克隆生长对被子植物传粉过程的影响

克隆植物与其传粉者的相互作用是植物繁殖生态学的重要研究领域之一。植物克隆生长与有性繁殖通常相伴进行, 往往产生较大的花展示与复杂的克隆空间结构, 通过传粉过程对有性繁殖过程产生影响, 共同决定植物的适合度。本文回顾了克隆生长对被子植物传粉过程影响的国内外研究进展, 从植物克隆大小、花资源空间配置、克隆构型与种群遗传结构四个方面讨论了克隆生长对传粉过程的影响及其生态学与进化生物学意义。早期研究预期, 随着克隆增大, 同株异花授粉水平增加, 因而通过增大自交率或花粉阻塞效应降低植物的适合度。但是, 后来的一些模拟与野外实验研究发现, 传粉者在同一克隆内访问的花数量并不会随克隆增大而一直增加, 访花行为也主要发生在分株内; 而且分子标记的自交率组分分析也表明自交主要发生在分株内。另一方面, 人工模型模拟以及传粉者访问行为研究表明, 当花朵数量相同时, 与所有花集中生长在同一分株上相比, 将花朵分散在多个分株上的克隆生长方式不会增加, 反而降低了同株异花授粉的发生水平。如果花序内花雌雄同熟, 花朵同时提供与接收花粉, 克隆生长会使植物接收到更高比例的异交花粉, 在提高后代质量的同时不增加同株异花授粉概率。这是从传粉生物学角度对植物克隆生长习性进化的一个全新的解释。今后, 克隆植物传粉生物学研究需要针对传粉者与克隆生长之间的相互作用建立理论模型, 探究克隆大小、克隆构型、花资源空间配置模式对传粉者访问频率和行为、花粉散布、交配格局的影响。同时, 需要在自然种群中, 尤其是克隆与非克隆的近缘类群、同一物种克隆与非克隆种群开展比较研究, 利用更高效的分子标记来研究克隆生长的生态与进化意义。     

An assessment of pollinator visitation to Banksia spinulosa

Abstract Remote photo-monitoring of Banksia spinulosa inflorescences indicated that birds, mammals and insects were regular visitors. These included the sugar glider, brown antechinus, eastern pygmy possum, eastern spinebill, honeybee and several moth species. Eastern spine-bills were recorded at all inflorescences monitored while visitation by other animals was more sporadic. Inflorescences were visited at least once every 24 h, and visitation frequency was approximately evenly distributed during the day and night. Examination of pollen tube growth from experimental treatments indicated that pollination success was similar from both nocturnal and diurnal visitors. However, nocturnal visitors were more effective at removing pollen from newly opened flowers. The behaviour of mammals at inflorescences was such that they would transfer much more pollen to flowers than other visitors, and probably contact receptive stigmas more often. Overall, mammals were considered to be slightly more effective pollinators than the more obvious daytime visitors, eastern spinebills. Although insects visited regularly, they were thought to be less effective at pollinating flowers than vertebrate visitors. Moths carried very little pollen, and the foraging behaviour of other insects was unlikely to promote much pollination.     

Agricultural landscapes with organic crops support higher pollinator diversity

Pollinators are traditionally thought to perceive non-flowering crop fields as hostile landscape matrix. In this study, we show that landscapes composed of higher proportions of organic crop fields support more bee species at greater abundances in fallow strips. An increase in organic cropping in the surrounding landscape from 5% to 20% enhanced bee species richness in fallow strips by 50%, density of solitary bees by 60% and bumble bee density by 150%. Bee species richness and bumble bee density responded strongest to organic cropping in landscape sectors with 500 m radius, solitary bee density in landscape sectors with 250 m radius. The most likely source of these results is that crop and noncrop habitats are strongly connected via bee foraging at the landscape scale. It seems likely that bees depending on nesting sites in fallow strips benefited from the more abundant flower resources provided by broadleaved weeds in organic crop fields. We conclude that the incorporation of organic crop fields into conventionally managed agricultural landscapes can provide food resources needed to sustain greater pollinator species richness in noncrop habitats.     

