Expression Of Floral Traits Research Articles

Drought and temperature stresses impact pollen production and autonomous selfing in a California wildflower, Collinsia heterophylla

AbstractOngoing climatic changes have altered growing conditions for plants by limiting water availability and inducing unprecedented temperature increases, eliciting plant functional responses that compromise floral trait expression, reduce pollen production and promote early self‐pollination. In a controlled greenhouse study, plants of Collinsia heterophylla, an annual mixed‐mated hermaphrodite, were grown under temperature and water stresses. Floral trait responses and related effects on plant reproductive success were recorded. Plants grown under temperature and water stresses were shorter and had fewer leaves at flowering than control plants. Temperature‐stressed plants flowered earlier and had smaller flowers that produced fewer per capita pollen grains than control and water‐stressed plants. While lifetime flower production in plants experiencing temperature stress alone or water stress alone was similar to control plants, those receiving combined temperature and water stresses had significantly lower lifetime flower production. Abiotic stress did not affect investment in female traits such as ovule number per flower, but impacted male traits seen as fewer pollen grains per flower. In plants experiencing abiotic stress, an increase in autonomous self‐pollination was facilitated through a compromise in herkogamy, a sexual interference mechanism that promotes cross‐pollination. Spatial separation between stigma and anther, a measure of herkogamy was zero or negative in stressed plants. Thus, abiotic stress responses in plants did not compromise plant reproductive success as autonomous selfing was augmented. In nature, such responses will likely reduce available nutrition for pollinators through decreased flower and pollen production.

Responses of floral traits to water limitation across soil types and along climatic gradients

Understanding how environmental changes affect plant-pollinator interactions is of increasing importance given that new environmental extremes continue to be reached. Drought may be a particularly destabilizing force for plant-pollinator interactions because moisture availability can affect floral trait expression. Past work has found variability in how some floral traits respond to moisture limitation, motivating work on the potential drivers of the variability in responses. Here we asked how flower size and nectar provisioning to pollinators change with moisture limitation, and if seed provenance mediates trait responses. Specifically, we conducted a greenhouse common garden using seeds of the serpentine-tolerant annual Antirrhinum vexillocalyculatum collected from pairs of populations on and off drought-prone serpentine soil and along moisture and temperature gradients. We experimentally quantified responses to two moisture treatments, and we found no evidence that seeds from different soil types vary in drought responses. However, plants from seeds from different temperature regimes varied in the way that water limitation affected nectar volume, and those from different precipitation regimes varied in the way that water limitation affected flower size. Water limitation reduced floral display and reward for plants from some populations but increased trait means in other populations. This suggests that drought will reduce floral resources for pollinators overall, but plants from some environments may continue to invest in pollination mutualisms under moisture limitation.

Floral traits and pollination biology ofEpimedium chlorandrumStearn (Berberidaceae)

AbstractThe evolution and expression of floral traits are responsive to selection pressure from biotic and abiotic factors. Although floral traits significantly vary among environments, the flower remains unchanged. We aimed to understand the adaptation of Epimedium chlorandrum of floral traits to a frequently nocturnally rainy and wet environments and the roles of floral traits in pollination and reproduction. We observed flowering phenology, measured floral characteristics including the number of pollen grains and ovules per flower, measured pollen viability and stigma receptivity, tested the volume and sugar concentration of nectar and conducted flower-visit observations in this species. Different pollination treatments were performed to characterize the breeding system. The inner sepals and highly curved longer spurs of E. chlorandrum jointly formed an umbrella that shielded the anthers and stigma from rain wash and prevented nectar dilution. Epimedium chlorandrum was visited by six species, while Bombus trifasciatus was the only effective pollinator and fed on the nectar. One flower secreted approximately 17.06 µL of nectar with a 29.19 g/100 mL sugar concentration, and the pollination efficiency of B. trifasciatus was positively associated with the nectar sugar concentration. The self- and open-pollination treatments resulted in fewer fertile seeds than the cross-pollination treatment. In contrast, the autonomous self-pollination treatment failed to yield fertile seeds. In summary, pollen limitation caused by harsh weather and pollinator shortage occurred during the pollination process of E. chlorandrum, which was partially alleviated by self-compatibility.

