Uni-directional Interaction and Plant–Pollinator–Robber Coexistence

A mathematical model for the plant-pollinator-robber interaction is studied to understand the factors leading to the widespread occurrence and stability of such interactions. In the interaction, a flowering plant provides resource for its pollinator and the pollinator has both positive and negative effects on the plant. A nectar robber acts as a plant predator, consuming a common resource with the pollinator, but with a different functional response. Using dynamical systems theory, mechanisms of species coexistence are investigated to show how a robber could invade the plant-pollinator system and persist stably with the pollinator. In addition, circumstances are demonstrated in which the pollinator's positive and negative effects on the plant could determine the robber's invasibility and the three-species coexistence.     

Invasibility of Nectarless Flowers in Plant–Pollinator Systems

This paper considers plant–pollinator systems in which plants are divided into two categories: The plants that secret a substantial volume of nectar in their flowers are called secretors, while those without secreting nectar are called nonsecretors (cheaters). The interaction between pollinators and secretors is mutualistic, while the interaction between pollinators and nonsecretors is parasitic. Both interactions can be described by Beddington–DeAngelis functional responses. Using dynamical systems theory, we show global dynamics of a pollinator–secretor–cheater model and demonstrate mechanisms by which nectarless flowers/nonsecretors can invade the pollinator–secretor system and by which the three species could coexist. We define a threshold in the nonsecretors’ efficiency in translating pollinator–cheater interaction into fitness, which is determined by parameters (factors) in the systems. When their efficiency is above the threshold, non-secretors can invade the pollinator–secretor system. While the nonsecretors’ invasion often leads to their persistence in pollinator–secretor systems, the model demonstrates situations in which the non-secretors’ invasion can drive secretors into extinction, which consequently leads to extinction of the nonsecretors themselves.     

Proteomics Study of COI1-regulated Proteins in Arabidopsis Flower

Jasmonates(JAs) are a new class of plant hormone that regulate expression of diverse genes to mediate various plant responses.The Arabidopsis F-box protein COI1 is required for plant defense and male fertility in JA signal pathway.To further investigate the regulatory role of COI1 in male fertility,we compared the proteomics profiles of Arabidopsis wild type(WT) flowers with coi1-1 mutan male-sterile flowers using two-dimensional difference gel electrophoresis coupled with matrix-assisted laser desoption/ionization-time-of-flight mass spectrometry.Sixteen proteins with potential function in specific biological processes such as metabolism processes and defense/stress responses were differentially expressed in WT and coi1-1 mutant flowers.Verification on a phi class glutathione transferase AtGSTF9,one out of these 16 identified proteins,revealed that the expression of AtGSTF9 was severely downregulated in flowers of coi1-1 mutant compared with that of WT.Further function analyses of these genes would provide new insights into the molecular basis of COI1-regulated male fertility.     

Virus Infection of Plants Alters Pollinator Preference: A Payback for Susceptible Hosts?

Plant volatiles play important roles in attraction of certain pollinators and in host location by herbivorous insects. Virus infection induces changes in plant volatile emission profiles, and this can make plants more attractive to insect herbivores, such as aphids, that act as viral vectors. However, it is unknown if virus-induced alterations in volatile production affect plant-pollinator interactions. We found that volatiles emitted by cucumber mosaic virus (CMV)-infected tomato (Solanum lycopersicum) and Arabidopsis thaliana plants altered the foraging behaviour of bumblebees (Bombus terrestris). Virus-induced quantitative and qualitative changes in blends of volatile organic compounds emitted by tomato plants were identified by gas chromatography-coupled mass spectrometry. Experiments with a CMV mutant unable to express the 2b RNA silencing suppressor protein and with Arabidopsis silencing mutants implicate microRNAs in regulating emission of pollinator-perceivable volatiles. In tomato, CMV infection made plants emit volatiles attractive to bumblebees. Bumblebees pollinate tomato by ‘buzzing’ (sonicating) the flowers, which releases pollen and enhances self-fertilization and seed production as well as pollen export. Without buzz-pollination, CMV infection decreased seed yield, but when flowers of mock-inoculated and CMV-infected plants were buzz-pollinated, the increased seed yield for CMV-infected plants was similar to that for mock-inoculated plants. Increased pollinator preference can potentially increase plant reproductive success in two ways: i) as female parents, by increasing the probability that ovules are fertilized; ii) as male parents, by increasing pollen export. Mathematical modeling suggested that over a wide range of conditions in the wild, these increases to the number of offspring of infected susceptible plants resulting from increased pollinator preference could outweigh underlying strong selection pressures favoring pathogen resistance, allowing genes for disease susceptibility to persist in plant populations. We speculate that enhanced pollinator service for infected individuals in wild plant populations might provide mutual benefits to the virus and its susceptible hosts.     