Linking pollinator visitation rate and pollen receipt

The majority of flowering plants require animals for pollination, a critical ecosystem service in natural and agricultural systems. However, quantifying useful estimates of pollinator visitation rates can be nearly impossible when pollinator visitation is infrequent. We examined the utility of an indirect measure of pollinator visitation, namely pollen receipt by flowers, using the hummingbird-pollinated plant, Ipomopsis aggregata (Polemoniaceae). Our a priori hypothesis was that increased pollinator visitation should result in increased pollen receipt by stigmas. However, the relationship between pollinator visitation rate and pollen receipt may be misleading if pollen receipt is a function of both the number of pollinator visits and variation in pollinator efficiency at depositing pollen, especially in the context of variable floral morphology. Therefore, we measured floral and plant characters known to be important to pollinator visitation and/or pollen receipt in I. aggregata (corolla length and width and plant height) and used path analysis to dissect and compare the effect of pollinator visitation rate vs. pollinator efficiency on pollen receipt. Of the characters we measured, pollinator visitation rate (number of times plants were visited multiplied by the mean percentage of flowers probed per visit) had the strongest direct positive effect on pollen receipt, explaining 36% of the variation in pollen receipt. Plant height had a direct positive effect on pollinator visitation rate and an indirect positive effect on pollen receipt. Despite the supposition that floral characters would directly affect pollen receipt as a result of changes in pollinator efficiency, corolla length and width only weakly affected pollen receipt. These results suggest a direct positive link between pollinator visitation rate and pollen receipt across naturally varying floral morphology in I. aggregata. Understanding the relationship between pollinator visitation rate and pollen receipt may be of critical importance in systems where pollinator visitation is difficult to quantify.     

Co-flowering neighbors influence the diversity and identity of pollinator groups visiting plant species

The generalization–specialization continuum exhibited in pollination interactions currently receives much attention. It is well-known that the pollinator assemblage of particular species varies temporally and spatially, and therefore the ecological generalization on pollinators may be a contextual attribute. However, the factors causing such variation and its ecological and evolutionary consequences are still poorly understood. This variation can be caused by spatial or temporal variation in the pollinator community, but also by variation in the plant community. Here, we examined how the floral neighbourhood influenced the generalization on pollinators and the composition of pollinators of six plant species differing in generalization levels and main pollinators. The diversity, identity and density of floral species affected both the level of generalization on pollinators and the composition of visitors of particular plant species. Although the relationships to floral neighbourhood varied considerably among species, generalization level and visitation by uncommon pollinators generally increased with floral diversity and richness. The generalization level of the neighbourhood was negatively related to the generalization level of the focal species in two species. The number of flowers of the pollinator-sharing species and the number of flowers of the focal species had different effects on the composition of visits in different species; attributable to differences in facilitation/competition for pollinator attraction. We propose that an important ecological implication of our results is that variation in species interactions caused by the pollination context may result in increased community stability. The main evolutionary implication of our results is that selection on flower and pollinator traits may depend, to an unknown extent, on the composition of the co-flowering plant community.     

The effects of floral display on pollinator visitation vary among populations ofPhacelia linearis (Hydrophyllaceae)

Summary Individual plants in gynodioecious populations ofPhacelia linearis (Hydrophyllaceae) vary in flower gender, flower size, and flower number. This paper reports the effects of variation in floral display on the visitation behaviour of this species' pollinators (mainly pollen-collecting solitary bees) in several natural and three experimental plant populations, and discusses the results in terms of the consequences for plant fitness. The working hypotheses were: (1) that because female plants do not produce pollen, pollen-collecting insects would visit hermaphrodite plants at a higher rate than female plants and would visit more flowers per hermaphrodite than per female; and (2) that pollinator arrival rate would increase with flower size and flower number, the two main components of visual display. These hypotheses were generally supported, but the effects of floral display on pollinator visitation varied substantially among plant populations. Hermaphrodites received significantly higher rates of pollinator arrivals and significantly higher rates of visits to flowers than did females in all experimental populations. Flower size affected arrival rate and flower visit rate positively in natural populations and in two of the three experimental populations. The flower size effect was significant only among female plants in one experimental population, and only among hermaphrodites in another. The effect of flower number on arrival rate was positive and highly significant in natural populations and in all experimental populations. In two out of three experimental populations, insects visited significantly more flowers per hermaphrodite than per female and visited more flowers on many-flowered plants than on few-flowered plants, but neither effect was detected in the third experimental population. Because seed production is not pollen-limited in this species, variation in pollinator visitation behaviour should mainly affect the male reproductive success of hermaphrodite plants. These findings suggest that pollinator-mediated natural selection for floral display inP. linearis varies in space and time.     

