: 0.025–0.05–0.013 g, N–P–K: 0.05–0.1–0.025 g) showing that reducing resource acceptors and increasing inorganic resources both helps to improve fruit set. We therefore conclude that reproductive success of E. songoricum is limited by both outcross pollen and available nutrients.     

Quantitative and qualitative consequences of reduced pollen loads in a mixed‐mating plant

Greater pollination intensity can enhance maternal plant fitness by increasing seed set and seed quality as a result of more intense pollen competition or enhanced genetic sampling. We tested experimentally these effects by varying the pollen load from a single pollen donor on stigmas of female flowers of Dalechampia scandens (Euphorbiaceae) and measuring the effects on seed number and seed mass. Seed set increased rapidly with pollen number at low to moderate pollen loads, and a maximum set of three seeds occurred with a mean pollen load of 19 pollen grains. We did not detect a trade‐off between the number of seeds and seed mass within a fruit. Seed mass increased with increasing pollen load, supporting the hypothesis of enhanced seed quality via increased pollen‐competition intensity or genetic sampling. These results suggest that maternal fitness increases with larger pollen loads, even when the fertilization success is already high. Our results further highlight the importance of high rates of pollen arrival onto stigmas, as mediated by reliable pollinators. Comparing the pollen‐to‐seed response curve obtained in this experiment with those observed in natural populations suggests that pollen limitation may be more severe in natural populations than predicted from greenhouse studies. These results also indicate that declines in pollinator abundance may decrease plant fitness through lowered seed quality before an effect on seed set is detected.     

Comparison of two cacao (Theobroma cacao L.) clones for the effect of pollination intensity on fruit set and seed content

We compared the influence of pollination intensity (PI) on fruit set and seed number per pod in two cacao (Theobroma cacao L.) clones, IFC5 (Forastero Lower-Amazon Amelonado), which is self-compatible and known to produce a high number of seeds per pod under open pollination, and SCA6 (Forastero Upper-Amazon), which is self-incompatible and known to produce a low number of seeds per pod under open pollination. With both clones, PI had a positive effect on fruit set, with the maximum rate requiring more than 150 pollen grains per pod. One-half of the maximum rate of fruit set was reached with 39 pollen grains per pod for SCA6, and 78 for IFC5. With SCA6, a significant positive effect of PI on seed number per pod was also observed, with maximum seed number requiring more than 200 pollen grains per pod. In contrast, seed numbers with IFC5 were approximately equal over the PI range 100–800 pollen grains per pod. Patterns of seed number per pod were compared after: (1) open insect pollination, (2) low-intensity hand pollination and (3) high-intensity hand pollination. The patterns obtained with IFC5 under open pollination showed a peak of 40–50 seeds per pod, whereas the distribution patterns were more even with SCA6. The pattern of seed number under open pollination was similar to that obtained with high-intensity hand pollination for IFC5, and with low-intensity hand pollination for SCA6. We concluded that the high number of seeds per pod observed with IFC5 under natural insect pollination may be explained by a high number of compatible self-pollen grains on the stigma and by a severe drop of low-pollinated flowers eliminating potential lowfilled fruits. With SCA6, however, the number of compatible pollen grains deposited on the stigma was probably low under open pollination, and the flowers required lower pollen quantities to set fruit, which resulted in a high frequency of low seed numbers per fruit. This difference in the capacity to set low-seeded fruits might be considered as an adaptive trait related to the mode of reproduction.     

Assessment of reproductive characteristics of Jatropha curcas L. in south Florida

Jatropha curcas L. (jatropha) is a species identified for biofuel production because of the high quality of the oil produced by its seeds. However, jatropha is undomesticated and little information is available about its reproductive characteristics. Breeding and genetic improvement programs are much needed for the jatropha development as a bioenergy crop. Information about floral display and mode of reproduction are considered essential for breeding programs. In this study, the total number of female flowers, male to female flower ratio, fruit set, in vitro pollen germination, and the formation of fruits by apomixis, self‐pollination, and natural pollination were evaluated in 17 jatropha accessions planted in South Florida. The total number of female flowers per inflorescence and male to female flower ration ranged from 2.8 to 9.1 and 9.9:1 to 55.4 : 1, respectively. During summer 2011, high fruit setting average was observed (75.5%). In vitro pollen germination varied from 64.6% during spring 2011 to 51.6% during fall 2011. The fruit set observed was from 10.1% to 64.0% through natural pollination and apomixes, respectively. Characteristics such as fruit fresh weight, number of seeds per fruit, seed dry weight, and oil content were influenced by mode of reproduction.     