Reduced seed set under water deficit is driven mainly by reduced flower numbers and not by changes in flower visitations and pollination

Abstract Water deficit can alter floral traits with cascading effects on flower‐visitor interactions and plant fitness. Water stress induction can diminish productivity, directly resulting in lower flower production and consequently seed set. Changes in floral traits, such as floral scent or reward amount, may in turn alter pollinator visitations and behaviour and consequently can reduce pollination services resulting in lower reproduction output. However, the relative contribution of this indirect in comparison to the direct effects to changes in seed set are not fully understood. We manipulated water availability using rain‐out shelters in a field experiment and measured effects on floral scent bouquet, morphology, phenology, flower‐visitor interactions, pollination and seed set. Plant individuals of Sinapis arvensis (Brassicaceae) were randomly assigned to one of three treatments: mean precipitation (=control), reduced mean precipitation or drought period treatment. Our results show that decreasing water availability lowers the number of flowers and seed set. This indicates a direct link between water stress and seed set, as seed mass increases with increasing flower number. The indirect link of water stress via floral traits, pollinator visits and pollination has weaker effects on seed set. However, floral traits remain relatively stable under decreased water availability, whereas plant growth and flower abundance decrease, potentially in order to allow investment in more resources in fewer flowers to maintain pollination success. Thus, plants are able to compensate for water stress and can maintain floral trait expression, such as a stable scent emission and bouquet, to retain pollinator attraction. These findings indicate that the direct link from water stress to seed set has a stronger impact on plants reproductive success than the indirect link through altered floral trait expression and pollinator visits in a generalist plant species. Read the free Plain Language Summary for this article on the Journal blog.

Soil water and nutrient availability interactively modify pollinator-mediated directional and correlational selection on floral display.

The individual and combined effects of abiotic factors on pollinator-mediated selection on floral traits are not well documented. To examine potential interactive effects of water and nutrient availability on pollinator-mediated selection on three floral display traits of Primula tibetica, we manipulated pollination and nutrient availability in a factorial experiment, conducted at two common garden sites with different soil water content (natural vs addition). We found that both water and nutrient availability affected floral trait expression in P. tibetica and that hand pollination increased seed production most when both nutrient content and water content were high, indicating joint pollen and resource limitation. We documented selection on all floral traits, and pollinators contributed to directional and correlational selection on plant height and number of flowers. Soil water and nutrient availability interactively influenced the strength of both pollinator-mediated directional and correlational selection, with significant selection observed when nutrient or water availability was high, but not when none or both were added. The results suggest that resource limitation constrains the response of P. tibetica to among-individual variation in pollen receipt, that addition of nutrients or water leads to pollinator-mediated selection and that effects of the two abiotic factors are nonadditive.

Flower traits associated with the visitation patterns of bees.

Plant-pollinator interactions are partially driven by the expression of plant traits that signal and attract bees to the nutritional resources within flowers. Although multiple physical and chemical floral traits are known to influence the visitation patterns of bees, how distinct bee groups vary in their responses to floral traits has yet to be elucidated. In this study, we used a common garden experiment to test for morphological floral traits associated with pollen quantity at the plant species level, and examined how the visitation patterns of taxonomically and functionally distinct bee groups are related to flower trait characteristics of 39 wildflower species. We also determined how floral traits influence the structure of wild bee communities visiting plants and whether this varies among geographic localities. Our results suggest that floral area is the primary morphological floral trait related to bee visitation of several distinct bee groups, but that wild bee families and functionally distinct bee groups have unique responses to floral trait expression. The composition of the wild bee communities visiting different plants was most strongly associated with variability in floral area, flower height, and the quantity of pollen retained in flowers. Our results inform wildflower habitat management for bees by demonstrating that the visitation patterns of distinct bee taxa can be predicted by floral traits, and highlight that variability in these traits should be considered when selecting plants to support pollinators.

