ALLOCATION OF REPRODUCTIVE RESOURCES WITHIN AND AMONG INFLORESCENCES OF LAVANDULA STOECHAS (LAMIACEAE)

Patterns of fruit set were studied in Lavandula stoechas, a Mediterranean shrub commonly occurring in southern Spain. The small, hermaphroditic flowers of this shrub are aggregated into dense, headlike inflorescences and exhibit extensive variations in fecundity. It was shown that as the number of developing fruits in the inflorescence increased, the probability of a flower setting fruit, the size of seeds, and their germinability decreased, most likely because of strong within-inflorescence resource limitation. An experiment was designed to ascertain whether increased fertility in late-opening flowers could be induced through reallocation of reproductive resources between different inflorescences. The experiment consisted of removing half of the inflorescence buds from a set of plants and comparing their fecundity with that of intact individuals. Thinning did not increase the proportion of flowers setting fruit which, in fact, was slightly lower than that of intact individuals (probably due to some reduction of floral display brought about by thinning). Although treated plants produced heavier seeds than controls, results suggest that inflorescences of L. stoechas behave as autonomous modules among which resources cannot be reallocated. Predispersal seed predation by insects accounted on average for a 31% reduction in fruit set. Predation was found to be nonrandomly distributed within inflorescences, with most damage concentrated on late fruits (i.e., those with smaller and less germinable seeds).     

Why Prunus mahaleb (Rosaceae) produces more flowers than fruits

Various mutually compatible hypotheses have been proposed to explain the low levels of fruit set observed in many hermaphrodite plants. I carried out trials to determine which of these hypotheses best explains the low fruit set occurring in Prunus mahaleb L. (Rosaceae). The results of insect exclusion, supplementary pollination, and flower removal experiments indicate that the ovary reserve hypothesis and the selective abortion hypothesis seem to be the most likely explanations for the production of excess flowers in P. mahaleb.     

REPRODUCTIVE SUCCESS AND INFLORESCENCE SIZE OF CALOPOGON TUBEROSUS (ORCHIDACEAE)

Reproductive success of Calopogon tuberosus, which produces no nectar, was investigated in relation to inflorescence size and dispersion pattern. Mean inflorescence size was 2.56 (range 1–10). A bagging experiment showed that insects are required for pollen transfer and that fruits are produced from self-, geitonogamous, and cross-pollinations; fruit set was not 100%. Fruit set of nonmanipulated plants was limited by the number of pollinator visits. Reproductive success increased with increasing inflorescence size, although not above theoretical predictions. However, the probability of producing no fruit or contributing no pollinia decreased with increasing inflorescence size since sequential flowering increased the probability of a pollinator visit to the inflorescence over the blooming period. Large inflorescences did not provide a greater pollinator attraction than small ones, because inflorescences only presented a few open flowers at a time. In addition, flowers on plants growing in clumps of 2–8 plants had a higher probability of setting fruit, apparently because of increased pollinator attraction. Although there are obvious selective advantages for large inflorescences, the sequential flowering habit, and low resource availability may reduce the advantages of large inflorescence size at our study site.     

Do local conspecific density and floral display size influence fruit set via pollinator visitation in Orchis militaris?

Plant density varies naturally, from isolated plants to clumped individuals, and this can influence pollinator foraging behaviour and plant reproductive success. In addition, the effect of conspecific density on reproduction may depend on the pollination system, and deceptive species differ from rewarding ones in this regard, a high density being often associated with low fruit set in deceptive plants. In our study, we aimed to determine how local conspecific density and floral display size (i.e. number of flowers per plant) affect fruit set in a deceptive orchid (Orchis militaris) through changes in pollinator visitation. We measured fruit set in a natural population and recorded pollinator abundance and foraging behaviour within plots of different O. militaris densities. Detailed data were recorded for the most abundant potential pollinators of O. militaris, i.e. solitary bees. Floral display size was negatively correlated to fruit set in medium‐density plots, but uncorrelated in low‐ and high‐density plots. Plot density had no effect on solitary bee abundance and visitation, which may be due to low pollinator abundance within the study site. The proportion of visited flowers per inflorescence was negatively influenced by floral display size, which is in line with previous studies. In addition, solitary bees spent decreasing time in successive flowers within an inflorescence, and the time spent per flower was negatively affected by ambient temperature. Our results suggest that pollinator behaviour during visitation is poorly linked to pollen deposition and reproductive success in O. militaris.     

