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The article discusses a study by L. Fishman and J. H. Willis on pollen limitation and natural selection on floral characters in the yellow monkeyflower. Fishman and Willis applied a factorial design of pollen supplementation and pollinator exclusion to plants in an Oregon population of Mimulus guttatus and found that pollen supplementation increased seed production and pollinator exclusion reduced it. The findings imply that Mimulus guttaus plants with different pollinator abundances will be subject to strongly divergent types of selection. According to the study, in declining pollinator populations, the flowers themselves may begin to evolve to be less attractive and less reliant on pollinators, which might lead to declines.

Flowers exhibit tremendous variation in size (>1000-fold), ranging from less than a millimeter to nearly a meter in diameter. Numerous studies have established the importance of increased floral size in species that exhibit relatively normal-sized flowers, but few studies have examined the evolution of floral size increase in species with extremely large flowers or flower-like inflorescences (collectively termed blossoms). Our review of these record-breakers indicates that blossom gigantism has evolved multiple times, and suggests that the evolutionary forces operating in these species may differ from their ordinary-sized counterparts. Surprisingly, rather than being associated with large-bodied pollinators, gigantism appears to be most common in species with small-bodied beetle or carrion-fly pollinators. Such large blossoms may be adapted to these pollinators because they help to temporarily trap animals, better facilitate thermal regulation, and allow for the mimicry of large animal carcasses. Future phylogenetic tests of these hypotheses should be conducted to determine if the transition to such pollination systems correlates with significant changes in the mode and tempo of blossom size evolution. [Copyright &y& Elsevier]

Flowers are phenotypically integrated units that may require a precise configuration of floral organs for proper function. Plants, however, are metameric organisms that often exhibit architectural and temporal variation in floral size and shape. Clearly, architectural and temporal effects could interrupt phenotypic integration within flowers, yet such effects have not been explicitly examined. We compared correlations among floral characters for the closely related but morphologically divergent species Nicotiana alata and Nicotiana forgetiana and their artificial F3 hybrids. Principal components analysis of 16 floral traits identified two suites of correlated characters that are maintained in both taxa and in the hybrids, regardless of architectural and temporal variation in floral characters. We interpret the maintenance of these suites as evidence of common developmental regulation of correlated characters. One suite includes measures of corolla tube, gynoecium, and androecium length, while the other includes corolla limb and corolla tube aperture characters. The correlated suites of characters are tightly integrated, independent from characters of the other factor, and they participate in a common function. As such, these character suites represent phenotypic modules that have been maintained despite evolutionary changes in morphology during the divergence of the two species. [ABSTRACT FROM AUTHOR]

The size and number of flowers displayed together on an inflorescence (floral display) influences pollinator attraction and pollen transfer and receipt, and is integral to plant reproductive success and fitness. Life history theory predicts that the evolution of floral display is constrained by trade-offs between the size and number of flowers and inflorescences. Indeed, a trade-off between flower size and flower number is a key assumption of models of inflorescence architecture and the evolution of floral display. Surprisingly, however, empirical evidence for the trade-off is limited. In particular, there is a lack of phylogenetic evidence for a trade-off between flower size and number. Analyses of phylogenetic independent contrasts (PIC5) of 25 1 angiosperm species spanning 63 families yielded a significant negative correlation between flower size and flower number. At smaller phylogenetic scales, analyses of individual genera did not always find evidence of a trade-off, a result consistent with previous studies that have examined the trade-off for a single species or genus. Ours is the first study to support an angiosperm-wide trade-off between flower size and number and supports the theory that life history constraints have influenced the evolution of floral display. [ABSTRACT FROM AUTHOR]

The structure of the plant inflorescence and flower is an important agronomic and ornamental trait studied for its potential economic applications. In particular, the capacity to modify flower size has always been a breeder’s goal. Genetic and molecular studies have shown that the Zea mays gene Ramosa1 ( Ra1) is involved in inflorescence branching regulation. In fact the ra1 loss of function mutation causes extra branching of the inflorescence. In this work we suggest a possible utilization of the Ramosa1 maize gene as a tool to modify inflorescence architecture and flower size in transgenic plants. In fact overexpression of this gene in Arabidopsis plants promotes an increase in reproductive organ size. Pollen, seeds, cotyledons, leaves and roots are also larger than those of the wild type. Analysis of organs from transformants showed that cell expansion was increased without apparently affecting cell division. These results suggest that the RA1 protein is able to up-regulate cell expansion in all organs of Arabidopsis plants. [ABSTRACT FROM AUTHOR]

1. Plants that display many open flowers usually receive higher pollinator visitation rates, but it is unclear whether pollinators select plants to visit based on the size of floral display (apparent size) or the value of the floral rewards (reward size). To examine how pollinators respond to apparent size and reward size, we observed bumble bees foraging among arrays of artificial plants. 2. We constructed two kinds of artificial flower: (i) rewarding flowers that produced nectar constantly; and (ii) unrewarding flowers that produced only water. Thus, we could construct plants that varied both in numbers of flowers (apparent size) and in numbers of rewarding flowers (reward size). 3. At the beginning of the experiments, bees made more visits to the plants with the most flowers, irrespective of the rewards they contained. However, after several hours of foraging, bees returned selectively to plants with the greater number of rewarding flowers, irrespective of the number of flowers the plant presented. After we replaced rewarding plants with non-rewarding plants, bees continued visiting plants at formerly-profitable locations for a while. 4. Our results demonstrate that bees initially showed preferences for plants with larger floral displays, but eventually bees were able to discriminate between rewarding and less-rewarding plants of equal display size by associating a plant's location with its reward size. Our results suggest that plants with many flowers can achieve higher visitation rates from pollinators in two ways: (i) by attracting inexperienced pollinators with large displays; and (ii) by encouraging experienced pollinators to return with the promise of greater rewards. [ABSTRACT FROM AUTHOR]