Functional flower traits and their diversity drive pollinator visitation

Recent studies have shown that the diversity of flowering plants can enhance pollinator richness and visitation frequency and thereby increase the resilience of pollination. It is assumed that flower traits explain these effects, but it is still unclear which flower traits are responsible, and knowing that, if pollinator richness and visitation frequency are more driven by mass‐ratio effects (mean trait values) or by trait diversity. Here, we analyse a three‐year data set of pollinator observations collected in a European grassland plant diversity experiment (The Jena experiment). The data entail comprehensive flower trait measurements, including reward traits (nectar and pollen amount), morphological traits (height, symmetry, area, colour spectra) and chemical traits (nectar‐amino acid and nectar‐sugar concentration). We test if pollinator species richness and visitation frequency of flower communities depend on overall functional diversity combining all flower traits within a community, single trait diversities (within trait variation) and community‐weighted means of the single traits, using Bayesian inference. Overall functional diversity did not affect pollinator species richness, but reduced visitation frequency. When looking at individual flower traits separately, we found that single trait diversity of flower reflectance and flower morphology were important predictors of pollinator visitation frequency. Moreover, independent of total flower abundance, community‐weighted means of flower height, area, reflectance, nectar‐sugar concentration and nectar‐amino acid concentration strongly affected both pollinator species richness and visitation frequency. Our results, challenge the idea that functional diversity always positively affects ecosystem functions. Nonetheless, we demonstrate that both single trait diversity and mass‐ratio effects of flower traits play an important role for diverse and frequent flower visits, which underlines the functionality of flower traits for pollination services.     

A meta-analysis of impacts of alien vs. native plants on pollinator visitation and reproductive success of co-flowering native plants

Alien plant species can alter pollinator visitation and, in turn, the sexual reproduction of natives. Using a conventional and a phylogenetically controlled meta-analytical approach on a data set of 40 studies, we evaluated the effect of alien neighbour plant species (aliens) on visitation to and reproduction of native co-flowering focal species (focals), and compared such effect to that of native neighbours (natives). An overall significantly negative effect of aliens on visitation to and reproduction of focals was confirmed. Interestingly, aliens differed from natives in their effect on visitation, but not on reproductive success. The negative effect of aliens on visitation and reproductive success increased at high relative alien plant abundance, but this increase was proportionally lower than the increase in relative plant abundance. Likewise, effect of aliens on visitation and reproductive success was most detrimental when alien and focal species had similar flower symmetry or colour. The phylogenetic relatedness between alien neighbours and focals influenced the reproductive success effect size. Results of the phylogenetic meta-analysis were only partly consistent with those of the conventional meta-analysis, depending on the response variable and on whether we controlled for the phylogeny of neighbour or focal species, which calls for special attention to control for species relatedness in this type of review. This study demonstrates the predominant detrimental impact of alien plants on pollination and reproduction of natives, and highlights the importance of phenotypic similarity to the outcome of the interaction.     

Molecular assays of pollen use consistently reflect pollinator visitation patterns in a system of flowering plants

Determining how pollinators visit plants vs. how they carry and transfer pollen is an ongoing project in pollination ecology. The current tools for identifying the pollens that bees carry have different strengths and weaknesses when used for ecological inference. In this study we use three methods to better understand a system of congeneric, coflowering plants in the genus Clarkia and their bee pollinators: observations of plant–pollinator contact in the field, and two different molecular methods to estimate the relative abundance of each Clarkia pollen in samples collected from pollinators. We use these methods to investigate if observations of plant–pollinator contact in the field correspond to the pollen bees carry; if individual bees carry Clarkia pollens in predictable ways, based on previous knowledge of their foraging behaviors; and how the three approaches differ for understanding plant–pollinator interactions. We find that observations of plant–pollinator contact are generally predictive of the pollens that bees carry while foraging, and network topologies using the three different methods are statistically indistinguishable from each other. Results from molecular pollen analysis also show that while bees can carry multiple species of Clarkia at the same time, they often carry one species of pollen. Our work contributes to the growing body of literature aimed at resolving how pollinators use floral resources. We suggest our novel relative amplicon quantification method as another tool in the developing molecular ecology and pollination biology toolbox.     