REPRODUCTIVE BIOLOGY OF ASCLEPIAS EXALTATA

An analysis of the relationships between plant size and survivorship and reproductive success was carried out by sampling four populations of the herbaceous perennial milkweed Asclepias exaltata in Virginia from 1980 to 1982. The annual survivorship rate (about 65%) is the lowest measured for any species of Asclepias. Survivorship was strongly size-dependent but showed no clear relationship with previous history of fruit production. Non-flowering plants were significantly smaller than flowering plants and showed very strong (r > 0.87) correlations between root dry weight and stem or leaf dry weight. Flowering plants were similar to nonflowering plants in root: shoot ratio (approximately 1:1) but differed in that root dry weight was not strongly correlated with stem or leaf dry weight. Components of inflorescence size were strongly correlated within a given level of comparison (e.g., stems per plant with flowers per plant) but less strongly correlated between levels (e.g., stems per plant with flowers per stem). Number of fruits per plant and percentage fruit-set were positively correlated with every component of inflorescence size. Although overall fruit-set was low (about 2%), fruits that were initiated had a high probability of surviving to maturity. There was no evidence of an early period of high fruit abortion: a relatively constant proportion of fruits aborted between each age class.     

Temporal and spatial variation in flower and fruit production in a food-deceptive orchid: a five-year study

Deceptive orchids are generally characterized by low levels of fruit set; however, there may be substantial variations in fruit set between sites and years. Within a single population, individual plants may also differ greatly in their reproductive output as a result of differences in inflorescence size or local density. In this study, we determined flower and fruit production over 5 years in two populations of the food-deceptive orchid, Orchis purpurea . All plants were monitored annually for survival and flowering at each site to determine whether flowering and fruiting induced costs. The number of flowers per inflorescence varied considerably from year to year (min: 36.6, max: 49.5). Average fruit set was low (7%) and varied considerably among years and populations. A considerable proportion of plants also failed to set any fruit. However, the probability of producing at least one fruit was not affected by inflorescence size or local density. The number of fruits was significantly related to inflorescence size, but proportional fruit set was not. Local density also did not affect the number of fruits, nor proportional fruit set. There was also no evidence that plants with large inflorescence size or high fruiting success had a larger probability of remaining vegetative the year after flowering than plants with small inflorescence size or low fruiting success. Our results suggest that pollinator-mediated selective forces on inflorescence size through female reproductive success alone are weak, most likely because of the low overall level of visitation and the resulting uncertainty of pollination at the individual level. Our results further demonstrate that investigation of patterns of fruit set over several years is needed to better understand the variability in female reproductive success that is typical of most plant–pollinator interactions.     

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 effect of inflorescence display size and flower position on pollination success in two deceptive and one rewarding orchid

• Inflorescence display size and flower position on the inflorescence play important roles in plant reproduction, in the formation of fruits and are primarily linked to pollinator behaviour. We used three orchids to determine how visitation rates and choice of pollinator depend on number and position of the flowers along the inflorescence.
• We measured reproductive success in (1) natural conditions, (2) hand-pollination experiments and (3) an experimental design, by modifying composition of inflorescences in populations of two deceptive orchids, Orchis anthropophora and O. italica, and one rewarding orchid, Anacamptis coriophora subsp. fragrans.
• There were no differences in natural fruit production in relation to flower position on the inflorescence (i.e. upper versus lower part), suggesting no preference of pollinators for different parts of the inflorescence. Hand-pollination experiments highlighted low pollen limitation in A. coriophora subsp. fragrans but high limitation in O. italica and O. anthropophora. Reproductive success of deceptive orchids in experimental plots decreased significantly when flowers on the upper half of the inflorescence were removed leading to reduced floral display, while reproductive success of the nectariferous species did not differ significantly.
• Our data highlight that in the examined orchids there is no clear relationship between fruit formation and flower position along inflorescences. Thus we can affirm that, for orchids, the entire inflorescence plays a dominant role in insect attraction but the part of the flower spike does not influence the choice of the insect. This implies that all flowers have the same possibility of receiving visits from pollinators, and therefore each flower has the same opportunity to set fruit.