We made a 5 × 5 diallel set of crosses between five inbred lines, four Japanese and one Chinese, and diallel analyses of radish floral morphology were conducted to estimate the genetic parameters underlying 13 floral traits. The modes of inheritance of 12 traits (but not anther length on the long stamen) followed the additive-dominance model. Broad-sense heritabilities of the 12 traits were 0.93–0.99, suggesting that floral morphology is highly heritable. But narrow-sense heritabilities differed among traits, ranging from 0.34 to 0.89. Pistil and stamen heights, tube length, and stigma position were explained mainly by additive effects, but anther, stigma, petal, and whole flower sizes were affected greatly by dominance effects as well as by additive effects. The correlations between the 13 traits suggested that some floral traits (e.g. stigma size and position) are more or less independent, indicating the possibility of creating lines with new floral morphologies by selecting for these genetically independent traits. A flower with a larger and higher stigma might improve the production of cross-pollinated seeds through enhancement of pollination efficiency. In that case, by selecting for stigma position in an early segregating generation and for stigma size in a later generation, it should be possible to produce new parental lines with effective floral morphology for F 1 seed production. [ABSTRACT FROM AUTHOR]

Andromonoecy is hypothesized to evolve as a mechanism enabling plants to independently allocate resources to female and male function, If staminate flower production is a mechanism to regulate allocation to female function (i.e., fruit production), then large-fruited species should be more strongly andromonoecious than smaller-fruited taxa because more resources are required to mature large fruit. We combined phylogenetically independent contrast analyses with extensive phenotypic characterization under common greenhouse conditions to examine the predicted relationship between fruit mass and the strength of andromonoecy among 13 species in Solanum sections Acanthophora and Lasiocarpa. The strength of andromonoecy, defined as the proportion of staminate flowers produced within inflorescences, was significantly and positively associated with fruit mass in both naïve and phylogenetically independent analyses. Our results are consistent with the hypothesis that andromonoecy functions as a mechanism to regulate allocation to female function and suggest that the strength of andromonoecy is also associated with resource limitation. In general, we find that strong andromonoecy appears to arise via reductions in hermaphroditic flower number. However, increases in staminate flowers have also contributed to transitions to strong andromonoecy in certain species. Finally, our analyses identified a suite of correlated characters (flower size, ovary width, fruit mass) that are associated with changes in the sexual expression of andromonoecy. [ABSTRACT FROM AUTHOR]

Floral display size represents a tradeoff between the benefits of increased pollinator visitation and the quantity of pollen received vs. the costs of increased self-pollination and reduced pollination quality. Plants with large floral displays often are more attractive to pollinators, but pollinators visit more flowers per plant. Intraplant foraging movements should increase self-pollination through geitonogamy, lowering outcrossing rates in large plants. Local genetic structure should also increase inbreeding and decrease outcrossing estimates, if pollinators move between neighboring, related plants. These predictions were tested in a population of larkspurs (Delphinium barbeyi) in Colorado. Allozymes were used to estimate outcrossing rates of plants varying in display size. Floral displays varied widely (2-1400 flowers; 1-26 inflorescences per plant), and outcrossing rate decreased significantly with increasing display size. Large, multistalked plants self over twice as frequently as single-stalked plants (46 vs. 21%). Local population structure is significant, and biparental inbreeding depresses outcrossing in plants surrounded by genetically similar neighbors. Protandry, coupled with stereotypical bottom-up pollinator foraging, reduces self- fertilization by autogamy or geitonogamy within inflorescences. Selfing is predominantly (>60%) by geitonogamy between inflorescences in large plants. Geitonogamy may be a significant cost to plants with large floral displays if inbreeding depression and/or pollen and ovule discounting results. If so, floral display size, particularly inflorescence number, may be under contrasting selection for pollination quantity vs. quality. [ABSTRACT FROM AUTHOR]

1. From Darwin to the present day, sexual selection has been shown to be widespread in both animals and plants. Attractive floral traits in hermaphrodite plants evolve in response to selection acting simultaneously through male and female sexual functions, but knowledge about the relative strength of gender-specific selection is scarce. 2. We experimentally altered flower size of the hermaphrodite Cistus salvifolius in three sites differing in physical characteristics, and measured the effect of this manipulation on both male and female success. 3. More pollen was dispersed from the non-manipulated flowers compared with the size-reduced flowers. Standardized selection differentials for flower size through pollen dispersal were significantly positive in all three populations. By contrast, flower size had little effect on female success. Fruit set was always high and selection for this trait through fruiting was not significant. The number of seeds per fruit from bigger flowers was significantly higher than that from smaller flowers, but this difference was attributable exclusively to one population in which selection via seeds per fruit was significant. Flower size reduction had no effect on offspring quality. 4. Our results show that phenotypic selection through male function generally supports that flower size would evolve primarily through selection on male fertility (male function hypothesis). Selection through female function was apparent only in one population, as pollen limitation was absent or weak. We show the relative strength of selection through male and female functions depends on the pollinator context. [ABSTRACT FROM AUTHOR]

The diversity of angiosperm flowers is astounding. The conventional explanation for this diversity is that it represents the great variety of ways in which flowers have adapted to attract an even greater diversity of animal pollinators. Many animal behaviourists are therefore interested in how changes in floral morphology affect pollinator behaviour. The establishment of well-characterised model plant species has greatly furthered our understanding of how floral morphology is generated and varied. Many of these model species are pollinated by animals and attract their pollinators through the production of colour, shape, scent, size and rewards. An understanding of the developmental plasticity of floral morphology, and the constraints upon it, should inform research into animal responses to flowers. The use of genetically characterised model species, and the isogenic and near-isogenic lines available in them, will allow dissection of the different components of floral attraction and reward in natural systems. [ABSTRACT FROM AUTHOR]

In animal-pollinated plants, two factors affecting pollen flow and seed production are changes in floral display and the availability of compatible mates. Changes in floral display may affect the number of pollinator visits and the availability of compatible mates will affect the probability of legitimate pollination and seed production. Distyly is a floral polymorphism where long-styled (pin) and short-styled (thrum) floral morphs occur among different individuals. Distylous plants frequently exhibit self and intra-morph incompatibility. Therefore changes in morph abundance directly affect the arrival of compatible pollen to the stigmas. Floral morph by itself may also affect female reproductive success because floral morphs may display differences in seed production. We explored the effects of floral display, availability of neighboring compatible mates, and floral morph on seed production in the distylous herb ARCYTOPHYLLUM LAVARUM. We found that floral display does not affect the mean number of seeds produced per flower. There is also no effect of the proportion of neighboring legitimate pollen donors on seed production in pin or thrum flowers. However, floral morphs differed in their female reproductive success and the thrum morph produced more seeds. Hand pollination experiments suggest that differences in seed production between morphs are the result of pollen limitation. Future research will elucidate if the higher seed production in thrum flowers is a consequence of higher availability of pollen donors in the population, or higher efficiency of the pin morph as pollen donor. [ABSTRACT FROM AUTHOR]