Going native? Flower use by bumblebees in English urban gardens

Background and Aims

Although urban gardens provide opportunities for pollinators in an otherwise inhospitable environment, most garden plants are not native to the recipient biogeographical region and their value to local pollinators is disputed. This study tested the hypothesis that bumblebees foraging in English urban gardens preferentially visited sympatric Palaearctic-range plants over species originating outside their native range.

Methods

Twenty-seven surveys of flower availability and bumblebee visitation (Bombus spp.) were conducted over a 3-month summer period. Plants were categorized according to whether they were native British, Palaearctic or non-Palaearctic in origin. A phylogeny of the 119 plant species recorded was constructed and the relationship between floral abundance and the frequency of pollinator visits investigated by means of phylogenetically independent contrasts. Differentiation in utilization of plant species by the five bumblebee species encountered was investigated using niche overlap analyses.

Key Results

There was conflicting evidence for preferential use of native-range Palaearctic plant species by bumblebees depending on which plants were included in the analysis. Evidence was also found for niche partitioning between species based on respective preferences for native and non-native biogeographical range plants. Two bumblebees (Bombus terrestris and B. pratorum) concentrated their foraging activity on non-Palaearctic plants, while two others (B. hortorum and B. pascourum) preferred Palaearctic species.

Conclusions

The long-running debate about the value of native and non-native garden plants to pollinators probably stems from a failure to properly consider biogeographical overlap between plant and pollinator ranges. Gardeners can encourage pollinators without consideration of plant origin or bias towards ‘local’ biogeographical species. However, dietary specialist bumblebees seem to prefer plants sympatric with their own biogeographical range and, in addition to the cultivation of these species in gardens, provision of native non-horticultural (‘weed’) species may also be important for pollinator conservation.     

‘Bee Hotels’ as Tools for Native Pollinator Conservation: A Premature Verdict?

Society is increasingly concerned with declining wild bee populations. Although most bees nest in the ground, considerable effort has centered on installing ‘bee hotels’—also known as nest boxes or trap nests—which artificially aggregate nest sites of above ground nesting bees. Campaigns to ‘save the bees’ often promote these devices despite the absence of data indicating they have a positive effect. From a survey of almost 600 bee hotels set up over a period of three years in Toronto, Canada, introduced bees nested at 32.9% of sites and represented 24.6% of more than 27,000 total bees and wasps recorded (47.1% of all bees recorded). Native bees were parasitized more than introduced bees and females of introduced bee species provisioned nests with significantly more female larva each year. Native wasps were significantly more abundant than both native and introduced bees and occupied almost 3/4 of all bee hotels each year; further, introduced wasps were the only group to significantly increase in relative abundance year over year. More research is needed to elucidate the potential pitfalls and benefits of using bee hotels in the conservation and population dynamics of wild native bees.     

Pollinator shifts between Ophrys sphegodes populations: might adaptation to different pollinators drive population divergence?

