Reconnecting plants and pollinators: challenges in the restoration of pollination mutualisms

Ecological restoration of plant-pollinator interactions has received surprisingly little attention, despite animal-mediated pollination underpinning reproduction of the majority of higher plants. Here, we offer a conceptual and practical framework for the ecological restoration of pollination mutualisms. Through the use of targeted restoration plantings to attract and sustain pollinators and increased knowledge of the ecological requirements of pollinators, we propose that pollination could be successfully restored in degraded ecosystems. The challenge for pollination biologists is to integrate their findings with those of plant restoration ecologists to ensure sustainable pollination in restored ecosystems.     

The secret pollinators: an overview of moth pollination with a focus on Europe and North America

Pollination is a crucial plant–animal interaction in ecosystems, and moths (Lepidoptera) are a widespread and species-rich group of flower visitors. In this article, plant and moth species connected via pollination interactions were identified from the literature, and information on the relevance of moth pollination in various ecosystems, including agro-ecosystems, was compiled, particularly for Europe and North America. Overall, 227 moth–flower pollination interactions were found, including certain specialized relationships between plants and pollinating seed predators. Most of the interactions could be attributed to the moth families Noctuidae (90 interactions, 56 species) and Sphingidae (85 interactions, 32 species), and to the plant families Orchidaceae (109 interactions, 22 species) and Caryophyllaceae (59 interactions, 16 species). Limited information is available on the role of moth pollination in natural ecosystems (7 studies). In temperate agro-ecosystems, moths are most likely not essential to crop pollination, but they can contribute to the pollination of non-crop plants, which are crucial to maintaining biodiversity in these ecosystems. In general, the role of moths as pollinators appears to be underestimated because only a few studies on moth pollination are available, and long-term, ecosystem-scale research is necessary to address temporal fluctuations in their abundance and community composition.     

When bigger is not better: intraspecific competition for pollination increases with population size in invasive milkweeds

One of the essential requirements for an introduced plant species to become invasive is an ability to reproduce outside the native range, particularly when initial populations are small. If a reproductive Allee effect is operating, plants in small populations will have reduced reproductive success relative to plants in larger populations. Alternatively, if plants in small populations experience less competition for pollination than those in large populations, they may actually have higher levels of reproductive success than plants in large populations. To resolve this uncertainty, we investigated how the per capita fecundity of plants was affected by population size in three invasive milkweed species. Field surveys of seed production in natural populations of different sizes but similar densities were conducted for three pollinator-dependent invasive species, namely Asclepias curassavica, Gomphocarpus fruticosus and G. physocarpus. Additionally, supplemental hand-pollinations were performed in small and large populations in order to determine whether reproductive output was limited by pollinator activity in these populations. Reproductive Allee effects were not detected in any of the study species. Instead, plants in small populations exhibited remarkably high levels of reproductive output compared to those in large populations. Increased fruit production following supplemental hand-pollinations suggested that the lower reproductive output of naturally pollinated plants in large populations is a consequence of pollen limitation rather than limitation due to abiotic resources. This is consistent with increased intraspecific competition for pollination amongst plants in large populations. It is likely that the invasion of these milkweed species in Australia has been enhanced because plants in small founding populations experience less intraspecific competition for pollinators than those in large populations, and thus have the ability to produce copious amounts of seeds.     

Secondary production and diet of an invasive snail in freshwater wetlands: implications for resource utilization and competition