Phenotypic expression of floral traits in hybrid zones provides insights into their genetic architecture.

Information on the genetic architecture of phenotypic traits is helpful for constructing and testing models of the ecoevolutionary dynamics of natural populations. For plant groups with long life cycles there is a lack of line cross experiments that can unravel the genetic architecture of loci underlying quantitative traits. To fill this gap, we propose the use of variation for phenotypic traits expressed in natural hybrid zones as an alternative approach. We used data from orchid hybrid zones and compared expected and observed patterns of phenotypic trait expression in different early-generation hybrid classes identified by molecular genetic markers. We found evidence of additivity, dominance, and epistatic interactions for different phenotypic traits. We discuss the potential of this approach along with its limitations and suggest that it may represent a realistic way to gain an initial insight into the heritability and genomic architecture of traits in organismal groups with complex life history, such as orchids and many others.

A comparison of coffee floral traits under two different agricultural practices

Floral traits and rewards are important in mediating interactions between plants and pollinators. Agricultural management practices can affect abiotic factors known to influence floral traits; however, our understanding of the links between agricultural practices and floral trait expression is still poorly understood. Variation in floral morphological, nectar, and pollen traits of two important agricultural species, Coffea arabica and C. canephora, was assessed under different agricultural practices (sun and shade). Corolla diameter and corolla tube length were larger and pollen total nitrogen content greater in shade plantations of C. canephora than sun plantations. Corolla tube length and anther filament length were larger in shade plantations of C. arabica. No effect of agricultural practice was found on nectar volume, sugar or caffeine concentrations, or pollen production. Pollen total nitrogen content was lower in sun than shade plantations of C. canephora, but no difference was found between sun and shade for C. arabica. This study provides baseline data on the influence of agronomic practices on C. arabica and C. canephora floral traits and also helps fill a gap in knowledge about the effects of shade trees on floral traits, which can be pertinent to other agroforestry systems.

Evolutionary transition between bee pollination and hummingbird pollination in Salvia: Comparing means, variances and covariances of corolla traits.

Covariation among traits can modify the evolutionary trajectory of complex structures. This process is thought to operate at a microevolutionary scale, but its long-term effects remain controversial because trait covariation can itself evolve. Flower morphology, and particularly floral trait (co)variation, has been envisioned as the product of pollinator-mediated selection. Available evidence suggests that major changes in pollinator assemblages may affect the joint expression of floral traits and their phenotypic integration. We expect species within a monophyletic lineage sharing the same pollinator type will show not only similarity in trait means but also similar phenotypic variance-covariance structures. Here, we tested this expectation using eighteen Salvia species pollinated either by bees or by hummingbirds. Our findings indicated a nonsignificant multivariate phylogenetic signal and a decoupling between means and variance-covariance phenotypic matrices of floral traits during the evolution to hummingbird pollination. Mean trait value analyses revealed significant differences between bee- and hummingbird-pollinated Salvia species although fewer differences were detected in the covariance structure between groups. Variance-covariance matrices were much more similar among bee- than hummingbird-pollinated species. This pattern is consistent with the expectation that, unlike hummingbirds, bees physically manipulate the flower, presumably exerting stronger selection pressures favouring morphological convergence among species. Overall, we conclude that the evolution of hummingbird pollination proceeded through different independent transitions. Thus, although the evolution of hummingbird pollination led to a new phenotypic optimum, the process involved the diversification of the covariance structure.

Oilseed rape (Brassica napus) as a resource for farmland insect pollinators: quantifying floral traits in conventional varieties and breeding systems.