Interspecific interactions can vary within and among populations and geographical locations, and this variation can influence the nature of the interaction (e.g. mutualistic versus antagonistic) and its evolutionary stability. Globeflowers are exclusively pollinated by flies whose larvae feed only on their seeds. Here we document geographical variability in costs and benefits in globeflowers in sustaining their pollinating flies throughout the range of this arctic-alpine European plant over several years. A total of 1,710 flower heads from 38 populations were analysed for their carpel, egg and seed contents. Individual and population analyses control for the confounding influences of variation in both: (1) population traits, such as fly density and egg distribution among flower heads; and (2) individuals traits, such as carpel and egg numbers per flower head. Despite considerable variation in ecological conditions and pollinator densities across populations, large proportions (range 33–58%) of seeds are released after predation, with a benefit-to-cost ratio of 3, indicating that the mutualism is stable over the whole globeflower geographical range. The stability of the mutualistic interaction relies on density-dependent competition among larvae co-developing in a flower head. This competition is revealed by a sharp decrease in the number of seeds eaten per larva with increasing larval number, and is intensified by non-uniform egg distribution among globeflowers within a population. Carpel number is highly variable across globeflowers (range 10–69), and flies lay more eggs in large flowers. Most plants within a population contribute to the rearing of pollinators, but some pay more than others. Large globeflowers lose more seed to pollinator larvae, but also release more seed than smaller plants. The apparent alignment of interests between fly and plant (positive relationship between numbers of seeds released and destroyed) is shown to hide a conflict of interest found when flower size is controlled for. [ABSTRACT FROM AUTHOR]

We analyze two mathematical models of adaptive investment in rewarding plant traits. In both models, the attractiveness of a particular trait value declines as the mean value in the population increases (asymmetric competition), giving relatively rewarding traits a competitive advantage. Including this competition for pollinator visits in a standard model of hermaphroditic sex allocation shifts additional allocation to pollinator rewards at the expense of allocation to pollen and seeds. In the second model, plants can invest additional resources in pollinator rewards but suffer reduced viability and rising costs due to excess pollen removal and within-plant selfing (geitonogamy). Despite these accumulating costs, increasing the magnitude of asymmetric competition exaggerates the ESS investment in rewards beyond the equilibrium in cases where attractiveness depends only on a plant's absolute reward value. We suggest that the type of frequency dependent selection modeled here is fundamentally equivalent to sexual selection in animal populations (with some unique exceptions). Testing the main assumptions of our models may reveal whether seemingly ‘extravagant’ floral traits are strictly analogous to the exaggerated secondary sexual traits of animals. [ABSTRACT FROM AUTHOR]

We investigated the responses of staminate and pistillate floral components of Prime-Jan and Prime-Jim primocane-fruiting blackberry (Rubus L. subgenus Rubus Watson) to three different growth chamber temperature regimens, 35.0/23.9 °C (HT), 29.4/18.3 °C (MT), and 23.9/12.8 °C (LT). Temperature was negatively related to flower size, and morphologically abnormal floral structures were evident in 41% and 98% of the MT- and HT-grown plants, respectively. Anthers of LT- and MT-grown plants dehisced. The viability of pollen (as deduced through staining) from Prime-Jan grown at LT or MT exceeded 70%, whereas that of Prime-Jim pollen was significantly reduced (<40%) by the MT regimen. In vitro pollen germinability (typically <50%) was negatively influenced by temperature but was unaffected by cultivar. Pollen useful life was diminished under HT conditions; LT-grown pollen held at 23.9 °C retained 63% of its original germinability over a 32-h period, while the germinability of that held at 35.0 °C for 16 hours decreased by 97%. Virtually all flowers cultured under HT conditions were male sterile, exhibiting structural or sporogenous class abnormalities including petaloidy and malformation of tapetal cells or microspores; HT anthers that were present, failed to dehisce. Stigma receptivity, pistil density, and drupelet set were also negatively influenced by increased temperature; values for these parameters of floral competency among control plants were reduced by 51%, 39%, and 76%, respectively, in flowers cultured under HT conditions. In this study, flowering and fruiting parameters, and presumably the yield potential of Prime-Jan and Prime-Jim, were adversely affected by increased temperature. However, their adaptive response to heat stress under field conditions awaits assessment. [ABSTRACT FROM AUTHOR]

Mutualisms are commonly exploited by cheater species that usurp rewards without providing reciprocal benefits. Yet most studies of selection between mutualist partners ignore interactions with third species and consequently overlook the impact of cheaters on evolution in the mutualism. Here, we explicitly investigate how the abundance of nectar-thieving ants (cheaters) influences selection in a pollination mutualism between bumble bees and the alpine skypilot, Polemonium viscosum. As suggested in past work with this species, bumble bees accounted for most of the seed production (78% ± 6% [mean ± SE]) in our high tundra study population and, in the absence of ants, exerted strong selection for large flowers. We tested for indirect effects of ant abundance on seed set through bumble bee pollination services (pollen delivery and pollen export) and a direct effect through flower damage. Ants reduced seed set per flower by 20% via flower damage. As ant density increased within experimental patches, the rate of flower damage rose, but pollen delivery and export did not vary significantly, showing that indirect effects of increased cheater abundance on pollinator service are negligible in this system. To address how ants affect selection for plant participation in the pollination mutualism we tested the impact of ant abundance on selection for bumble bee-mediated pollination. Results show that the impact of ants on fitness (seed set) accruing under bumble bee pollination is density dependent in P. viscosum. Selection for bumble bee pollination declined with increasing ant abundance in experimental patches, as predicted if cheaters constrain fitness returns of mutualist partner services. We also examined how ant abundance influences selection on flower size, a key component of plant investment in bumble bee pollination. We predicted that direct effects of ants would constrain bumble bee selection for large flowers. However, selection on flower size was significantly positive over a wide range of ant abundance (20—80% of plants visited by ants daily). Although high cheater abundance reduces the fitness returns of bumble bee pollination, it does not completely eliminate selection for bumble bee attraction in P. viscosum. [ABSTRACT FROM AUTHOR]