Invasive species can monopolize resources and thus dominate ecosystem production. In this study we estimated secondary production and diet of four populations of Pomacea canaliculata, a freshwater invasive snail, in wetlands (abandoned paddy, oxbow pond, drainage channel, and river meander) in monsoonal Hong Kong (lat. 22°N). Apple snail secondary production (ash-free dry mass [AFDM]) ranged from 165.9 to 233.3 g m⁻² year⁻¹, and varied between seasons. Production was lower during the cool dry northeast monsoon, when water temperatures might have limited growth, but fast growth and recruitment of multiple cohorts were possible throughout much (7–10 months) of the year and especially during the warm, wet southwest monsoon. The diet, as revealed by stomach-content analysis, consisted mainly of detritus and macrophytes, and was broadly consistent among habitats despite considerable variation in the composition and cover of aquatic plants. Apple snail annual production was >10 times greater than production estimates for other benthic macroinvertebrates in Hong Kong (range 0.004–15 g AFDM m⁻² year⁻¹, n = 29). Furthermore, annual production estimates for three apple snail populations (i.e. >230 g AFDM m⁻² year⁻¹) were greater than published estimates for any other freshwater snails (range 0.002–194 g AFDM m⁻² year⁻¹, n = 33), regardless of climatic regime or habitat type. High production by P. canaliculata in Hong Kong was attributable to the topical climate (annual mean ~24°C), permitting rapid growth and repeated reproduction, together with dietary flexibility including an ability to consume a range of macrophytes. If invasive P. canaliculata can monopolize food resources, its high productivity indicates potential for competition with other macroinvertebrate primary consumers. Manipulative experiments will be needed to quantify these impacts on biodiversity and ecosystem function in wetlands, combined with management strategies to prevent further range extension by P. canaliculata.     

Optimal cell size for resource uptake in fluids: a new facet of resource competition

Planktonic microorganisms are affected by various size-dependent processes both from the bottom up and from the top down. We developed a simple resource-consumer model to explore how size-dependent resource uptake and resource loss influence the growth of, and competition between, planktonic microorganisms. We considered three steps of resource uptake: diffusive transport of resource molecules, uptake by membrane transporters, and cellular enzymatic catalysis, and we investigated optimal cell size when one, two, or three of those steps limit resource uptake. Optimal cell size depends negatively on the size of resource molecules when resource uptake is limited by diffusive transport and membrane uptake. When competing for two resources of different molecular sizes, two different-sized consumers can coexist if the inputs of resources and sizes of consumers are correctly chosen. The model suggests that mixtures of various-sized resources can promote coexistence and size diversity of microorganisms even if the availability of one element, such as carbon, nitrogen, or phosphorus, limits the whole community. Model predictions include that bacteria grown on maltose or polysaccharides should be smaller compared with those grown on glucose under carbon limitation. Our results suggest that size of resource molecules can be an important factor in microbial resource competition in aquatic environments.     

谁是腊梅的主要传粉者:昆虫行为与传粉作用

访花昆虫不同的访花与飞行行为导致其访花频率小同,进而对植物传粉产生不同的影响.意大利蜜蜂(Apis mellifera ligustica)、苍蝇和食蚜蝇是腊梅(Chimonanthus praecox)最常见的访花类群(或种类),但是它们的访花行为却有很大不同.意大利蜜蜂主要以快速飞行为主,偶见爬行,苍蝇访花时主要以快速飞行、爬行和跳跃为主,而食蚜蝇以长时间单花访问和悬空飞行以及间歇性休息为主.根据访花类群(或种类)在一朵花上的访花时间和花间飞行时间进行推算,每只意大利蜜蜂、苍蝇和食蚜蝇每分钟分别可以访问4.57、2.65和0.53朵花.结合每种(类)昆虫的访花数量推算出意大利蜜蜂、苍蝇和食蚜蝇每分钟分别可以访问498.19、1,089.74和99.78朵花.传粉效力(相同条件下单位时间内相同数目访花者能够授粉的花朵数)实验结果证明苍蝇和意大利蜜蜂分别是93％、100％(n=30),而食蚜蝇只有13％(n=30).苍蝇和意大利蜜蜂可携带大量花粉且具有很高的传粉效力,因此,访花速度的快慢是二者访花能力强弱的主要限制因素.由此,我们认为苍蝇可能是腊梅最主要的传粉者,意大利蜜蜂次之(数量偏少),食蚜蝇再次(访花与飞行行为的影响以及访花效力较低是限制其访花能力的主要因素).     