     

Causes of spatial patterns of fruit set in waratah: Temporal vs. spatial interactions between flowers on an inflorescence

Spatial patterns of fruit set within inflorescences may be controlled by pollination, nutrient allocation, or inflorescence architecture. Generally, flowers that have spatial and/or temporal precedence are more likely to set fruits. We sought to separate these factors by comparing patterns of fruit set on inflorescences of two species of Telopea (Proteaceae); one that flowers from the tip to the base of the rachis, the other from base to tip. In both species, most fruits were set at the top of the inflorescence (the last flowers to open for T. speciosissima) and this was extreme for T. mongaensis, where the top flowers open first. Fruit set was not generally limited by inadequate pollination for either T. mongaensis or T. speciosissima, as hand pollinations did not increase fruit set and many abscised flowers contained pollen tubes. In T. speciosissima, we tested whether removal of developing topmost fruits would ‘release’ those that had initiated but not yet aborted lower down. There was no significant effect. Plant hormones can increase the degree to which a developing fruit is a sink for nutrients, so we applied cytokinin to the developing lower fruits on some inflorescences. There was no significant effect of the hormone treatment. We conclude that temporal precedence may contribute to the skewed pattern of fruit set in T. mongaensis, because there was an extreme concentration of fruit set on the distal part of the inflorescences, but it cannot explain this pattern of fruit set in T. speciosissima, where the distal flowers are the last to open. Some other process must therefore constrain fruit set to the topmost flowers in an inflorescence. While cytokinin application had no significant effect, the power of this experiment was low and we consider that the hypothesis of hormonal control is worth further exploration.     

WHY HERMAPHRODITIC PLANTS PRODUCE MANY MORE FLOWERS THAN FRUITS: EXPERIMENTAL TESTS WITH AGAVE MCKELVEYANA

Five hypotheses explaining the low fruit-to-flower ratios in self-incompatible hermaphroditic plants are tested for Agave mckelveyana. Results from pruning, bagging, and hand-pollination experiments indicate that “excess” flowers (i.e., flowers that fail to produce mature fruits) do not contribute to female fitness (i.e., fruit or seed production). These flowers are aborted regardless of their pollination history. Additional data on nectar production imply that these flowers act as pollen donors and contribute to male fitness.     

An exception to Darwin's syndrome: floral position,protogyny, and insect visitation in Besseya bullii (Scrophulariaceae)

Darwin pointed out that plants with vertical inflorescences are likely to be outcrossed if the inflorescence is acropetalous (flowers from the bottom up), the flowers are protandrous (pollen is dispersed before stigmas are receptive), and pollinators move upward on the inflorescence. This syndrome is common in species pollinated by bees and flies, and very few exceptions are known. We investigated flowering phenology and pollinator behavior in Besseya bullii (Scrophulariaceae) and found that it did not fit Darwin's syndrome. The vertical inflorescence was acropetalous but the flowers were distinctly protogynous, so flowers with newly receptive stigmas appeared on the inflorescence above those with dehiscing anthers. A number of small insects visited B. bullii; bees in the family Halictidae (Augochlorella striata and Dialictus spp.) were most common. When insects moved between gender phases within inflorescences, they moved up more often than down (61% versus 39% of observations, respectively) but this difference was only marginally significant. Most visits were to male-phase flowers only, and this preference was more pronounced for pollen-foraging insects than for nectar-foraging insects. B. bullii was self-compatible, so its flowering characteristics potentially could result in considerable self-pollination. However, an average of 38% of the lowermost flowers opened before any pollen was available on the same inflorescence; these solo females had a high probability of outcrossing (though fruit set was relatively low in the bottom portion of the inflorescence). Upper flowers may also be outcrossed because downward insect movement was not uncommon. Therefore protogyny in B. bullii may not necessarily lead to more selfing than would protandry.     

The effects of herbivory and resource variability on the production of a second inflorescence by the desert lily, Pancratium sickenbergeri

We investigated the effects of herbivory by the dorcas gazelle, Gazella dorcas, on the production of a second inflorescence in a desert lily, Pancratium sickenbergeri (Amaryllidaceae), after the first inflorescence was eaten. In three populations exposed to different levels of herbivory, we conducted an inflorescence-clipping experiment in the flowering seasons of 1997, 1998 and 1999. The experiment was performed on inflorescences at both the emerging stage and the anthesis stage. Plants in each control group were undamaged. Clipped plants had a greater probability of producing a second inflorescence stalk than unclipped plants. In all the populations, the probability of producing a second inflorescence was greater when the first inflorescence was cut in the emerging stage than when it was cut in anthesis, indicating that there was a cost of compensation. In the population with the highest level of herbivory, the production of a second inflorescence stalk was related to resource availability in all 3 years. Plants produced as many flowers or fruits on the second stalk as on the first, indicating that potential reproductive output is double that usually achieved. Although we found no evidence for a trade-off between investment in first and second inflorescence stalks, the lower probability of producing a second inflorescence if heavy investment had been made in the first inflorescence indicates that compensation is costly. These results contradict the mutualism hypothesis that herbivory is beneficial to plants and provide support for the trade-off hypothesis that benefits of herbivory are proximal and are attained at an evolutionary cost.     