Local adaptation to different pollinators is considered one of the possible initial stages of ecological speciation as reproductive isolation is a by‐product of the divergence in pollination systems. However, pollinator‐mediated divergent selection will not necessarily result in complete reproductive isolation, because incipient speciation is often overcome by gene flow. We investigated the potential of pollinator shift in the sexually deceptive orchids Ophrys sphegodes and Ophrys exaltata and compared the levels of floral isolation vs. genetic distance among populations with contrasting predominant pollinators. We analysed floral hydrocarbons as a proxy for floral divergence between populations. Floral adoption of pollinators and their fidelity was tested using pollinator choice experiments. Interpopulation gene flow and population differentiation levels were estimated using AFLP markers. The Tyrrhenian O. sphegodes population preferentially attracted the pollinator bee Andrena bimaculata, whereas the Adriatic O. sphegodes population exclusively attracted A. nigroaenea. Significant differences in scent component proportions were identified in O. sphegodes populations that attracted different preferred pollinators. High interpopulation gene flow was detected, but populations were genetically structured at species level. The high interpopulation gene flow levels independent of preferred pollinators suggest that local adaptation to different pollinators has not (yet) generated detectable genome‐wide separation. Alternatively, despite extensive gene flow, few genes underlying floral isolation remain differentiated as a consequence of divergent selection. Different pollination ecotypes in O. sphegodes might represent a local selective response imposed by temporal variation in a geographical mosaic of pollinators as a consequence of the frequent disturbance regimes typical of Ophrys habitats.     

Searching for Spatial Patterns in a Pollinator–Plant–Herbivore Mathematical Model

This paper deals with the spatio-temporal dynamics of a pollinator–plant–herbivore mathematical model. The full model consists of three nonlinear reaction–diffusion–advection equations defined on a rectangular region. In view of analyzing the full model, we firstly consider the temporal dynamics of three homogeneous cases. The first one is a model for a mutualistic interaction (pollinator–plant), later on a sort of predator–prey (plant–herbivore) interaction model is studied. In both cases, the interaction term is described by a Holling response of type II. Finally, by considering that the plant population is the unique feeding source for the herbivores, a mathematical model for the three interacting populations is considered. By incorporating a constant diffusion term into the equations for the pollinators and herbivores, we numerically study the spatiotemporal dynamics of the first two mentioned models. For the full model, a constant diffusion and advection terms are included in the equation for the pollinators. For the resulting model, we sketch the proof of the existence, positiveness, and boundedness of solution for an initial and boundary values problem. In order to see the separated effect of the diffusion and advection terms on the final population distributions, a set of numerical simulations are included. We used homogeneous Dirichlet and Neumann boundary conditions.     

Pollinator attraction in Anacamptis papilionacea (Orchidaceae): a food or a sex promise?

The Mediterranean orchid Anacamptis papilionacea , despite showing a typical food-deceptive floral display, has also been reported to frequently attract male pollinators, suggesting a potentiality for sexual attraction. In a survey from a southern Italian population of A. papilionacea and their hybrids with Anacamptis morio , we collected 37 pollinators belonging to five bee species carrying 126 orchid pollinia. The main pollinator of A. papilionacea was Anthophora crinipes male (48.6%), but the number of females was not negligible (22.9%). We also found pollinator sharing between the hybrid and the parental species. Our findings confirm that, contrary to other food-deceptive species, A. papilionacea mainly attracts male insects, but also that, in contrast to sexually deceptive species, this attraction is not specific. We suggest that A. papilionacea adopts a complex mix of food and sexually deceptive pollination and could represent a helpful model for studying the transition between different pollination strategies.     

Stigmatic exudate in the Annonaceae: Pollinator reward,pollen germination medium or extragynoecial compitum?