Fruit size decline from the margin to the center of capitula is the result of resource competition and architectural constraints

Plants produce repeated structures, such as leaves, flowers, and fruits, which differ in size and shape. One example of this is fruit size, which is commonly observed to decrease from proximal to distal positions within an inflorescence. The resource limitation hypothesis proposes that because proximal fruits usually develop first, they have temporal priority on access to resources over distal fruits. The non-mutually exclusive architectural effects hypothesis suggests that these position effects in fruit size may also be due to inherent architectural variation along infructescence axes. We separated out the effects of resource competition and inflorescence architecture by removing the outer or the inner flowers within capitula of Tragopogon porrifolius. We also studied if fruit position influenced germination and seedling performance in order to assess fitness consequences of position effects. Inner fruits were significantly heavier when outer flowers were removed. However, outer fruits did not significantly increase when inner flowers were removed, suggesting later fruits were limited by the development of early fruits. Our findings also suggest that architectural constraints restricted the size of inner fruits in comparison with outer ones. We found that both resource competition and inflorescence architecture affected the fruit size of T. porrifolius, even though this species does not have linear, indeterminate inflorescences. We advance the hypothesis that, when such effects on fitness occur, resource competition-mediated position effects could turn, in evolutionary time, into architectural position effects.     

Beyond the pollination syndrome: nectar ecology and the role of diurnal and nocturnal pollinators in the reproductive success of Inga sessilis (Fabaceae)

Inga species present brush‐type flower morphology allowing them to be visited by distinct groups of pollinators. Nectar features in relation to the main pollinators have seldom been studied in this genus. To test the hypothesis of floral adaptation to both diurnal and nocturnal pollinators, we studied the pollination ecology of Inga sessilis, with emphasis on the nectar secretion patterns, effects of sequential removals on nectar production, sugar composition and the role of diurnal and nocturnal pollinators in its reproductive success. Inga sessilis is self‐incompatible and pollinated by hummingbirds, hawkmoths and bats. Fruit set under natural conditions is very low despite the fact that most stigmas receive polyads with sufficient pollen to fertilise all ovules in a flower. Nectar secretion starts in the bud stage and flowers continually secreting nectar for a period of 8 h. Flowers actively reabsorbed the nectar a few hours before senescence. Sugar production increased after nectar removal, especially when flowers were drained during the night. Nectar sugar composition changed over flower life span, from sucrose‐dominant (just after flower opening, when hummingbirds were the main visitors) to hexose‐rich (throughout the night, when bats and hawkmoths were the main visitors). Diurnal pollinators contributed less than nocturnal ones to fruit production, but the former were more constant and reliable visitors through time. Our results indicate I. sessilis has floral adaptations, beyond the morphology, that encompass both diurnal and nocturnal pollinator requirements, suggesting a complementary and mixed pollination system.     

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

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

How to be an attractive male: floral dimorphism and attractiveness to pollinators in a dioecious plant

Background  

Sexual selection theory predicts that males are limited in their reproductive success by access to mates, whereas females are more limited by resources. In animal-pollinated plants, attraction of pollinators and successful pollination is crucial for reproductive success. In dioecious plant species, males should thus be selected to increase their attractiveness to pollinators by investing more than females in floral traits that enhance pollinator visitation. We tested the prediction of higher attractiveness of male flowers in the dioecious, moth-pollinated herb Silene latifolia, by investigating floral signals (floral display and fragrance) and conducting behavioral experiments with the pollinator-moth, Hadena bicruris.     

Density dynamics of bumblebees in subalpine meadows: competition and resource limitation

Densities of Bombus flavifrons and B. rufocinctus were measured over one summer in 23 and 11 discrete subalpine meadows, respectively. I assessed the relation between observed bee densities and meadow size, elevation, floristics, temperature, and time of day by multiple regression. Multivariate regression solutions for each week of observation accounted for 38 to 61% of the variation in density for B. flavifrons and 47 to 87% for B. rufocinctus . Temperature or time of day consistently accounted for the most variation in density of the two species. In early summer meadow size or elevation also accounted for a large proportion of explained variation in B. flavifrons density; as the summer progressed meadow floristics increasingly contributed to the explanation of variation in both species.
In mid- to late summer a statistically significant portion of the variation in densities not explained by habitat characteristics was accounted for by densities of congeners: i.e., densities of B. flavifrons and B. rufocinctus were negatively related in eight meadows where they co-occurred. In experimental meadows where all Bombus but one of the two target species were removed. B. rufocinctus underwent significant positive density compensation but B. flavifrons did not. To the extent that B. flavifrons appeared to limit the density of B. rufocinctus these statistical and experimental results are consistent with the hypothesis that competition limits local bumblebee densities.     