The tolerance of various species of plant to abiotic stress has been enhanced by genetic engineering with certain genes. However, the use of such transgenes is often associated with negative effects on growth and productivity under non-stress conditions. The codA gene from Arthrobacter globiformis is of particular interest with respect to the engineering of desirable productive traits in crop plants. The expression of this gene in tomato plants resulted in significantly enlarged flowers and fruits under non-stress conditions. The enlargement of flowers and fruits was associated with high levels of glycinebetaine that accumulated in reproductive organs, such as flower buds and fruits. The enlargement of flowers was related to an increase in the size and number of cells, and reflected the pleiotropic effect of the codA transgene on the expression of genes involved in the regulation of cell division. [ABSTRACT FROM AUTHOR]

Variation in flower size is an important aspect of a plant’s life history, yet few studies have shown how flower size varies with environmental conditions and to what extent foliar responses to the environment are correlated with flower size. The objectives of this study were to (1) develop a theoretical framework for linking flower size and leaf size to their costs and benefits, as assessed using foliar stable carbon isotope ratio (δ13C) under varying degrees of water limitation, and then (2) examine how variation in flower size within and among species growing along a naturally occurring moisture availability gradient correlates with variation in δ13C and leaf size. Five plant species were examined at three sites in Oregon. Intra- and inter-specific patterns of flower size in relation to moisture availability were the same: the ratios of the area of flower display to total leaf area and of individual flower area to leaf area were greater at sites with more soil moisture compared to those sites with less soil moisture. The increase in flower area per unit increase in leaf area was greater at sites with more soil moisture than at sites where water deficit can occur. Values of δ13C, an index of water-use efficiency, were greater for plants with larger floral size. The patterns we observed generalize across species, irrespective of overall plant morphology or pollination system. These correlations between flower size, moisture availability, and δ13C suggest that water loss from flowers can influence leaf responses to the environment, which in turn may indirectly mediate an effect on flower size. [ABSTRACT FROM AUTHOR]

To determine whether population differentiation in flowering time is consistent with differences in current selection, we quantified phenotypic selection acting through female reproductive success on flowering phenology and floral display in two Scandinavian populations of the outcrossing, perennial herb Arabidopsis lyrata in two years. One population was located in an alpine environment strongly affected by grazing, whereas the other was close to sea level and only moderately affected by herbivory. Multiple regression models indicated directional selection for early end of flowering in one year in the lowland population, and directional selection for early start of flowering in one year in the alpine population. As expected, there was selection for more inflorescences in the lowland population. However, in the alpine population, plants with many inflorescences were selectively grazed and the number of inflorescences produced was negatively related to female fitness in one year and not significantly related to female fitness in the second year. The results are consistent with the hypothesis that genetic differentiation in flowering phenology between the study populations is adaptive, and indicate that interactions with selective grazers may strongly influence selection on floral display in A. lyrata. [ABSTRACT FROM AUTHOR]

Two non-mutually exclusive hypotheses regarding the benefits of andromonoecy (producing perfect and female-sterile flowers on the same plant) are tested using Solanum carolinense. Results indicate that (1) staminate flowers are cheaper to produce than perfect flowers, even after correcting for their relative position in the inflorescence; (2) the resources saved by producing staminate flowers are not re-allocated to other fitness enhancing functions; and (3) the main morphological characteristic of staminate flowers, pistil reduction, does not increase either pollinator visitation or siring success of open-pollinated plants. These results indicate that neither the resource savings hypothesis nor the increased pollen donation hypothesis explains the evolution and maintenance of andromonoecy in S. carolinense. [ABSTRACT FROM AUTHOR]

1. Animal-pollinated plants influence their mating success through characteristics of their individual flowers and the arrangement of flowers into inflorescences. Previous studies of inflorescence function have focused on flower number, so the influences of traits of individual flowers on pollinator attraction and self-pollination between flowers remain unknown. 2. To investigate the effects of flower size and number on pollinator attraction and behaviour on inflorescences, we reduced the perianth size of flowers of Delphinium bicolor Nuttall and Delphinium glaucum S. Watson. 3. Reduction in flower size decreased the number of visits per inflorescence by bumble bees ( Bombus spp.), but increased the number of probes per visit. In contrast, both attraction and probes per visit increased in a decelerating manner with number of open flowers. The average number of probes per flower, which combines the effects of pollinator attraction and behaviour on inflorescences, did not differ significantly between small- and large-flowered plants, or with flower number. 4. The absence of significant variation among plants with different floral and inflorescence characteristics in visits per flower and nectar standing crop per flower indicate that bees achieved an ideal free distribution. 5. Our results suggest that large flowers reduce the incidence of geitonogamous pollination without reducing the frequency of probes per flower. [ABSTRACT FROM AUTHOR]

• Background and Aims Despite considerable investment in elaborate floral displays, Tacca chantrieri populations are predominantly selfing. It is hypothesized that this species might possess considerable spatial or temporal variation in outcrossing rates among populations. To test this hypothesis, genetic variability and genetic differentiation within and among T. chantrieri populations were investigated to find out if they are in agreement with expectations based on a predominantly inbred mating system.• Methods Genetic diversity was quantified using inter-simple sequence repeats (ISSR) in 303 individuals from 13 populations taken from known locations of T. chantrieri in China, and from one population in Thailand.• Key Results Of the 113 primers screened, 24 produced highly reproducible ISSR bands. Using these primers, 160 discernible DNA fragments were generated, of which 145 (90·62 %) were polymorphic. This indicated considerable genetic variation at the species level. However, there were relatively low levels of polymorphism at population levels, with percentages of polymorphic bands (PPB) ranging from 8·75 % to 55 %. A high level of genetic differentiation among populations was detected based on different measures (Nei''s genetic diversity analysis: GST = 0·5835; AMOVA analysis: FST = 0·6989). Furthermore, based on levels of genetic differentiation, the 14 populations clustered into two distinct groups separated by the Tanaka Line.• Conclusions High levels of differentiation among populations and low levels of diversity within populations at large spatial scales are consistent with earlier small-scale studies of mating patterns detected by allozymes which showed that T. chantrieri populations are predominantly selfing. However, it appears that T. chantrieri has a mixed-mating system in which self-fertilization predominates, but there is occasional outcrossing. Significant genetic differences between the two distinct regions might be attributed to vicariance along the Tanaka Line. Finally, possible mechanisms of geographic patterns based on genetic differentiation of T. chantrieri are discussed. [ABSTRACT FROM AUTHOR]