Although “dry‐type” stigmas are widely regarded as ancestral in angiosperms, the early‐divergent family Annonaceae has copious stigmatic exudate. We evaluate three putative functions for this exudate: as a nutritive reward for pollinators; as a pollen germination medium; and as an extragynoecial compitum that enables pollen tube growth between carpels. Stigmatic exudate is fructose dominated (72.2%), but with high levels of glucose and sucrose; the dominance of hexose sugars and the diversity of amino acids observed, including many that are essential for insects, support a nutritive role for pollinators. Sugar concentration in pre‐receptive flowers is high (28.2%), falling during the peak period of stigmatic receptivity (17.4%), and then rising again toward the end of the pistillate phase (32.9%). Pollen germination was highest in sugar concentrations <20%. Sugar concentrations during the peak pistillate phase therefore provide optimal osmolarity for pollen hydration and germination; subsequent changes in sugar concentration during anthesis reinforce protogyny (in which carpels mature before stamens), enabling the retention of concentrated exudate into the staminate phase as a pollinator food reward without the possibility of pollen germination. Intercarpellary growth of pollen tubes was confirmed: the exudate therefore also functions as a suprastylar extragynoecial compitum, overcoming the limitations of apocarpy.     

Does Plant Origin Influence the Fitness Impact of Flower Damage? A Meta-Analysis

Herbivory has been long considered an important component of plant-animal interactions that influences the success of invasive species in novel habitats. One of the most important hypotheses linking herbivory and invasion processes is the enemy-release hypothesis, in which exotic plants are hypothesized to suffer less herbivory and fitness-costs in their novel ranges as they leave behind their enemies in the original range. Most evidence, however, comes from studies on leaf herbivory, and the importance of flower herbivory for the invasion process remains largely unknown. Here we present the results of a meta-analysis of the impact of flower herbivory on plant reproductive success, using as moderators the type of damage caused by floral herbivores and the residence status of the plant species. We found 51 papers that fulfilled our criteria. We also included 60 records from unpublished data of the laboratory, gathering a total of 143 case studies. The effects of florivory and nectar robbing were both negative on plant fitness. The methodology employed in studies of flower herbivory influenced substantially the outcome of flower damage. Experiments using natural herbivory imposed a higher fitness cost than simulated herbivory, such as clipping and petal removal, indicating that studies using artificial herbivory as surrogates of natural herbivory underestimate the real fitness impact of flower herbivory. Although the fitness cost of floral herbivory was high both in native and exotic plant species, floral herbivores had a three-fold stronger fitness impact on exotic than native plants, contravening a critical element of the enemy-release hypothesis. Our results suggest a critical but largely unrecognized role of floral herbivores in preventing the spread of introduced species into newly colonized areas.     

Mutation in Flower Colour and Shape of Chrysanthemum morifolium Induced by γ-Radiation

Flowers of Chrysanthemum morifolium Ramat cv. Lalima were greyed red and florets were flat spoon shaped. Ray florets after inoculation on the Murashige and Skoog's medium supplemented with 1.07 μM α-naphthaleneacetic acid and 8.87 μM benzyladenine were irradiated with γ-radiation (0.5 Gy and 1 Gy). All the regenerated shoots either from control or from γ-irradiated florets were isolated, rooted and transplanted in the field after hardening. Two mutants were obtained in the γ-irradiated plants (0.5 Gy). Both the mutants were yellow coloured but one having flat spoon shaped ray florets similar to the original cultivar, while the other having tubular florets. Both the mutants were propagated vegetatively and have produced true-to-type flowers. This revised version was published online in July 2006 with corrections to the Cover Date.     

Pollinator Shift and Reproductive Performance of the Qinghai–Tibetan Plateau Endemic and Endangered Swertia przewalskii (Gentianaceae)

