Ants in your plants: effects of nectar-thieves on pollen fertility and seed-siring capacity in the alpine wildflower, Polemonium viscosum

Flowers of the alpine skypilot, Polemonium viscosum , are attacked by nectar thieving ants of Formica neorufibarbus gelida . Ants exert selection on flower scent, size and shape in skypilots by damaging the pistils. Here, I report on the frequency and nature of contact between ants and pollen-bearing anthers and determine the consequences of such contact for pollen performance and pollen donor paternity. In laboratory trials, ants entered flowers with full intact anthers and emasculated (female) flowers equivalently. Similarly, flower visitation rates of ants foraging naturally were not affected by the frequency of male phase flowers per plant. Ants actively interacted with the pollen-bearing anthers during 21% of flower visits, on average. The rate at which such interactions occurred was predicted by the proportion of flowers in the male phase, under a random foraging model. The effect of ants on pollen fertility was tested experimentally by enclosing ants in male-phase flowers on intact inflorescences. Adjacent control flowers were left un-occupied. Pollen from flowers with a history of ant occupancy had significantly lower germination on virgin recipient stigmas than pollen from unoccupied control flowers. With hand-pollination, sufficient pollen was transferred from ant-occupied flowers to saturate seed set. However, a model based on the relationship between seed set and compatible pollen delivery by natural pollinators indicated that ant damage to pollen should reduce paternity accruing per flower visit by 20–26% on average, in nature. Results support the hypothesis that in P. viscosum , selection on floral traits by nectar-thieving ants operates through male as well as female function.     

Pollen precedence and stigma closure: a mechanism of competition for pollination between Delphinium nelsonii and Ipomopsis aggregata

Summary Previous experiments showed that the sympatric herbs Delphinium nelsonii and Ipomopsis aggregata compete for hummingbird pollination and that deleterious effects of the former species on seed set of the latter involve interspecific pollen transfer. However, seed set was not reduced when pollen of both species was applied simultaneously to I. aggregata stigmas. Hence a competitive effect may require arrival of foreign pollen before conspecific pollen. To explore this possibility we subjected I. aggregata flowers to a competition treatment in which they received D. nelsonii pollen 6 h before I. aggregata pollen, or to a control in which they received only the conspecific pollen. Foreign pollen precedence decreased mean seed set by almost 50%, which is consistent with effects observed in previous experiments. Reduced seed set can be explained by the fact that foreign pollen often caused stigma lobes to close together within 1.5–6 h, reducing subsequent receptivity. Stigma closure was also elicited by conspecific pollen, but not by mechanical stimulation, and was influenced by size of the pollen load and identity of the plant being pollinated.     

Reproduction in Polemonium: factors influencing outbreeding potential

Summary Three measures of potential outbreeding efficiency were quantified throughout two flowering seasons in a population of the mass-flowering Polemonium foliosissimum: 1) the distances flown between plants by bumblebee pollinators; 2) the proximity of visited plants; and 3) the diversity of individuals visited. Results were consistent between years. Individual plants did not function differentially with respect to female (i.e., pollen receipt) and male (i.e., pollen donation) function. Both female and male components of outbreeding, as estimated by all three measures, were positively correlated with the number of flowers per plant. Significantly more outbreeding could occur during the final phase of flowering when most individuals had relatively few blossoms. Although the potential for outbreeding was the same at any time through an individual's blooming regime, individuals for which the peak of blooming occurred after the peak for the population as a whole have significantly more outbreeding potential via both female and male function. These factors may combine such that any pressure to increase outbreeding in this self-incompatible species manifests itself in directional selection for delayed flowering time.     

Distance-dependent pollen limitation of seed set in some insect-pollinated dioecious plants

By applying hand pollination, we studied pollen limitation of seed set of female plants in four dioecious plant species with insect-pollination. The effect of hand pollination increased with distance to the nearest male plant. Distances at which seed set was 50% of its maximal value (after hand pollination) were: 2.3 m for Valeriana dioica, 5.3 m for Salix repens, 8.5 m for Asparagus officinale and 10.6 m for Bryonia dioica. We discuss to what extent the reduced seed set was caused by either fewer visits or by visits of a lower quality. We argue that quantifying distance-dependent seed set in dioecious plants may be a good way of studying effects of habitat fragmentation on the breakdown of mutualistic pollination systems.     