Crop management affects pollinator attractiveness and visitation in oilseed rape

Ecological intensification of agriculture implies managing ecological processes to improve performance of agricultural systems. However, impacts on relevant ecological functions such as insect pollination from other crop management factors are poorly explored. Pest insects and crop resources such as water availability can directly affect crop yields, but it is unknown if there are indirect effects through effects on insect pollination. With a factorial experiment, we examined how irrigation and control of pollen beetles affected crop attractiveness and pollinator visitation in an open-pollinated spring oilseed rape cultivar. We studied how irrigation and pest control modified the production of flowers and nectar in oilseed rape, and if this in turn affected the flower-visitation of honey bees and bumble bees. Pest control increased the number of oilseed rape flowers by 69%, and the amount of nectar per flower with 36%, but for the latter only in non-irrigated plots. Furthermore, we found higher pollinator densities in plots with reduced pollen beetle densities. Pest control also reduced the number of non-legitimate flower visits, suggesting higher pollination efficiency in plots with reduced pollen beetle densities. We show that crop management affects the value of mass-flowering crops as a resource for pollinating insects. Development of pest control tools that are harmless to pollinators could increase the value of flowering crops as food resources for pollinating insects.     

Co-flowering neighbor alters pollinator composition and influences reproductive success in a plant pollinated by multiple insects

Plant Ecology - The effect of the interactions among co-flowering plants for pollination has always been explained by changes in pollinator visiting frequency and/or foraging behavior. For most...     

持续干旱对樱桃根际土壤细菌数量及结构多样性影响

以1年生吉塞拉实生容器苗为试材,采用绿色荧光蛋白基因标记技术,研究了干旱胁迫(连续干旱0、7、14、21、28 d和35 d)对樱桃根际促生细菌YT3的标记菌YT3-gfp数量的影响,同时结合平板计数法和末端限制性片段长度多态性分析(terminal restriction fragment length polymorphism,T-RFLP)技术,研究了干旱对樱桃土壤中的微生物数量及细菌群落结构多样性影响。结果表明:樱桃根际土壤中的YT3-gfp数量是非根际土壤中的8.75—28.77倍,随着持续干旱强度的增加,YT3-gfp的数量先增加后减小。干旱对根际土壤中YT3-gfp数量的影响大于对非根际土壤的影响,分别在持续干旱至第21天和28天时,YT3-gfp的数量达到最大值。随着持续干旱强度的增加,根际土壤中细菌和放线菌数量先增加后减小,而真菌的数量一直减少。此外,持续干旱至第21天或28天时,樱桃根际土壤具有最高的丰富度指数、多样性指数和最低的优势度指数。基于T-RFLP的主成分分析结果显示持续干旱14—35 d时,其细菌群落结构成为一个相对独立的群,群落结构趋于多样性;而持续干旱7 d和42 d构成另外两个相对独立的群,群落结构趋于简单。以上分析可知,干旱对土壤微生物影响显著,一定强度的干旱可提高细菌和放线菌数量,提高细菌群落结构多样性,适当干旱对维持根际土壤细菌群落结构多样性是有益的。     