The role of termites in the decomposition of wood and leaf litter in the Southern Guinea savanna of Nigeria

Summary The number of flowers produced by inflorescences of Yucca whipplei (Agavaceae) consistently exceeds the number of fruits produced by about one order of magnitude. To determine the factors responsible for low fruit set, the relation between pollinator availability, the amount of resources spent on reproduction (as indicated by inflorescence size), and the number of fruits matured was studied during 1978 and 1979 at 18 locations in chaparral, coastal sage scrub, and desert scrub communities of southern California.The following results support the conclusion that pollinators do not usually limit fruit production in Yucca whipplei. Rather, fruit production is limited by the amount of resources available to support developing fruits. (1) Fruit production is positively correlated with inflorescence size both within and between populations. The average size of inflorescence for a population is an excellent predictor of mean fruit production. Furthermore, 54% of the total variance in fruit production of individual plants can be explained by inflorescence size. (2) In contrast, although fruit production within most populations is positively correlated with an index of the number of pollinator visits to an inflorescence, the relative abundance of pollinators for a population is a poor predictor of mean fruit production, and only 9% of the total variance in fruit production can be explained by the visitation index. Furthermore, at four sites studied for two years, there was little change in average inflorescence size or fruit production from 1978 to 1979, despite large differences in relative abundance of pollinators at each of the sites. (3) Based on geographic proximity, and physiographic and vegetational similarities, study sites were grouped into regional clusters. Both inflorescence size and fruit production varied considerably between regions. Of the total variation in fruit production, 27% can be attributed to differences between regions. Most of this variation is the result of regional differences in inflorescence size, which in turn influence fruit production.Why does Yucca whipplei produce such large inflorescences if so few fruits can be supported? Two relevant hypotheses are discussed: (1) the floral display is the result of selection for pollen dissemination at the expense of fruit set; and (2) the floral display is the result of selection for a bet-hedging strategy either to increase the probability of adequate pollination when pollinators are unusually rare, or to allow individuals to support more fruits when resources are unusually abundant.     

Regulation of the mutualism between yuccas and yucca moths: intrinsic and extrinsic factors affecting flower retention

Yuccas initiate far more flowers than they can mature as fruit, thereby providing opportunities for them to mature flowers of the highest quality. Flower quality in yuccas has both intrinsic and extrinsic components. Intrinsic components relate to flower morphology and inflorescence architecture. Yucca moths (Tegeticula spp., Incurvariidae), the sole pollinators and primary seed predators of most yuccas (Yucca spp., Agavaceae), mediate extrinsic components of flower quality through their ovipositions in flowers, and the quantity and quality of pollen that they transfer. In addition, intrinsic and extrinsic components interact as a function of flowering phenology and moth activity within inflorescences.
We investigated selective abscission of flowers in Y. kanabensis with respect to various combinations of intrinsic and extrinsic factors. First, we considered the effect of high and low pollen loads delivered to different subsets of flowers and in different presentation orders. In the absence of moth ovipositions, Y. kanabensis is sensitive to the amount of pollen that moths deliver and tends to retain high pollen flowers, even when all flowers receive sufficient pollen for full fertilization. However, pollen delivery sequence and the position of flowers with an inflorescence modify this high pollen effect. We then considered the interplay between high and low pollen combined with moth ovipositions and found that the number of ovipositions dominated the pollen effect. Finally, we considered number of ovipositions in conjunction with different flowers in the blooming sequence while controlling pollen levels and found that the clear effect of ovipositions on flower fate can be tempered by where the flower is in the blooming sequence.
These results have implications for the regulation of the mutualism between yuccas and yucca moths, indicating that yuccas are capable of regulating costs, retaining flowers of relative high quality and selectively abscising the rest. Yucca sensitivity to several intrinsic and extrinsic factors allows the plant to respond flexibly to the pollination environment and several species of moths.     

Fruit set,fruit reduction,and fruiting strategy in Cornus sanguinea (Cornaceae)

We studied various aspects of the fruiting biology of Cornus sanguinea (Cornaceae), a fleshy-fruit-producing deciduous shrub, in four populations in northwest Spain. One population was studied over a 5-yr period (1989-1994), and the remaining populations in 1994 only. Fruit-set level varied among years (range 11-18%) and among populations (range 8-22%), but was in all cases low. Within plants, fruit-set level did not vary significantly among inflorescences, indicating that inflorescence fruit set is independent of inflorescence position and inflorescence, phenology. To investigate the function of surplus flowers, we carried out flower removal experiments. Inflorescence fruit-set level was unaffected by removal of up to =75% of flowers. These results suggest that Cornus sanguinea regulates its fruit-set level via plasticity in the number of fruits aborted: if flower mortality has been high, fewer fruits will be aborted. Within the inflorescence, surplus flowers thus act as insurance against flower loss.     