The significance of pollination time and frequency and of purity of pollen loads for seed set in Rhinanthus angustifolius (Scrophulariaceae) and Viscaria vulgaris (Caryophyllaceae)

Summary The effects of time of the day and frequency of application and of purity of artificially deposited pollen loads on fruiting and seed set were studied in Rhinanthus angustifolius (diurnally visited by bumblebees) and Viscaria vulgaris (diurnally visited by bumblebees, nocturnally by sphingid moths).Time of the day did not influence pollination success in either species. Increase of pollination frequency increased fruiting and seed set in Rhinanthus but had no affect on Viscaria. Five successive artificial pollinations were needed to achieve seed set equal to that observed naturally in Rhinanthus while a single artificial pollination was sufficient in Viscaria. Mixing Lupinus and Viscaria pollen did not reduce fruiting and seed set in Viscaria. The results are discussed in relation to observed seed sets in early and late flowers, and small and large patches of Viscaria vulgaris and among Rhinanthus flowers in populations of different densities.     

Seed set after pollination with in-vitro-matured,isolated pollen of Triticum aestivum

Summary Immature pollen of two varieties of Triticum aestivum, at the stage right after the first pollen mitosis, was isolated from individual anthers and cultured in microcultures of microliter droplets. In a specifically designed medium, some of the pollen grains developed to maturity. These were applied to excised stigmas on agar, where they produced pollen tubes. Application to flowers in vivo led to seed set. Pollen was matured in vitro from a variety that produced a different protein banding pattern on SDS-PAGE as compared to the variety that was pollinated. The protein banding in the produced seeds showed the hybrid pattern, demonstrating that the seeds were not produced by self-pollination in this in-breeding species but by pollination with the in-vitro-matured pollen.     

Interspecific competition for seed production between two naturalized dandelions under different levels of fertilizer supply

Two naturalized dandelions,Taraxacum officinale andTaraxacum laevigatum, were grown in mixed stands at various plant densities and mixing ratios with various levels of fertilizer supply until seeds were produced. The effect of fertilizer supply on the competitive relationship for seed production between the species was analyzed. In pure stands, the simultaneous effects of plant density and fertilizer supply on mean plant weight were formulated for each species. By introducing the density conversion factor, the results of plant weight growth in mixed stands were formulated successfully.T. laevigatum produced more seeds thanT. officinale at the same plant weight. The relationship between mean plant weight and mean number of mature seeds produced per plant was also formulated for each species. Ratio diagrams were predicted with these formulas. The density conversion factors showed thatT. officinale was always dominant overT. laevigatum for plant weight growth within a generation. The ratio diagrams showed thatT. officinale became dominant overT. laevigatum for seed production with repeating generations at higher levels of fertilizer supply, but that the species with a larger ratio of plant density became dominant over the other species at lower levels of fertilizer supply.     

Within-population spatial variation in pollinator visitation rates, pollen limitation on seed set, and flower longevity in an alpine species

Pollen limitation through insufficient pollen deposition on stigmas caused by too infrequent pollinator visitation may influence the reproductive outcome of plants. In this study we investigated how pollinator visitation rate, the degree of pollen limitation, and flower longevity varied spatially among three sites at different altitudes within a population of the dwarf shrub Dryas octopetala L. in alpine southern Norway. Significant pollen limitation on seed set only occurred at the mid-elevation site, while seed set at the other sites appeared to be mainly resource limited, thus indicating a spatial variation in pollen limitation. There was no association between the spatial variation in the extent of pollen limitation and pollinator visitation rate to flowers. However, pollinator visitation rates were related to flower longevity of Dryas; sites with low visitation rates had long-lived flowers and vice versa. Thus, our results suggest within-population spatial co-variation between pollinator visitation rates, pollen limitation, and a developmental response to these factors, flower longevity.     

Pollination by deceit,floral sex ratios and seed set in dioecious Rubus chamaemorus L.

Summary Male and female flowers of the dioecious perennial herb Rubus chamaemorus L. are similar in general appearance. However, female flowers are somewhat smaller, do not produce any pollen, and contain very small amounts of nectar. Syrphids and bumblebees, which are important pollinators of R. chamaemorus, showed a strong preference for male flowers. Male flowers were also less often rejected by flower visitors than were female flowers, and two different groups of syrphid species stayed longer in male than in female flowers. These observations suggest that female flowers of R. chamaemorus attract pollinators by deceit.Hand-pollination experiments indicated that pollen availability limited seed production of R. chamaemorus in female dominated habitats but not in areas with an equal floral sex ratio. We suggest that the relative importance of factors limiting female reproductive success is not constant, but is influenced by the floral sex ratio of the population. This should apply also to other dioecious species that show variable sex ratios on either a local or regional scale.     