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

Abstract: Many cool season garden crops, including Viola × wittrockiana Gams. (pansy), exhibit reduced flowering outdoors during the warm summer months. Twelve pansy cultivars varying in summer garden performance were grown under either 20±1.5 or 30±1°C (air temperature) to determine growth and flowering responses to prolonged high-temperature exposure and to identify selection criteria to screen pansies for flowering heat tolerance. Increasing temperature from 20 to 30°C increased leaf number below the first flower on ‘Crystal Bowl Primrose’ and ‘Skyline White’ only. Flower bud number reduction at 30°C versus 20°C varied from 20% for ‘Crystal Bowl Purple’ to 77% for ‘Majestic Giants Red and Yellow’. Flower diameter reduction at 30°C versus 20°C ranged from 14% for ‘Skyline Beaconsfield’ to 44% for ‘Super Majestic Giants Ocean’. The percentage reduction in total color (flower number×estimated flower area) ranged from 60% for ‘Crystal Bowl Primrose’ to 88% for ‘Majestic Giants Rose Shades’. Based on a weighted base selection index, ‘Super Majestic Giants Canary’ and ‘Delta Yellow’ were identified as the most heat-tolerant cultivars, while ‘Super Majestic Giants Ocean’ and ‘Majestic Giants Rose Shades’ were identified as the most heat-sensitive. In a second experiment, root and shoot dry mass were determined after 10, 20, or 30d when grown at 20 or 30°C. Relative growth rate and root:shoot ratio were also calculated. After 30d, ‘Crystal Bowl Primrose’, ‘Crystal Bowl Sky Blue’ and ‘Skyline White’ relative growth rates were lower at 30°C versus 20°C. Root:shoot ratio on day 30 was lower at 30°C compared to 20°C for six cultivars, but similar across temperature for five cultivars and higher for ‘Crystal Bowl Primrose’. Flower bud number at first flower was positively correlated with branch number, shoot dry mass at flowering, but not correlated with root dry mass at flowering, and negatively correlated with flower diameter and root:shoot ratio (either at flowering, or after 10, 20 or 30d at 30°C), suggesting that these traits may be useful when screening pansies for flowering heat tolerance. [Copyright &y& Elsevier]

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Gynodioecious populations (i.e. populations with female and hermaphrodite individuals) often contain a third phenotype with an intermediate sex expression. In Silene italica, this phenotype is characterized by a mixture of pistillate and perfect flowers and is thus gynomonoecious. To characterize sexual functions of these gynomonoecious individuals and their potential influence in the maintenance of gynodioecy, reproductive characters of the three sexual phenotypes were compared over 2 years in several families of S. italica produced by crossing. We found that gynomonoecious individuals were intermediate for flower production and female fertility characters, although they did not always significantly differ from female individuals. Perfect flowers of gynomonoecious and hermaphrodite plants were similar in size and pollen production. Gynomonoecious individuals were thus intermediate in female and in male functions. Family effects were found for most of the characters. The female advantage (i.e. the fertility of females compared to the female fertility of pollen producing plants) was not dramatically different when gynomonoecious plants were taken into account. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 87, 583–591. [ABSTRACT FROM AUTHOR]

We looked at whether flower visitation rates, pollen limitation on seed production, and phenotypic selection on flower size through female function varied across a sharp gradient in light intensity (open meadow vs. forest) within a population of the perennial plant Campanula persicifolia L. (Campanulaceae). Flower visitation rates of putative pollinators were similar in both habitats. Seed number per fruit was strongly pollen limited, with no difference in the magnitude of pollen limitation between the two habitats. This strong pollen limitation set the basis for significant phenotypic selection, through female function, on a trait that probably is important for pollinator attraction: flower size. This was revealed by path analysis and structural equation modelling. The lack of difference in pollen limitation on seed production in the two habitats may be explained by the similarity in flower visitation rates in the two habitats. Moreover, the similarity in pollen limitation in the two habitats probably resulted in a similar magnitude and direction of selection on flower size through female function. Our results suggest that pollen limitation and selection through female function may vary little across space within a popualtion despite large variation in the environmental conditions experienced by plants. [ABSTRACT FROM AUTHOR]

Plants interact simultaneously with a diversity of visitors, including herbivores and pollinators. Correlations among traits associated with herbivory and pollination may constrain the degree to which plants can evolve in response to any one interactor. Using the distylous plant, Gelsemium sempervirens, we tested the hypothesis that traits typically associated with pollination (distyly) and herbivore resistance (secondary compounds) were phenotypically correlated and examined how these traits influenced plant interactions with floral visitors. The flowers of G. sempervirens are visited by pollinators and a nectar robber, and the leaves and flowers express gelsemine, an alkaloid that is deterrent and sometimes toxic to visitors. Using an observational approach across five populations, we found the thrum floral morph (short-styled) expressed more leaf gelsemine than the pin morph (long-styled). Leaf gelsemine concentrations were positively correlated with flower gelsemine; however, there were no correlations between gelsemine and other floral morphological traits. Trait expression influenced pollination more so than robbing. Thrums received two times less pollen than pins. Moreover, across both morphs, pollen receipt was lower in plants that expressed higher levels of leaf gelsemine in two sites. These results imply that traits associated with pollination and herbivore resistance may not he independent. [ABSTRACT FROM AUTHOR]

Plants need not participate passively in their own mating, despite their immobility and reliance on pollen vectors. Instead, plants may respond to their recent pollination experience by adjusting the number of flowers that they display simultaneously. Such responsiveness could arise from the dependence of floral display size on the longevity of individual flowers, which varies with pollination rate in many plant species. By hand-pollinating some inflorescences, but not others, we demonstrate plasticity in display size of the orchid Satyrium longicauda. Pollination induced flower wilting, but did not affect the opening of new flowers, so that within a few days pollinated inflorescences displayed fewer flowers than unpollinated inflorescences. During subsequent exposure to intensive natural pollination, pollen removal and receipt increased proportionally with increasing display size, whereas pollen-removal failure and self-pollination accelerated. Such benefitcost relations allow plants that adjust display size in response to the prevailing pollination rate to increase their attractiveness when pollinators are rare (large displays), or to limit mating costs when pollinators are abundant (small displays). Seen from this perspective, pollination-induced flower wilting serves the entire plant by allowing it to display the number of flowers that is appropriate for the current pollination environment. [ABSTRACT FROM AUTHOR]