A ballistic pollen dispersal system influences pollination success and fruit‐set pattern in pollinator‐excluded environments for the endangered species Synaphea stenoloba (Proteaceae)

The critically endangered Synaphea stenoloba (Proteaceae) has numerous scentless flowers clustered in dense inflorescences and deploys a ballistic pollen ejection mechanism to release pollen. We examined the hypothesis that active pollen ejection and flowering patterns within an inflorescence influence the reproductive success (i.e. fruit formation) of individual flowers within or among inflorescences of S. stenoloba in a pollinator‐excluded environment. Our results showed that: (1) no pollen grains were observed deposited on the stigma of their own flower after the pollen ejection system was manually activated, indicating self‐pollination within an individual flower is improbable in S. stenoloba; (2) fruit set in the indoor open pollination treatment and the inflorescence‐closed pollination treatment indicated that S. stenoloba is self‐compatible and pollen ejection can potentially result in inter‐floral pollination success; (3) fruit set in the inflorescence‐closed pollination treatment was significantly lower than that of indoor open pollination, indicating within‐ and between‐flower pollination events in an inflorescence are most likely limited, with pollination between inflorescences providing the highest reproductive opportunity; and (4) analysis of the spatial distribution of cumulative fruit set on inflorescences showed that pollen could reach any flower within an inflorescence and there was no functional limitation on seed set among flowers located at various positions within the inflorescence. These data suggest that the pollen ejection mechanism in S. stenoloba can enhance inter‐plant pollination in pollinator‐excluded environments and may suggest adaptation to pollinator scarcity attributable to habitat disturbance or competition for pollinators in a diverse flora. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 59–68.     

The evolution of ovule number and flower size in wind-pollinated plants

In angiosperms, ovules are "packaged" within individual flowers, and an optimal strategy should occur depending on pollination and resource conditions. In animal-pollinated species, wide variation in ovule number per flower occurs, and this contrasts with wind-pollinated plants, where most species possess uniovulate flowers. This pattern is usually explained as an adaptive response to low pollen receipt in wind-pollinated species. Here, we develop a phenotypic model for the evolution of ovule number per flower that incorporates the aerodynamics of pollen capture and a fixed resource pool for provisioning of flowers, ovules, and seeds. Our results challenge the prevailing explanation for the association between uniovulate flowers and wind pollination. We demonstrate that when flowers are small and inexpensive, as they are in wind-pollinated species, ovule number should be minimized and lower than the average number of pollen tubes per style, even under stochastic pollination and fertilization regimes. The model predicts that plants benefit from producing many small inexpensive flowers, even though some flowers capture too few pollen grains to fertilize their ovules. Wind-pollinated plants with numerous flowers distributed throughout the inflorescence, each with a single ovule or a few ovules, sample more of the airstream, and this should maximize pollen capture and seed production.     

Breeding system and factors limiting fruit production in the nectarless orchid Broughtonia lindenii

Low fruit set values in most orchids (especially epiphytic and tropical species) are normally thought to be the consequence of pollination constraints and limited resources. In particular, pollination constraints are modulated by pollinator visitation rates, pollinator visitation behaviour (promoting crossing or selfing), the type and number of pollinia deposited on stigmas (in the case of orchids with subequal pollinia) and the amount of pollen loaded per inflorescence. In order to assess to what extent these factors can affect fruit set in specific orchid-pollinator systems, the repercussions of some of these aspects on reproduction of Broughtonia lindenii were examined in a coastal population in western Cuba. The study focused on plant breeding system, importance of pollen load and type of pollinia on subsequent fruit and seed, limiting factors of seed production and interaction with pollinators. This species presents long-lasting flowers that senesce after all forms of effective visit. Pollinator dependence for fruit production was demonstrated, while hand-pollination experiments revealed self-compatibility and inbreeding depression at seed level. More pollinia on stigmas enhance the proportion of well-developed seeds. In contrast, the pollinia type used in pollination is not important for seed quality of fruits, suggesting that small pollinia are not rudimentary. Natural fruit set in two consecutive years was substantially affected by pollinator activity, and also by systematic depredatory activity of ants and a caterpillar. Considering that this orchid completely lacks nectar and that the local assemblage of pollinators and predators influenced its reproduction, a minor importance of resource constraints in this epiphyte (with long-lasting reserve structures) is confirmed at least for a short time.     