Oilseed rape (OSR; Brassica napus L.) is a major crop in temperate regions and provides an important source of nutrition to many of the yield‐enhancing insect flower visitors that consume floral nectar. The manipulation of mechanisms that control various crop plant traits for the benefit of pollinators has been suggested in the bid to increase food security, but little is known about inherent floral trait expression in contemporary OSR varieties or the breeding systems used in OSR breeding programmes. We studied a range of floral traits in glasshouse‐grown, certified conventional varieties of winter OSR to test for variation among and within breeding systems. We measured 24‐h nectar secretion rate, amount, concentration and ratio of nectar sugars per flower, and sizes and number of flowers produced per plant from 24 varieties of OSR representing open‐pollinated (OP), genic male sterility (GMS) hybrid and cytoplasmic male sterility (CMS) hybrid breeding systems. Sugar concentration was consistent among and within the breeding systems; however, GMS hybrids produced more nectar and more sugar per flower than CMS hybrid or OP varieties. With the exception of ratio of fructose/glucose in OP varieties, we found that nectar traits were consistent within all the breeding systems. When scaled, GMS hybrids produced 1.73 times more nectar resource per plant than OP varieties. Nectar production and amount of nectar sugar in OSR plants were independent of number and size of flowers. Our data show that floral traits of glasshouse‐grown OSR differed among breeding systems, suggesting that manipulation and enhancement of nectar rewards for insect flower visitors, including pollinators, could be included in future OSR breeding programmes.

Strong Phenotypic Variation in Floral Design and Display Traits of an Annual Tarweed in Relation to Small-Scale Topographic Heterogeneity in Semiarid Chile

Floral trait expression in wild populations varies in relation to environmental gradients. Variation can be observed among and within plant populations. We investigated the changes in floral phenotype within populations and the probability of plant pollinator visits in relation to small-scale variation in soil moisture and plant cover in a semiarid ecosystem. We measured the variability of floral phenotypes of three wild populations of Madia chilensis along a gradient constituted by three topographic positions (south-facing slope, north-facing slope, and ravine). Changes in soil moisture, plant density, leaf water content, and internode elongation were measured for one population over two study years. Pollinator visit probabilities were also estimated. Floral phenotypes were strongly segregated among topographic positions but less segregated among populations. Plants with the lowest water contents and the smallest or least-conspicuous flower heads grew in the drier north-facing slope, while plants with the highest water contents and showier flower design and display grew in the wetter ravine. Probabilities of pollinator visits were higher on the north-facing slope. Differences in floral phenotypes and probabilities of pollinator visits of M. chilensis were strongly connected to topographic variations at small spatial scale. Topographic heterogeneity at small spatial scale could thus have important consequences for floral trait evolution and pollination ecology.

Experimental floral and inflorescence trait manipulations affect pollinator preference and function in a hummingbird‐pollinated plant

Controversy is ongoing regarding the importance of pollinator-mediated selection as a source of observed patterns of floral diversity. Although increasing evidence exists of pollinator-mediated selection acting on female reproductive success, there is still limited understanding of pollinator-mediated selection on floral traits via male reproductive success. Here we quantify potential selection by the ruby-throated hummingbird, Archilochus colubris, on four floral traits of hermaphroditic Silene exerted through male floral function. In single trait manipulative experiments we quantified hummingbird visitation preference and/or fluorescent dye (a pollen analog) donation as a function of number of flowers displayed (inflorescence size), height of the floral display (inflorescence height), floral color, and corolla tube length. Hummingbirds preferred to visit larger floral displays and floral displays at greater height, likely representing a general pollinator preference for larger, more visible signals and/or greater rewards. In addition, hummingbirds preferred to visit red flowers, and male function was greater in flowers manipulated to have longer corolla tubes. Selection pressures exerted by hummingbirds on S. virginica floral and inflorescence design through male reproductive success are consistent with the contemporary expression of floral traits of S. virginica relative to related Silene species with different pollinators, and they are consistent with the hummingbird syndrome of traits expressed by S. virginica.