• Background and Aims Flowers are relatively invariant organs within species, but quantitative variation often exists among conspecifics. These variations represent the raw material that natural selection can magnify, eventually resulting in morphological divergence and diversification. This paper investigates floral variability in Rosmarinus officinalis, a Mediterranean shrub. [ABSTRACT FROM PUBLISHER]

Sexual dimorphism is one of the most widespread and recognizable patterns of phenotypic variation in the biotic world. Sexual dimorphism in floral display is striking in the dioecious plant Silene latifolia, with males making many, small flowers compared to females. We investigated this dimorphism via artificial selection on two populations to determine whether genetic variation exists within populations for flower size and the extent of the between-sex correlation, whether a flower size and number trade-off exists within each sex, and whether pollen and ovule production vary with flower size. We selected for decreased flower size (calyx width) in females and increased flower size in males and measured the response to selection in size and correlated responses in flower dry mass, flower number, and pollen or ovule number per flower. Four bouts of selection in each of two selection programs were performed, for a total of three selection lines to decrease size, three to increase it, and two control lines. Flower size always significantly responded to selection and we always found a significant correlated response in the sex not under selection. Selection decreased but did not eliminate the sexual dimorphism in flower dry mass and number. A negative relationship between flower size and number within each sex was revealed. Whereas ovule number showed a significant correlated response to selection on flower size, pollen number did not. Our results indicate that although substantial additive genetic variation for flower size exists, the high between-sex genetic correlation would likely constrain flower size from becoming more sexually dimorphic. Furthermore, floral display within each sex is constrained by a flower size and number trade-off. Given this trade-off and lack of variation in pollen production with flower size, we suggest that sexual dimorphism evolved via sexual selection to increase flower number in males but not females. [ABSTRACT FROM AUTHOR]

Two separate studies were initiated in January 1998 on 'Tifblue' rabbiteye (Vaccinium ashei Reade) and 'Magnolia' southern highbush (V. corymbosum L.) blueberry plants that had received > 500 chilling hours (7°C). In each study, the terminal 2 flower buds from both excised stems and paired stems on plants were forced in a greenhouse (17-23°C and natural daylength) and rated for floral bud development. In 'Magnolia,' leaf removal from excised stems and intact stems resulted in an increased rate of floral bud development. After 7 days of forcing, excised stems with leaves removed showed no significant differences in floral bud development from intact plants with unabscised leaves. After 40 days of forcing, flower buds from all treatments had reached stage 3, an easily recognizable stage of development. 'Tifblue' stems with 5 mm basal sections removed weekly to retard vascular blockage did not differ in floral bud development from stems without basal pruning. Excised stems did not differ from matching intact stems for the first 3.5 weeks of forcing. At this time 'Tifblue' flower buds had reached 3.7 on the floral development scale. After>4.5 weeks of forcing in the greenhouse floral bud development ratings on excised stems were significantly lower than those on intact stems. These studies indicate that with sufficient chilling, floral bud development in excised stems without leaves approximate that of intact plants for a period of 3.5 weeks and reach at least stage 3, an easily identifiable floral stage. [ABSTRACT FROM PUBLISHER]

Assumed trade-offs between male and female functions in hermaphroditic flowers have been difficult to demonstrate. Collinsia parviflora (Scrophulariaceae) is a winter annual that exhibits significant among-population variation in corolla size in British Columbia, Canada. We asked whether reduction in secondary male allocation (i.e., the attractive corolla), a preliminary indicator of mating system, was matched by a reduction in primary male allocation (i.e., pollen production), and whether there were allocation trade-offs between male and female function both within and among six study populations. Larger-flowered populations allocated more to male function (androecium and corolla biomass), and because populations did not vary in female biomass allocation (gynoecium and calyx), population differences were not due to simple allometric scaling. Populations also differed in per-flower gamete production (pollen and ovules). We found male-female trade-offs within populations between androecium and gynoecium mass and between corolla and calyx mass. Among populations, there was a marginal trade-off between pollen and ovule production and a significant within-male trade-off between pollen grain size and number. Trade-offs between the sexes were primarily apparent when we controlled for flower size in the analysis. Variation among populations in sex allocation may reflect different optima related to the mating system. [ABSTRACT FROM AUTHOR]

Although pollinator-mediated natural selection has been measured on many floral traits and in many species, the extent to which selection is constrained from producing optimal floral phenotypes is less frequently studied. In particular, negative correlations between flower size and flower number are hypothesized to be a major constraint on the evolution of floral displays, yet few empirical studies have documented such a trade-off. To determine the potential for genetic constraints on the adaptive evolution of floral displays, I estimated the quantitative genetic basis of floral trait variation in two populations of Lobelia siphilitica. Restricted maximum likelihood (REML) analyses of greenhouse-grown half-sib families were used to estimate genetic variances and covariances for flower number and six measures of flower size. There was significant genetic variation for all seven floral traits in both populations. Flower number was negatively genetically correlated with four measures of flower size in one population and three measures in the other. When the genetic variance-covariance matrices were combined with field estimates of phenotypic selection gradients, the predicted multivariate evolutionary response was less than or opposite in sign to the selection gradient for flower number and rive of six measures of flower size, suggesting genetic constraints on the evolution of these traits. More generally, my results indicate that the adaptive evolution of floral displays can be constrained by tradeoffs between flower size and number, as has been assumed by many theoretical models of floral evolution. [ABSTRACT FROM AUTHOR]

A model was developed describing the influence of irradiance and temperature in the greenhouse on the size of Chrysanthemum flowers. In the model, flower diameter increment was related to a development index ranging from zero (start of SD) to unity (anthesis). The growth of the visible flower was divided into two different phases. In the first phase a linear function described the growth and development of the early visible flower bud, while the second phase, representing the opening process of the flower was best described with a monomolecular growth function. The effect of the climate on the two growth phases was modelled using empirical climate functions. Data, collected from a light and a temperature experiment, showed that low light integrals and temperatures above 20 °C had a retarding effect on flower growth. When the model was fitted to the observed data from the light experiment the R2-values varied from 0.999 to 0.965. Even the simulated diameter values matched the observed values from the temperature experiment well (R2-values from 0.998 to 0.966). When validated on independently collected data from two trials, the model could simulate the variations in the data with R2-values of 0.993 and 0.997. [Copyright &y& Elsevier]