Effects of Seed Number on Competition and Dominance among Fruits in Capsicum annuum L.

The effects of seed number on set, development and growth ofa fruit, and on inhibition of later-developed fruits were studiedby varying the pollen load on the stigma of sweet pepper flowers(Capsicum annuum L.). Despite much variation, a linear increasein individual fruit weight with seed number could be observed.Seed number affected the growth rate rather than the growingperiod of fruit. When seed numbers were low, the probabilityof fruit setting was positively related to seed number. However,a relatively low seed number (50–100 seeds/fruit: 20–30%of the maximum seed number) was sufficient for maximal fruitset. An increase in seed number increased the inhibitory effect ofa fruit on set and growth of later-developing fruits. As a result,when pollination treatments were applied to all the flowersof a plant, results could be quite different to those obtainedwhen only a limited number of flowers were treated. Fruit setof the second fruit was reduced by the application of a highpollen load to the first flower, even when the first fruit abortedbefore it had accumulated much dry matter. Our results suggestthat growth inhibition of the second fruit by seed number ofthe first fruit is controlled both by competition for limitedassimilates, as well as by dominance due to the production ofplant growth regulators by the developing fruit. Sweet pepper; Capsicum annuum L.; pollination; fruit set; abortion; abscission; fruit growth; first-fruit dominance; sink strength     

Plants,position and pollination: Planting arrangement and pollination limitation in a revegetated eucalypt woodland

The spatial arrangement of plants is a key determinant of pollination services in natural ecosystems. Despite this, plant arrangements are rarely considered an important characteristic of revegetated communities and this may be limiting successful pollination dynamics in these re‐created systems. We assessed if aggregated South Australian Blue Gum (Eucalyptus leucoxylon, F. Muell) had greater pollination success than more dispersed trees by measuring fruit set, seed production and germination in a revegetated eucalypt woodland. The proportion of buds that developed into fruit (fruit set) was similar between aggregated (five flowering Blue Gum within 30 m) and dispersed (no flowering Blue Gum within 30 m) trees (<14%). Aggregated and dispersed trees produced a similar number of seeds per fruit in 2015, when fewer trees flowered and those that flowered produced fewer flowers (ca 8 seeds/fruit). In 2016, when flowering was more abundant, aggregated trees produced more seeds per fruit than dispersed trees (12 seeds/fruit compared to 4 seeds/fruit). Despite differences in seed number, there was no difference in the quality of seed produced, indicating outcrossed pollen was received by both aggregated and dispersed trees. Although outcrossing was likely, this did not prevent pollination limitation, with the addition of outcrossed pollen increasing fruit and seed set for trees in both arrangements. Consequently, we propose that planting individuals of the same species (conspecifics) in clumps, rather than intermixing species, can increase seed set in revegetated eucalypts, particularly in good flowering years. However, aggregated plantings will be most effective if the needs and foraging behaviour of pollinators are also considered, to attract and sustain pollinators in revegetated systems, thereby minimizing the risk of pollination limitation.     

Pollen viability and limitation of seed production in a population of the circumpolar cushion plant, Silene acaulis (Caryophyllaceae)

Pollen viability among genders and limitation of female seed production in a natural trioecious population of the circumpolar cushion plant Silene acaulis was examined. Pollen viability was estimated by an in vitro pollen germination experiment. Both male and hermaphrodite flowers displayed large variation in pollen viability (0–53% in hermaphrodite and 0–54% in male flowers). There was a significant difference between genders in pollen viability: male plants had on average higher pollen viability than hermaphrodite plants. Resource and pollen limitation of seed production was studied by an experiment consisting of three treatments; (I) hand-pollination and removal of all other flowers on the cushion, (II) hand-pollination without removal of other flowers, and (III) open pollination without removal of flowers. Hand-pollination increased seed production, whereas removal of flowers had no effect on seed production. Abortion of pollinated ovules during seed development and seed mass did not differ among treatments. To control for effect of fruit number on seed production, data from naturally pollinated individuals was used. There was a positive correlation between both total number of seeds and fruit number, mean seed number per fruit and fruit number, respectively. These results indicate that seed production of 5. acaulis is mainly limited by pollen availability whereas resource competition between fruits is not important as a limiting factor. The possible role of male quality differences between genders and pollen limitation of seed production for maintenance of trioecious reproductive systems is discussed.