Experimental evidence that wildflower strips increase pollinator visits to crops

Wild bees provide a free and potentially diverse ecosystem service to farmers growing pollination‐dependent crops. While many crops benefit from insect pollination, soft fruit crops, including strawberries are highly dependent on this ecosystem service to produce viable fruit. However, as a result of intensive farming practices and declining pollinator populations, farmers are increasingly turning to commercially reared bees to ensure that crops are adequately pollinated throughout the season. Wildflower strips are a commonly used measure aimed at the conservation of wild pollinators. It has been suggested that commercial crops may also benefit from the presence of noncrop flowers; however, the efficacy and economic benefits of sowing flower strips for crops remain relatively unstudied. In a study system that utilizes both wild and commercial pollinators, we test whether wildflower strips increase the number of visits to adjacent commercial strawberry crops by pollinating insects. We quantified this by experimentally sowing wildflower strips approximately 20 meters away from the crop and recording the number of pollinator visits to crops with, and without, flower strips. Between June and August 2013, we walked 292 crop transects at six farms in Scotland, recording a total of 2826 pollinators. On average, the frequency of pollinator visits was 25% higher for crops with adjacent flower strips compared to those without, with a combination of wild and commercial bumblebees (Bombus spp.) accounting for 67% of all pollinators observed. This effect was independent of other confounding effects, such as the number of flowers on the crop, date, and temperature. Synthesis and applications. This study provides evidence that soft fruit farmers can increase the number of pollinators that visit their crops by sowing inexpensive flower seed mixes nearby. By investing in this management option, farmers have the potential to increase and sustain pollinator populations over time.     

Effects of flower size and number on pollinator visitation to wild radish,Raphanus raphanistrum

Plant traits that increase pollinator visitation should be under strong selection. However, few studies have demonstrated a causal link between natural variation in attractive traits and natural variation in visitation to whole plants. Here we examine the effects of flower number and size on visitation to wild radish by two taxa of pollinators over 3 years, using a combination of multiple regression and experimental reductions in both traits. We found strong, consistent evidence that increases in both flower number and size cause increased visitation by syrphid flies. The results for small bees were harder to interpret, because the multiple regression and experimental manipulation results did not agree. It is likely that increased flower size causes a weak increase in small-bee visitation, but strong relationships between flower number and small-bee visitation seen in 2 years of observational studies were not corroborated by experimental manipulation of this trait. Small bees may actually have responded to an unmeasured trait correlated with flower number, or lower small-bee abundances when the flower number manipulation was conducted may have reduced our ability to detect a causal relationship. We conclude that studies using only 1 year, one method, or measuring only one trait may not provide an adequate understanding of the effects of plant traits on pollinator attraction.     

Habitat fragmentation leads to reduced pollinator visitation,fruit production and recruitment in urban mangrove forests

Mangrove forests worldwide undergo anthropogenic fragmentation that may threaten their existence, and yet there have been few tests of the effects of fragmentation on demographic processes critical for mangrove regeneration. Predicting the effects of habitat fragmentation on mangroves is problematic as pollinators may move more freely across water than terrestrial habitat, and propagules can be widely dispersed by water. Here, within each of two estuaries, we compared pollinator diversity and activity, reproductive effort and output, and rates of recruitment for sets of three large (>1500 trees), medium (300–500) and small (<50) stands. As predicted, most measures of reproductive activity and success were inversely related to stand size with large stands typically producing significantly more and larger fruit, and significantly more seedlings. Most strikingly, we found the effect of fragmentation on the abundance of pollinators (honeybees), the production and quality of fruit and the survival rate of seedlings to be similar, showing significant reduction of recruitment in small stands. This study provides the first rigorous evidence that recruitment of mangroves, like for many terrestrial plants, is negatively impacted by habitat fragmentation. From a management perspective, we argue that in the short term our data imply the importance of conserving the largest possible stands. However, additional work is needed to determine (1) the proportion of recruits within small stands that originate within large stands, (2) how seedling performance varies with fruit size and genotype, and (3) how seedling size and performance vary with the abundance and diversity of pollen.     

Linkage disequilibrium in French wild cherry germplasm and worldwide sweet cherry germplasm

A basic knowledge on linkage disequilibrium (LD) is necessary in order to determine resolution of association studies. We investigated the extent and patterns of LD in a self-incompatible species (Prunus avium L.), in 3 groups (wild cherry, sweet cherry landraces and sweet cherry modern varieties), using a set of 35 microsatellite markers and the gametophytic self-incompatibility locus. Since population structure might create spurious LD, we thus used the information provided by a structure analysis published in a previous study to perform the LD analysis. In the current study, we detected a greater LD extent in sweet cherry than in wild cherry, which is plausibly due to the bottleneck associated with domestication and breeding. Higher LD values in sweet cherry sub-groups may be explained by smaller sample sizes. We also showed that the remaining structure in the groups of sweet cherry, in particular landraces, is responsible for a part of the LD extent. Intra-group relatedness may also account for extensive LD in two sub-groups. These results demonstrate, if ever necessary, the importance of controlling the genetic structure and relatedness when estimating LD. Moreover, LD decays very rapidly with genetic linkage distance in both wild and sweet cherries, which seems promising for future association studies.     