Although many diclinous plants have a flower size dimorphism, the causes and ecological correlates of dimorphism in flower size remain poorly understood. In this paper we quantify the frequency and distribution of flower size dimorphism in the native flora of La Réunion Island. The frequency of flower size dimorphism is lower than in a previous global study suggesting that the dimorphism may limit colonisation success. Flower size dimorphism is significantly more frequent in endemic taxa compared to indigenous taxa, a trend that is particularly evident at the species level, due to the large number of species with male flowers bigger than female flowers and is more common in species-rich lineages. These results suggest that flower size dimorphism may be associated with species divergence. The direction of dimorphism varies between dioecious and monoecious species, suggesting that the mechanisms acting on flower size variation differ in species with these different sexual systems. [ABSTRACT FROM AUTHOR]

A common gender dimorphism in angiosperms is gynodioecy, in which hermaphrodites and females co-occur. Females are at an inherent disadvantage because they can transmit their genes only through ovule production. One mechanism by which females can compensate for the loss of male function is by producing more seeds than hermaphrodites. As such, females should: (1) increase resource uptake to support higher seed production; and (2) allocate resources saved by the loss of male function to seed production. To test this hypothesis, we measured physiological and floral traits of gynodioecious Lobelia siphilitica, controlling for both environmental and genetic variation through a comparison of greenhouse-grown siblings. Pre-reproductive females had 14% higher area-based (Z=2.14; P=0.04) and 32% higher mass-based (Z=1.96; P=0.05) photosynthetic rate than hermaphrodites, suggesting that they have increased carbon acquisition by altering photosynthetic physiology. Female L. siphilitica produced flowers with 4–8% smaller corollas than hermaphrodites (all P<0.05), suggesting that females allocate resources away from floral structures used for pollinator attraction. The genetic correlation between genders for four floral and four physiological traits was significantly less than one but greater than zero, indicating that the evolution of gender dimorphism in response to sex-differential selection will be constrained. The allocation of resources saved by the loss of male function has been viewed as the most important mechanism allowing females of gynodioecious species to support higher seed production. Our data suggest that increased resource acquisition by females at pre-reproductive stages can also contribute to the maintenance of gender dimorphism in gynodioecious species. [ABSTRACT FROM AUTHOR]

There has been very little empirical study of quantitative genetic variation in flower size in sexually dimorphic plant species, despite the frequent occurrence of flower size differences between sexual phenotypes. In this study we quantify the nature of quantitative flower size variation in females and hermaphrodites of gynodioecious Thymus vulgaris . In a field study, females had significantly smaller flowers than hermaphrodites, and the degree of flower size dimorphism varied significantly among populations. To quantify the genetic basis of flower size variation we sampled maternal progeny from 10 F 0 females in three populations (across the range of variation in flower size in the field), performed controlled crosses on F 1 offspring in the glasshouse and grew F 2 progeny to flowering in uniform field conditions. A significant population * sex interaction was again observed, hence the degree of sexual dimorphism shows genetic variation among populations. A significant family * sex interaction was also observed, indicating that the degree of sexual dimorphism shows genetic variation among families. Females showed significantly greater variation among populations and among families than hermaphrodites. Female flower size varied significantly depending on the degree of stamen abortion, with morphologically intermediate females having flowers more similar to hermaphrodites than to other females. The frequency of female types that differ in the degree of stamen abortion varied among populations and families and mean family female flower size increased as the proportion of intermediate female types increased across families. Variation in the degree of flower size dimorphism thus appears to be a result of variation in the degree of stamen abortion in females, the potential causes of which are discussed. [ABSTRACT FROM AUTHOR]

The evolution of a showy floral display as an advertisement to pollinators could simultaneously advertise the availability of resources to pre-dispersal seed-predators. The hypotheses tested here are that the incidence of seed predation by bud-infesting insect larvae in capitula of Asteraceae is positively related to (1) capitulum size among species, (2) capitulum size within species, (3) capitulum lifespan, and (4) the degree of flowering asynchrony on individual plants. Three populations of each of 20 common herbaceous species of Asteraceae from disturbed ground and grassland habitats were monitored for the presence of pre-dispersal, seed-eating insect larvae. Mean capitulum size (receptacle width) of each species was measured. In a sub-set of eight species, individual capitula were tagged to determine their flowering phenology and lifespan (from anthesis to seed shedding). From these data an index of flowering synchrony on individual plants was derived. Among species, the incidence of larval infestation increased with capitulum size. Small-flowered species such as Achillea millefolium were largely free of bud-infesting larvae, whilst large-flowered species such as Arctium minus were heavily infested. In three cases investigated in greater detail, bud infestation was found to increase with capitulum size within species, suggesting a potential for natural selection to favour smaller capitula. No relationship was found between infestation levels and either capitulum lifespan or degree of flowering synchrony, and there was no evidence that the relationship between capitulum size and infestation was confounded by correlations with these other features. The results support hypotheses 1 and 2, but not 3 and 4. It is suggested that the characteristic capitulum size of each species may represent a trade-off between the opposing selection pressures of pollinators and pre-dispersal seed predators. [ABSTRACT FROM AUTHOR]

We studied the interrelation between nectary structure (13 parameters), nectar characteristics (yield, chemical composition), and flower size of 11 Labiatae species in a Mediterranean shrub community near Athens, Greece. We also explored whether the above attributes are affected by the Mediterranean summer drought constraints. Our findings show that among all nectary parameters studied, nectary size and stomatal opening are the most important in (positively) shaping nectar secretion, nectary size being the most meaningful. Nectary structure is correlated to quantity of the nectar secreted, not its quality. Wide flowers bear wide nectaries with large stomatal openings, whereas deep flowers are not related to any nectary size. Corolla size (both length and width) and nectary stomatal opening decrease with flowering time. This applies also to nectary size, nectar volume and sugar content of the perennials (9 species). All above cases of time dependence show that there is a constraint effect of Mediterranean climate on floral and nectary structure, reflected also as a decrease in nectar secretion. Nectary structure in Labiatae is largely shaped by both phylogenetic and climate constraints. On the other hand, although nectar is largely influenced by nectary structure, it is to a large extent ecologically biased, implying that, in addition to phylogeny, there are many other ecological parameters interfering in its secretion such as time within the season, life history, and light requirements. [ABSTRACT FROM AUTHOR]