The effect of petal-size manipulation on pollen removal,seed set,and insect-visitor behavior in Campanula americana

We investigated the effect of petal-size variation in tall bellflower, Campanula americana, on pollen removal rates, seed set, and pollinator behavior. Pollen removal, seed set, and the behavior of two bee visitors (Bombus and Halictus) were assessed in flowers that had 0%, 50%, or 100% of their petal lobes removed. Pollen removal rates did not differ significantly among the three treatment groups, probably due to the presence of an ineffective (ugly) pollinator (Halictus), which quickly removed pollen from completely reduced flowers. Seed set was significantly positively correlated with petal size. Bumblebees discriminated against reduced flowers, thereby lowering pollen movement to reduced flowers. Halictid bees rarely contacted stigmatic surfaces and thus pollen delivery to reduced flowers was limited. The presence of an ugly pollinator probably obscured the effect of petal reduction on pollen receipt, thus masking the impact of attractive structures on male fitness. Given the positive relationship between seed set and petal size, we caution against ascribing expenditure on attractive structures exclusively to male function. In plants with many olules, expenditure on attractive structures may affect seed set.     

Pollen-limited seed set in pseudogamous blackberries (Rubus L. subgen. Rubus)

Summary Pseudogamous blackberry species are polyploid and usually exhibit meiotic irregularities causing severe reduction in pollen viability. When species means were compared, relative seed set was strongly correlated with the number of good pollen grains produced per flower divided by the number of ovules (good-pollen/ovule ratio).Partial correlation analysis for percentage viable pollen and relative seed set, controlling for good-pollen/ovule ratio, revealed an almost significant relationship, presumably because meiotic irregularities have a similar effect on both pollen viability and the viability of meiotically derived embryo sacs. Seed germination, on the other hand, showed no relationship with relative seed set or with pollen viability.The automatic selfing capacity exhibited by most of these blackberry species does not appear to be related to either pollen production or relative seed set.     

Pollination in vitro: effects on the growth of pollen tubes,seed set and gametophytic self-incompatibility in Trifolium pratense L. and T. repens L.

Summary Growth of pollen tubes and seed set were compared after hand pollination in situ and in vitro in two self-incompatible species, Trifolium pratense and Trifolium repens. Adhesion of pollen grains to the stigma was greater in vitro for both species. After cross-pollination, in vitro culture gave a significant increase in the cumulative growth of pollen tubes in pistils of T. pratense compared to in situ conditions. After selfing in T. repens, pollen tube growth was significantly increased by in vitro culture of florets. Seed set after crossing in situ and in vitro was similar for both species. Seed set after selfing in vitro was not increased in T. pratense. Several genotypes of T. repens were classified as very good, good and poor selfers based on their capacity for seed set following selfing in situ. In vitro pollination increased self seed formation by 1.7-, 18.0- and 31.0-fold for each class, respectively. Ovules located nearest to the style were fertilized more often after selfing than after crossing.     

Dosage-dependent impacts of a floral volatile compound on pollinators, larcenists, and the potential for floral evolution in the alpine skypilot Polemonium viscosum

All volatile organic compounds (VOCs) vary quantitatively, yet how such variation affects their ecological roles is unknown. Because floral VOCs are cues for both pollinators and floral antagonists, variation in emission may have major consequences for costs and benefits in plant-pollinator interactions. In Polemonium viscosum, the emission rate for the floral VOC 2-phenylethanol (2PE) spans more than two orders of magnitude. We investigated the ecological and evolutionary impacts of this immense phenotypic variation. The emission rate of 2PE varies independently of nectar rewards and thus is uninformative of profitability. Emission is elevated in flowers that are morphologically vulnerable to ant larcenists, suggesting that chemical deterrence may compensate for weak physical barriers. In nature, plants emitting more 2PE than their neighbors escape ant damage. Flower-damaging ants die when exposed to 2PE in the laboratory, and they avoid high 2PE emitters in the field. High 2PE also reduces bumblebee visitation and pollination, suggesting an ecological cost of defense in pollinator service. However, at more moderate emission rates, 2PE enhances the amount of nectar left in flowers, at no pollination cost. In conclusion, repellency of 2PE is highly sensitive to dosage, giving it a key role in shaping ecological interactions between skypilot plants and their floral visitors.     