Reproductive failure results in many plant species becoming endangered. However, little is known of how and to what extent pollinator shifts affect reproductive performance of endangered species as a result of the artificial introduction of alien insects. In this study we examined breeding systems, visitor species, visiting frequency and seed set coefficients of Swertia przewalskii in two years that had different dominant pollinator species (native vs. alien). Flowers of this species were protandrous and herkogamous and insects were needed for the production of seeds. The stigmatic receptivity of this species was shorter than for other gentians. No significant difference in seed set coefficient was found for hand-pollinated plants between the two years, indicating that pollinator shift only had a minor effect on this plant’s breeding system. The commonest pollinators in 2002 were native bumblebees, alien honeybees and occasional solitary bees, however, only alien honeybees were observed in 2004. The flower visitation rate in both years was relatively high, although the total visit frequency decreased significantly in 2004. The control flowers without any treatment produced significantly fewer seed sets in 2004 than in 2002. In the past decade the seed production of this species may have partly decreased due to pollination by alien honeybees, however, we suggest that they might have acted as alternative pollinators ensuring seed production of S. przewalskii when native pollinators were unavailable. The main reason that this plant is endangered is probably the result of habitat destruction, but changes in land use, namely intensified agricultural practice and unfavorable animal husbandry have also contributed to its decline. We recommend that in-situ conservation, including the establishment of a protected area, is the best way to preserve this species effectively.     

Flower choice by naïve young crab spiders and the effect of subsequent experience

Initial responses of na?ve individuals to critical environmental stimuli provide important information about the innate contribution to behaviour, and subsequent responses to the same stimuli may show the role of experience in mediating those initial responses. To test the role of these factors, I measured initial patch choices and giving-up responses of just-emerged, na?ve, second-instar crab spiders, Misumena vatia, on several hunting sites they encountered after leaving their natal nests. In follow-up tests I measured the effects of these experiences on subsequent patch choice decisions. The choice of hunting sites is a vital decision at all stages of the life cycle for sit-and-wait predators such as Misumena. In their initial tests these spiderlings remained more frequently on goldenrod (Solidago spp.) flowers than on green or yellow goldenrod buds, a preference they retained through tests run on 5 consecutive days. Individuals on green and yellow buds shifted sites more quickly and frequently than those from flowers, and made most of these moves to flowers, which attracted many more prey than did buds. These differences were not affected by age, energetic condition, or loss of information over the period of the experiment. Once spiderlings moved from buds, they showed a high, increasing tendency to move from buds in subsequent runs, those from flowers showed a consistently low tendency. These results suggest that spiderlings retain their innate behavioural patterns through the second instar, but that experience also plays a modest role in patch choice at this stage. Copyright 2000 The Association for the Study of Animal Behaviour.     

Pollinator abundance,morphology, and flower visitation rate: analysis of the “quantity” component in a plant-pollinator system

Summary Abundance and flower visitation rate of the pollinators of Lavandula latifolia (Labiatae), an insect-pollinated shrub, were studied over a 6-year period. The objective was to elucidate interspecific patterns in the quantity component of the plant-pollinator interaction. A total of 54 insect taxa are considered in the analyses, including hynenopterans, dipterans and lepidopterans. Most pollinators were comparatively scarce, with a few taxa acounting collectively for the majority of individuals. Pollinators differed broadly in flower visitation rate (0.2–30 flowers/min). Most of this variation was explained by differences in flower handling time (HT). Regardless of proboscis length, hymenopterans had intrinsically shorter handling times than lepidopterans. Within each group, HT decreased exponentially with increasing proboscis length. Abundance and visitation rate were uncorrelated across pollinator taxa. The total number of visits that each pollinator contributed to the plant (NFV) was estimated as the product of abundance x visitation rate. NFV values spanned four orders of magnirade. A small, taxonomically diverse group of species (1 moth, 1 butterfly, 4 bees) accounted for most visits and thus could effectively exert some selection on floral features. Nevertheless, the morphological diversity represented in this group of dominant pollinators probably constrains plant specialization, as they will most likely select for different floral features or in opposing directions on the same traits.     

Pollinator aggregative and functional responses to flower density: does pollinator response to patches of plants accelerate at low‐densities?