Leaf herbivory and drought stress affect floral attractive and defensive traits in Nicotiana quadrivalvis

Adaptive phenotypic plasticity allows sessile organisms such as plants to match trait expression to the particular environment they experience. Plasticity may be limited, however, by resources in the environment, by responses to prior environmental cues, or by previous interactions with other species, such as competition or herbivory. Thus, understanding the expression of plastic traits and their effects on plant performance requires evaluating trait expression in complex environments, rather than across levels of a single variable. In this study, we tested the independent and combined effects of two components of a plant's environment, herbivory and water availability, on the expression of attractive and defensive traits in Nicotiana quadrivalvis in the greenhouse. Damage and drought did not affect leaf nicotine concentrations but had additive and non-additive effects on floral attractive and defensive traits. Plants in the high water treatment produced larger flowers with more nectar than in the low water treatment. Leaf damage induced greater nectar volumes in the high water treatment only, suggesting that low water limited plastic responses to herbivore damage. Leaf damage also tended to induce higher nicotine concentrations in nectar, consistent with other studies showing that leaf damage can induce floral defenses. Our results suggest that there are separate and synergistic effects of leaf herbivory and drought on floral trait expression, and thus plasticity in response to complex environments may influence plant fitness via effects on floral visitation and defense.

A global test of the pollination syndrome hypothesis.

'Pollination syndromes' are suites of phenotypic traits hypothesized to reflect convergent adaptations of flowers for pollination by specific types of animals. They were first developed in the 1870s and honed during the mid 20th Century. In spite of this long history and their central role in organizing research on plant-pollinator interactions, the pollination syndromes have rarely been subjected to test. The syndromes were tested here by asking whether they successfully capture patterns of covariance of floral traits and predict the most common pollinators of flowers. Flowers in six communities from three continents were scored for expression of floral traits used in published descriptions of the pollination syndromes, and simultaneously the pollinators of as many species as possible were characterized. Ordination of flowers in a multivariate 'phenotype space' defined by the syndromes showed that almost no plant species fall within the discrete syndrome clusters. Furthermore, in approximately two-thirds of plant species, the most common pollinator could not be successfully predicted by assuming that each plant species belongs to the syndrome closest to it in phenotype space. The pollination syndrome hypothesis as usually articulated does not successfully describe the diversity of floral phenotypes or predict the pollinators of most plant species. Caution is suggested when using pollination syndromes for organizing floral diversity, or for inferring agents of floral adaptation. A fresh look at how traits of flowers and pollinators relate to visitation and pollen transfer is recommended, in order to determine whether axes can be identified that describe floral functional diversity more successfully than the traditional syndromes.

Modification of flower architecture during early stages in the evolution of self-fertilization.

The evolution of selfing from outcrossing is characterized by a series of morphological changes to flowers culminating in the selfing syndrome. However, which morphological traits initiate increased self-pollination and which are accumulated after self-fertilization establishes is poorly understood. Because the expression of floral traits may depend on the conditions experienced by an individual during flower development, investigation of changes in mating system should also account for environmental and developmental factors. Here, early stages in the evolution of self-pollination are investigated by comparing floral traits among Brazilian populations of Eichhornia paniculata (Pontederiaceae), an annual aquatic that displays variation in selfing rates associated with the breakdown of tristyly to semi-homostyly. Thirty-one Brazilian populations under uniform glasshouse conditions were compared to investigate genetic and environmental influences on flower size and stigma-anther separation (herkogamy), two traits that commonly vary in association with transitions to selfing. Within-plant variation in herkogamy was also examined and plants grown under contrasting environmental conditions were compared to examine to what extent this trait exhibits phenotypic plasticity. In E. paniculata a reduction in herkogamy is the principal modification initiating the evolution of selfing. Significantly, reduced herkogamy was restricted to the mid-styled morph and occurred independently of flower size. Significant genetic variation for herkogamy was detected among populations and families, including genotypes exhibiting developmental instability of stamen position with bimodal distributions of herkogamy values. Cloned genets exposed to contrasting growth conditions demonstrated environmental control of herkogamy and genotypic differences in plasticity of this trait. The ability to modify herkogamy independently of other floral traits, genetic variation in the environmental sensitivity of herkogamy, and the production of modified and unmodified flowers within some individuals, reveal the potential for dynamic control of the mating system in a species that commonly confronts heterogeneous aquatic environments.