Bumblebee-pollination and temporal change of the calyx tube length in Clematis stans(Ranunculaceae)

Clematis stans is dioecious semi-arboreal, with pale purple–blue, nodding, tubulous flowers in a paniculate inflorescence. Both male and female flowers produce nectar from the base of the calyx tube during a flowering period of 3 or 4 days, and are pollinated by two bumblebee species, Bombus diversus and B. honshuensis, with different proboscis lengths. When the flowers open, four sepals constructing a calyx tube separate at the top and their respective tips gradually curl up, so that a tubular part shortens. Observations at two field sites showed that B. diversus (with a longer proboscis) most often visits the flowers with a longer calyx tube, and B. honshuensis (with a shorter proboscis) the flowers with a shorter calyx tube, i.e., later in the flowering period. By changing the calyx tube length, the flowers of C. stans accept the two bumblebee species with different proboscis length as pollinators and thus increase the chance of pollination for each flower. It was also found that the two bumblebee species prefer the male flowers to the female flowers, although the female flowers secrete more nectar as a reward than male flowers. This is likely because they visit the male flowers to collect pollen grains in addition to nectar. Electronic Publication     

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.     

NEW INTERPRETATION OF THE INFLORESCENCE OF FAGUS DRAWN FROM THE DEVELOPMENTAL STUDY OF FAGUS CRENATA,WITH DESCRIPTION OF AN EXTREMELY MONSTROUS CUPULE

The inflorescence of Fagus is generally considered to be a determinate one, i.e., an axillary dichasium, in contrast to those of most genera in the family, which are indeterminate, dichasial, or simple catkins. To understand the relationship between the two types, ontogenetic development of the inflorescence of Fagus crenata was investigated. The early developmental stages are similar in both the male and the female inflorescences. At first, the inflorescence is oval-shaped, then a swelling forms at the distal side of it. Subsequently, another swelling forms at the proximal side. The more or less conspicuous residual part of the primary inflorescence axis remains between the two swellings. The inflorescence becomes heart-shaped and the first flower forms at the summit of each swelling. Subsequently, higher-ordered flowers form dichasially in the male inflorescence, and the cupule valves differentiate in the female one. This organogenetic manner suggests that the inflorescence of Fagus is an indeterminate one, consisting of two dichasia arranged alternately on the primary axis. The scale leaves surrounding the inflorescence were also given a new interpretation. They were considered to be stipules of the bracts, because sometimes they constitute a continuous structure, together with an inconspicuous swelling between them. A proliferous-type monstrous cupule was interpreted as supporting evidence for the hypothesis.     

Selective fruit filling in relation to pollen load size in Alstroemeria aurea (Alstroemeriaceae)

 We conducted an experiment in a natural population of Alstroemeria aurea, a clonal perennial, to determine (1) if reproduction was resource limited, and (2) if fruits would be selectively filled based on differences in pollination intensity when pollen loads were adequate for full seed set. Under these conditions, differences in pollination intensity are unlikely to affect seed number, but could affect seed quality, providing an interesting test of the gametophytic competition hypothesis. To test for resource limitation, percent fruit maturation, number of seeds per fruit and average seed weight were compared to paired controls for ramets in which all but one fruit was removed. To test the effect of pollination intensity on selective resource allocation, three types of pollination treatments were performed: (1) all flowers of the single inflorescence received a low pollen load, (2) all flowers received a high pollen load, (3) alternate flowers of the inflorescence received either a high or a low pollen load. We determined the percentage of fruit that reached maturity, counted the number of seeds and ovules and calculated the average seed weight for all capsules in each treatment. Resources appeared to limit reproduction in this population since seed number and weight were significantly higher than in controls when competing capsules were removed. At the whole ramet level, a four fold difference in pollen loads had no significant effect on any of the parameters measured. However, when pollination intensity varied within an inflorescence, the number of seeds per fruit increased by about 10% in flowers that received the higher pollen load. We observed the same trend in each of 2 years, but the increase was significant in only 1 year. The differences, although not great, were only slightly smaller than when all competing fruits were removed, and were consistent with selective resource allocation based on pollination intensity independent of seed set. Received: 28 September 1997 / Accepted: 30 April 1998