Plant reproduction is often reduced at low densities, due to reduced pollinator visitation rates. Recent theory suggests that a disproportionate increase in pollinator visits to patches of plants as heterospecific plant density increases (i.e. if visitation is a sigmoid function of patch density) can rescue sparse populations of a focal plant species from reduced reproductive success or population decline. A field experiment was performed to determine the shape of the pollinator visitation response to patches of differing density of the common weed Brassica rapa . Both the aggregative and functional response for the entire pollinator community were saturating rather than sigmoid, indicating that pollinator response does not accelerate when density increases. The results for the entire pollinator community were consistent among temporal and spatial replicates. Aggregative response curves for specific pollinator taxa were either linear (bombyliid flies) or saturating (syrphid flies, solitary bees, and Lepidoptera). Functional responses for these taxa were saturating (syrphid flies and solitary bees) or flat (bombyliid flies and Lepidoptera). Individual pollinators visited more plants during foraging bouts in high-density patches, but visits per plant decreased. Seeds per fruit and seeds per flower increased with increasing density. There is no evidence that pollinators disproportionately visit denser patches, or that the conditions for this mechanism of pollination facilitation are likely to be met in this generalist pollinator system.     

Flower formation in the short-day plant Kalanchoë by grafting with a long-day and a short-long-day Echeveria

Flower formation was induced in the shortday plant Kalanchoë blossfeldiana Poellnitz under long-day conditions by grafting with flowering shoots of the short-long-day plant Echeveria harmsii (Rose) MacBr. and of the long-day plant Echeveria pulvoliver (E. pulvinata Rose x E. harmsii). Vegetative shoots from induced Echeveria plants failed to cause a flowering response in Kalanchoë. The presence of flowering and vegetative shoots side by side on induced Echeveria plants provides evidence for physiological chimeras in this genus.Abbreviations LD long day(s) - SD short day(s) - LDP long-day plant(s) - LSDP long-short-day plant(s) - SDP short-day plant(s) - SLDP short-long-day plant(s)     

Plant visibility through mobile learning? Implementation and evaluation of an interactive Flower Hunt in a botanic garden

Plants have an enormous importance for life on earth but are often overlooked. This phenomenon called plant blindness is reinforced as students prefer to spend their recreation time with modern media rather than in nature. Mobile devices have become an omnipresent part of students’ everyday lives. Nevertheless, direct experiences with plants are needed to create an empathic link. We decided to combine mobile learning with plant education in the context of an excursion. Therefore, we created an interactive treasure hunt-like quiz for mobile devices through a botanic garden as an approach to positively influence students’ knowledge about and attitude towards plants and nature. The main topic was the variety of flowers which is why we called it the flower hunt. Besides the comprehension of plant diversity, its endangerment and protection, attention was given to flower morphology and ecology as well as to systematics. Our results showed increased performances especially in systematics. Students were motivated to take part and enjoyed the implementation. Although the unit did not entirely cure plant blindness it increased students’ environmental awareness. Thus, the flower hunt provides a new educational approach for promoting conservation.     

Flower symmetry and flower size variability: an examination of Berg’s hypotheses in an alpine meadow

植物花对称性与传粉系统密切相关, 花特征的变异性受到传粉者的选择作用。由于特化的传粉者对两侧对称的花的稳定选择, Berg假说认为两侧对称植物花大小的变异性比辐射对称植物的更低; 而且, 在传粉者的选择作用下花特征比植物营养特征明显有更低的变异性, 因为后者更易受环境影响。该文对青藏高原东部高山草甸植物群落的50种开花植物(包括19种两侧对称植物和31种辐射对称植物)的花和叶特征进行了测定和分析。结果表明不论是两侧对称植物还是辐射对称植物, 花大小的变异性都显著低于叶片大小的变异性, 表明传粉者对花施加的稳定选择有利于花的稳定性。但是, 辐射对称物种花大小的变异程度和两侧对称物种的相似, 即使在控制物种系统发育的影响后, 也没有发现显著的差异, 这与Berg假说不一致。高山生态系统中传粉者种类相对较少, 以熊蜂和蝇类为主, 传粉者的活动受局域气候环境影响较大, 因此传粉昆虫对植物花的选择作用强度可能有较大的变异性。