Foliar damage modifies floral attractiveness to pollinators in Alstroemeria exerens

Pollination is a requisite for sexual reproduction in plants and its success may determine the reproductive output of individuals. Pollinator preference for some floral designs or displays that are lacking or poorly developed in focal plants may constrain the pollination process. Foliar herbivory may affect the expression of floral traits, thus reducing pollinator attraction. Natural populations of the Andean species Alstroemeria exerens (Alstromeriaceae) in central Chile show high levels of foliar herbivory, and floral traits show phenotypic variation. In the present field study, we addressed the attractive role of floral traits in A. exerens and the effect of foliar damage on them. Particularly, we posed the following questions: (1) Is there an association between floral display and design traits and the number and duration of pollinator visits? and (2) Does foliar damage affect the floral traits associated with pollinator visitation? To assess the attractiveness of floral traits for pollinators, we recorded the number and duration of visits in 101 focal plants. To evaluate the effects of foliar damage on floral traits, 100 plants of similar size were randomly assigned to control or damage groups during early bud development. Damaged plants were clipped using scissors (50% of leaf area) and control plants were manually excluded from natural herbivores (<5% of leaf area eaten). During the peak of flowering, we recorded the number of open flowers, and estimated corolla and nectar guide areas. The number and duration of pollinator visits was statistically associated with floral design and display traits. Plants with larger displays, corollas and nectar guide areas received more visits. Visits lasted longer as display increases. In addition, foliar damage affected attractive traits. Damaged plants had fewer open flowers and smaller nectar guide areas. We conclude that foliar damage affects plant attractiveness for pollinators and hence may indirectly affect plant fitness.

PLASTICITY AND ENVIRONMENT-SPECIFIC COVARIANCES: AN INVESTIGATION OF FLORAL–VEGETATIVE AND WITHIN FLOWER CORRELATIONS

Floral traits are commonly thought to be more canalized than vegetative ones. In addition, floral and vegetative traits are hypothesized to be genetically decoupled, enabling vegetative structures to respond plastically to environmental heterogeneity, and to evolve in response to selection without disrupting the reproductive function of flowers. To test these hypotheses, we evaluate the genetic architecture of floral and vegetative traits in natural populations of Arabidopsis thaliana raised under variable light-quality environments. Plants were grown either under high or low ratios of red to far-red (R:FR) light, an aspect of light quality that varies with neighbor proximity and regulates competitive shade-avoidance responses. Across environments, we detected significant genetic variation for the average expression of all measured floral traits (petal length and width, stamen length, pistil length, stigma-anther separation, and exsertion of both the stamen and pistil beyond the corolla). Light quality significantly influenced the absolute size of several floral traits as well as the allometry (i.e., relative scaling) of all floral traits, and genotypes differed in the plasticity of floral traits to the light treatments. Exposure to low relative to high R:FR resulted in significantly greater elongation in the vegetative trait, petiole length, and genotypes again differed in the plasticity of this trait to R:FR. Consistent with prior studies, most floral traits were less plastic than the vegetative trait; herkogamy (i.e., stigma-anther separation) was the exception and expressed more variable trait values across environments than petiole length, apparently as a consequence of the independent responses of stamens and pistils. Flowers also showed strong phenotypic integration; genotypic correlations were significantly positive among floral traits within each light treatment. Although floral-vegetative correlations were not significant in the high R:FR light treatment, significant correlations were detected between petal traits, pistil length, and petiole length under low R:FR, in contrast to the widely held hypothesis that floral and vegetative traits are genetically independent. Finally, we detected selection for reduced herkogamy in the low R:FR light treatment. The observed correlation between functional trait groups suggest that vegetative plasticity may affect the expression of floral traits in some environments, and that environment-specific constraints may exist on the evolution of floral and vegetative traits.