Flowering phenology,pollen flow and fruit production in the andean shrub Befaria resinosa

Summary In the eastern Andes of Colombia, the shrub Befaria resinosa (Ericaceae) has peaks of flowering that are separated by extended periods of low flower production. The effect that these fluctuations in flower production have on pollen flow was investigated by using fluorescent dye as a pollen analog. Dye applied to open flowers was dispersed over long distances more often during low flower production than during high flower production. Whether enhanced pollen dispersal during flowering lows is of benefit to individual plants is not clear. The proportion of flowers that set fruit is positively correlated with flower abundance, negating the possibility that increased pollen dispersal results in a higher rate of fruit production due to outbreeding effects. It is also difficult to attribute the pattern of fruit production to changes in pollinator visitation rates, which are negatively correlated with flower abundance in the case of hummingbirds and not correlated at all with flower abundance in the case of insects. An opportunistic, large-bodied hummingbird (Colibri coruscans) visits B. resinosa during high flowering and may be a particularly effective pollinator, accounting for some of the increase in the proportion of flowers setting fruit. Rainfall is positively correlated with flower production and may be an important factor in shaping flowering phenology, but it is not significantly correlated with the proportion of flowers setting fruits. The possibility that low-level flowering may counteract inbreeding that results from peak flowering is discussed.     

Effect of pollination intensity on fruit and seed set in cacao (Theobroma cacao L.)

We studied the functional relationship between pollination intensity and fruit survival as well as the number of seeds per pod in the tropical tree Theobroma cacao L. on a Forastero Upper-Amazon clone (UPA 409) in Ivory Coast. Cutting the style 24 h after pollination allowed for counting the number of pollen grains deposited on a stigma without affecting fruit set and seed development. Forty-three pollen grains were necessary to reach 50% of maximum fruit set 28 days after pollination. Above 115 pollen grains, the proportion of developing ovaries reached a maximum of 88% 28 days after pollination and 75% at maturity. With fewer than 238 pollen grains per stigma, there was a close relationship between pollination intensity and number of seeds per pod; the pollenseed ratio increased from 1.61 to 3.81 for PI increasing from 30 to 238 pollen grains. For higher pollination intensities, the average number of seeds per pod reached a maximum of 58. The relationship between pollination intensity and seed content was modelled. Results are consistent with the hypothesis that ovules attracted pollen tubes in a similar way regardless of whether or not they had already been reached by another pollen tube.     

Effects of parentage,prior fruit set and pollen load on fruit and seed production in Campanula americana L

Summary We conducted a controlled crossing experiment to examine the effects of maternal and paternal parentage, the size of the pollen load, and prior fruit production on the proportion of flowers that set fruit, seed number per fruit and seed weight in a natural population of Campanula americana. Effects due to the maternal parent were large for all measures of fruit and seed production, while the paternal parent had a significant effect only upon mean seed weight. As the number of prior fruits on the maternal plant increased the probability that a flower would produce a mature fruit, the number of seeds per fruit, and total seed weight per fruit all decreased. We found no effect of the size of the pollen loads used in this study on fruit or seed production. These results are consistent with those of other studies that suggest in natural plant populations maternal effects, especially environmental maternal effects, can have an overwhelming effect on fruit and seed production and on seed characteristics.     

Gigantism and the potential for interference competition in the rotifer genus Asplanchna

Summary Laboratory experiments showed that A. bright-welli could outreproduce, or coexist with, A. silvestrii only when the absence of dietary tocopherol prevented the latter from transforming to the giant cruciform and campanulate morphs. When tocopherol permitted polymorphic transformations, as often occurs in nature, the giant morphs of A. silvestrii ingested and rapidly excluded the much smaller, only slightly polymorphic A. brightwelli. Such interference (or encounter) competition from trimorphic Asplanchna species is known to occur in nature and must limit the distribution and abundance of monomorphic or only slightly polymorphic species. The ability to eat congeneric competitors may have provided some selective pressure for the evolution of gigantism in the genus.     

Herbivory and intraspecific competition in a stream caddisfly population

Summary Field experiments were conducted to assess the effects of different population densities of the herbivorous caddisfly Helicopsyche borealis on periphyton biomass and on its own growth rate in Big Sulphur Creek, a third-order stream in northern California, USA. Stream enclosures were used to vary grazer density from one-eighth to twice natural density (1/8X–2X) in two experiments (35 d and 60 d), which spanned the period of most rapid larval growth. Periphyton biomass and chlorophyll a were inversely related to grazer density. Grazer densities of 1/8X–1/2X moderately reduced periphyton when compared to an ungrazed control, whereas densities of 1X–2X greatly depleted periphyton. The growth rate of H. borealis larvae declined with increasing larval density. Growth rate was highest at densities of 1/8X–1/2X; larvae grew more slowly at 1X and showed no growth at 2X. Weekly supplementation of periphyton to 1/4X, 1X, and 2X densities significantly increased the final body mass of larvae at 1X and 2X (by 26% and 50%, respectively, compared to unsupplemented larvae) but did not change the body mass of larvae at 1/4X. These results suggest that periphyton is a limiting resource in Big Sulphur Creek and that H. borealis larvae compete exploitatively for that resource. Intraspecific competition may be an important, but often overlooked, feature of many herbivore populations in streams.