The orchid market is a dynamic horticultural business in which novelty and beauty command high prices. The two main interests are the development of flowers, from the miniature to the large and showy, and their fragrance. Overall organ size might be modified by doubling the chromosome number, which can be accomplished by careful study of meiotic chromosome disjunction in hybrids or species. Meiosis is the process in which diploid (2n) pollen mother cells recombine their DNA sequences and then undergo two rounds of division to give rise to four haploid (n) cells. Thus, by interfering in chromosome segregation, one can induce the development of diploid recombinant cells, called unreduced gametes. These unreduced gametes may be used for breeding polyploid progenies with enhanced fertility and large flower size. This review provides an overview of developments in orchid polyploidy breeding placed in the large context of meiotic chromosome segregation in the model plants Arabidopsis thaliana and Brassica napus to facilitate molecular translational research and horticultural innovation.

Plant pathogens evade basal defense systems and attack different organs and tissues of plants. Genetic engineering of plants with genes that confer resistance against pathogens is very effective in pathogen control. Conventional breeding for disease resistance in ornamental crops is difficult and lagging relative to that in non-ornamental crops due to an inadequate number of disease-resistant genes. Therefore, genetic engineering of these plants with defense-conferring genes is a practical approach. We used rice BSR2 encoding CYP78A15 for developing transgenic Torenia fournieri Lind. lines. The overexpression of BSR2 conferred resistance against two devastating fungal pathogens, Rhizoctonia solani and Botrytis cinerea . In addition, BSR2 overexpression resulted in enlarged flowers with enlarged floral organs. Histological observation of the petal cells suggested that the enlargement in the floral organs could be due to the elongation and expansion of the cells. Therefore, the overexpression of BSR2 confers broad-spectrum disease resistance and induces the production of enlarged flowers simultaneously. Therefore, this could be an effective strategy for developing ornamental crops that are disease-resistant and economically more valuable.

Geographic differences in floral traits may reflect geographic differences in effective pollinator assemblages. Independent local adaptation to pollinator assemblages in multiple regions would be expected to cause parallel floral trait evolution, although sufficient evidence for this is still lacking. Knowing the intraspecific evolutionary history of floral traits will reveal events that occur in the early stages of trait diversification. In this study, we investigated the relationship between flower spur length and pollinator size in 16 populations of Aquilegia buergeriana var. buergeriana distributed in four mountain regions in the Japanese Alps. We also examined the genetic relationship between yellow- and red-flowered individuals, to see if color differences caused genetic differentiation by pollinator isolation. Genetic relationships among 16 populations were analyzed based on genome-wide single-nucleotide polymorphisms. Even among populations within the same mountain region, pollinator size varied widely, and the average spur length of A. buergeriana var. buergeriana in each population was strongly related to the average visitor size of that population. Genetic relatedness between populations was not related to the similarity of spur length between populations; rather, it was related to the geographic proximity of populations in each mountain region. Our results indicate that spur length in each population evolved independently of the population genetic structure but in parallel in response to local flower visitor size in different mountain regions. Further, yellow- and red-flowered individuals of A. buergeriana var. buergeriana were not genetically differentiated. Unlike other Aquilegia species in Europe and America visited by hummingbirds and hawkmoths, the Japanese Aquilegia species is consistently visited by bumblebees. As a result, genetic isolation by flower color may not have occurred., Competing Interests: There are no conflicts of interest., (© 2022 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

We estimated the genetic basis for variation in floral traits and the phenotypic and genetic correlations between these traits using clonal propagation in Echium vulgare (Boraginaceae). For all floral traits measured (nectar volume, nectar sugar concentration, sugar production, corolla size, style length, stamen length) we found significant clonal repeatabilities in a greenhouse experiment. Variation among clonal families in nectar volume was surprisingly high (seven-fold difference between the highest and lowest family mean) and not correlated with flower size. Variation among clonal families in pollen production was even higher (28-fold) and correlated to flower size traits. Corolla length, style length and stamen length were all highly correlated. Flower size is not a good predictor for nectar production rate, while it is for the number of pollen grains produced. [ABSTRACT FROM AUTHOR]

Water loss through inflorescences may place extreme demands on plant water status in arid environments. Here we examine how corolla size, a trait known to influence pollination success, affects the water cost of flowering in the alpine skypilot, Polemonium viscosum. In a potometry experiment, water uptake rates of inflorescences were monitored during bud expansion and anthesis. Corolla volume of fully expanded flowers predicted water uptake during bud expansion ( R2=0.61, P=0.0375) and corolla surface area predicted water uptake during anthesis ( R2=0.59, P=0.044). To probe mechanisms underlying the relationship between corolla size and water uptake, cell dimensions and densities were measured in several regions of fully expanded corollas. Corolla length was positively correlated with cell length in the middle of the corolla tube and cell diameter in the corolla lobe (Pearson's r from 0.26–0.33, n=86, P ≤ 0.05). Cell density was negatively correlated with cell dimensions in the upper corolla tube and lobe (Pearson's r from –0.39 to –0.42, P ≤ 0.0015). These findings suggest that more water may be required to maintain turgor in large corollas in part because their tissues have lower cell wall densities. The carbon cost of water use by flowers was assessed in krummholz and tundra habitats for P. viscosum flowering, respectively, during dry and wet portions of the growing season. For plants in full flower, average leaf water potentials were significantly more negative ( P=0.0079) at mid-day in the krummholz (June) than in the tundra (July), but were similar before dawn ( P=0.631). Photosynthetic rate at the time of flowering declined significantly with increasing corolla size in the krummholz ( P=0.0376), but was unrelated to corolla size on the tundra ( P>0.72). Plants losing water through large corollas may close leaf stomata to maintain turgor. If photosynthesis limits growth in this perennial species, then the water cost of producing large flowers should exacerbate the cost of reproduction under dry conditions. Such factors could select for flowers with smaller corollas in the krummholz, countering pollinator-mediated selection and helping maintain genetic variation in corolla size components of P. viscosum. [ABSTRACT FROM AUTHOR]