Nectar reward and advertisement in hummingbird‐pollinated Silene virginica (Caryophyllaceae)

We tested for an association between nectar and various floral traits and investigated their roles as primary and secondary pollinator attractants in hummingbird-pollinated Silene virginica. Our goal was to gain insight into the mechanisms of pollinator-mediated selection that underlies floral trait divergence within the genus. In a field population of S. virginica, we measured five floral and eight vegetative traits and quantified nectar volume, nectar sugar concentration, and total sugar reward (nectar volume × nectar sugar concentration). All three components of nectar reward were positively correlated to flower size, and nectar volume varied significantly among individuals within the population. To ascertain whether the correlation of specific floral traits with nectar reward influences the behavior of the primary pollinator of S. virginica, the ruby-throated hummingbird, Archilochus colubris, we investigated whether A. colubris preferred the expression of floral traits associated with high nectar volume and total sugar reward. We accomplished this by constructing floral arrays consisting of artificial flowers that had equal nectar quantity and total sugar reward but that differed in petal area and corolla tube diameter, which were positively correlated with nectar quantity and total sugar reward in our field study. In observations of visitation frequencies to the various floral-trait combinations, hummingbirds preferentially visited artificial floral phenotypes with larger petal displays, with the greatest preference for floral phenotypes with both larger petals and wider corolla-tube diameters. This association between primary and secondary floral attractants and hummingbird discrimination of floral features supports the concept that the floral traits of S. virginica reflect pollinator-mediated selection by the principal pollinator.

Floral trait expression and plant fitness in response to below- and aboveground plant–animal interactions

Although plant–animal interactions like pollination and herbivory are obviously interdependent, ecological investigations focus mainly on one kind of interaction ignoring the possible significance of the others. Plants with flowers offer an extraordinary possibility to study such mutualistic and antagonistic interactions since it is possible to measure changes in floral traits and fitness components in response to different organisms or combinations of them. In a three factorial common garden experiment we investigated single and combined effects of root herbivores, leaf herbivores and decomposers on flowering traits and plant fitness of Sinapis arvensis. Leaf herbivory negatively affected flowering traits indicating that it could significantly affect plant attractiveness to pollinators. Decomposers increased total plant biomass and seed mass indicating that plants use the nutrients liberated by decomposers to increase seed production. We suggest that S. arvensis faced no strong selection pressure from pollen limitation, for two reasons. First, reduced nutrient availability through leaf herbivory affected primarily floral traits that could be important for pollinator attraction. Second, improved nutrient supply through decomposer activity was invested in seed production and not in floral traits. This study indicates the importance of considering multiple plant–animal interactions simultaneously to understand selection pressures underlying plant traits and fitness.

The neighborhood matters: effects of neighbor number and sibling (or kin) competition on floral traits in Spergularia marina (Caryophyllaceae).

Effects of abiotic factors on the expression of floral and gender traits have been well documented in wild plant species; by contrast, little is known of the effect(s) on an individual's floral phenotype of the genetic composition of its neighboring conspecifics. Here we report the results of a greenhouse experiment conducted to detect the effects of genetic attributes of an individual's local environment on the expression of floral traits in the selfing annual, Spergularia marina (Caryophyllaceae). First, to test the hypothesis that negative effects of intraspecific competition are stronger when an individual competes with genetically similar individuals than when it competes with unrelated genotypes, we evaluated the effects of the number of nearest-neighbor kin (vs. unrelated individuals) on floral traits. Plants adjacent to two kin produced significantly fewer stamens per flower than plants adjacent to one or no siblings, indicating that kin competition reduced allocation to male function. Second, to test the hypothesis that the genetic diversity of a neighborhood influences the phenotype of a focal plant, we determined whether the number of maternal families represented among an individual's nearest neighbors influences its phenotype. The number of maternal families surrounding a focal plant did not affect floral trait expression. These results suggest that in S. marina, male function is more sensitive than female function to the genetic environment and that as an individual's genetic similarity to its neighbors increases, so do the negative effects of competition for limited resources.