Floral visitation by the Argentine ant reduces pollinator visitation and seed set in the coast barrel cactus, Ferocactus viridescens

Mounting evidence indicates that trade-offs between plant defense and reproduction arise not only from resource allocation but also from interactions among mutualists. Indirect costs of plant defense by ants, for example, can outweigh benefits if ants deter pollinators. Plants can dissuade ants from occupying flowers, but such arrangements may break down when novel ant partners infiltrate mutualisms. Here, we examine how floral visitation by ants affects pollination services when the invasive Argentine ant (Linepithema humile) replaces a native ant species in a food-for-protection mutualism with the coast barrel cactus (Ferocactus viridescens), which, like certain other barrel cacti, produces extrafloral nectar. We compared the effects of floral visitation by the Argentine ant with those of the most prevalent native ant species (Crematogaster californica). Compared to C. californica, the Argentine ant was present in higher numbers in flowers. Cactus bees (Diadasia spp.), the key pollinators in this system, spent less time in flowers when cacti were occupied by the Argentine ant compared to when cacti were occupied by C. californica. Presumably as a consequence of decreased duration of floral visits by Diadasia, cacti occupied by L. humile set fewer seeds per fruit and produced fewer seeds overall compared to cacti occupied by C. californica. These data illustrate the importance of mutualist identity in cases where plants balance multiple mutualisms. Moreover, as habitats become increasingly infiltrated by introduced species, the loss of native mutualists and their replacement by non-native species may alter the shape of trade-offs between plant defense and reproduction.     

Congruence between visitation and pollen-transport networks in a California plant–pollinator community

Most recent studies describing pollination networks are based on observed flower visits, and few have explicitly tested if the floral visitors actually carry pollen. Since floral visitors can vary in their ability to remove and transfer pollen, it is important to show that visitation patterns reflect effective pollination. Given the difficulty of measuring per-visit pollen deposition at the community scale, a first step is to examine the amount of conspecific pollen carried by insect visitors. Here I compared the plant–animal visitation network with the pollen-transport network, estimated from insect pollen loads, for a montane meadow community from southern California, USA. Visitation and pollen-transport networks were positively associated with each other in both 2001 and 2002. However, the exclusion of visitors that do not carry any conspecific pollen reveals that pollen-transport networks are more specialized from the plants' perspective and that species are involved in fewer mutualistic interactions compared with estimates derived from visitation frequencies. Although conspecific pollen loads were smaller in 2002, bees tended to carry the largest conspecific loads in both years and were responsible for transporting the most pollen. These results suggest that, although visitation networks are suitable first-order approximations of pollination networks, information on which visitors carry conspecific pollen, and in what amounts, is crucial for distinguishing between antagonistic and mutualistic interactions.     

Influence of pollinator abundance and flower visitation on seed yield in red clover

Red clover (Trifolium pratense) is an important forage crop in temperate regions. However, the seed yield in red clover, especially in tetraploid cultivars, constrains the commercial success of this crop. The relationship between seed yield and several flower and inflorescence characteristics has been investigated previously, but up to now no clear link has been identified. The central questions in this article are as follows: (1) Whether nectar production and composition are correlated with seed yield? (2) Which is the main pollinator found on red clover in Flanders? (3) Whether pollinator preferences can explain differences between high and low seed-yielding cultivars and genotypes? We found that Bombus pascuorum was the most abundant pollinator on red clover in the region considered. Nectar volume, nectar composition or pollinator preference did not explain the variation in seed yield between genotypes, nor between diploid and tetraploid cultivars. To explain seed yield differences among red clover genotypes and ploidy levels better, we believe that plant-related characteristics like problems with meiosis, low pollen or seed viability or embryo abortion need to be investigated.