Outline of HIV replication and its cellular factors: the track of an invader in the cell

A number of novel findings with reference to HIV replication have been reported even though it passed more than 20 years after a first HIV isolation. Although many cellular factors are known to be involved in the HIV replication, recently investigators discovered novel HIV-suppressive cellular factors such as APOBEC or TRIM5 alpha. Here, I describe and discuss how HIV uses the cellular machinery for its replication.     

The effects of competition and the presence of an audience on weight lifting performance

The purpose of this investigation was to examine the effects of the presence of an audience and competition on maximal weight lifting performance. Thirty-two recreationally trained participants (15 men, 17 women; 21 +/- 2.5 years) performed a 1 repetition maximum (1 RM) bench press during 3 different situations (coaction, competitive coaction, and audience condition). Subjects also completed the Activation-Deactivation Adjective Checklist Short Form following the 3 trials to measure arousal state during each of the 3 trials. Significant differences (p < 0.05) were found between competitive coaction and coaction trials as well as between audience and coaction trials. Both men and women demonstrated the highest performance in front of an audience (105 +/- 48 kg) followed by competition (103 +/- 46 kg) and then the coaction trial (93 +/- 43 kg). No significant difference in arousal was measured between trials. The data suggest that performing a maximal lift in the presence of an audience or in competition facilitates performance and support the self-presentation and self-awareness theories. Social facilitation effects should be controlled in research settings and may aid the performance of weight lifting activities during events or competition.     

The consequences of direct versus indirect species interactions to selection on traits: pollination and nectar robbing in Ipomopsis aggregata

Organisms experience a complex suite of species interactions. Although the ecological consequences of direct versus indirect species interactions have received attention, their evolutionary implications are not well understood. I examined selection on floral traits through direct versus indirect pathways of species interactions using the plant Ipomopsis aggregata and its pollinators and nectar robber. Using path analysis and structural equation modeling, I tested competing hypotheses comparing the relative importance of direct (pollinator-mediated) versus indirect (robber-mediated) interactions to trait selection through female plant function in 2 years. The hypothesis that provided the best fit to the observed data included robbing and pollination, suggesting that both interactors are important in driving selection on some traits; however, the direction and intensity of selection through robbing versus pollination varied between years. I then increased my scope of inference by assessing traits and species interactions across more years. I found that the potential for temporal variation in the direction and intensity of selection was pronounced. Taken together, results suggest that assessing the broader context in which organisms evolve, including both direct and indirect interactions and across multiple years, can provide increased mechanistic understanding of the diversity of ways that animals shape floral and plant evolution.     

The sweet jelly of Combretum lanceolatum flowers (Combretaceae): a cornucopia resource for bird pollinators in the Pantanal, western Brazil

 The pollination biology of the neotropical scandent shrub Combretum lanceolatum was studied in the seasonally-flooded Pantanal region in western Brazil. This plant bears horizontally oriented inflorescences, whose yellowish green flowers begin to expand at dusk and are fully open at dawn. Instead of fluid nectar the flowers produce sweet gelatinous secretion in form of pellets. The glandular complex of the flower is composed of the inner wall of the receptacle and its tubular extension, being equivalent to the nectariferous disk of the nectar-producing species within the genus. The jelly is produced at night, contains mannan and is imbibed by free hexoses. It originates by swelling and disintegration of the inner wall, after contact with the nectar generated concomitantly in the mesophyll. Combretum lanceolatum is unique within the genus in its production of jelly pellets instead of liquid nectar. A new term, the jelly-flower, is proposed for flowers with this kind of reward. The pellet is not replaced once removed by a bird, and thus resembles a fruit in its availability to consumers, another unique feature that distinguishes this species within the genus. The jelly pellets offered by the many flowered branches attract a great diversity of bird visitors (28 species from eight families), which feed on this copious food resource and pollinate the flowers. The most effective pollinators probably are thrushes, tanagers, and orioles. Flocking parakeets and macaws sometimes feed on the petals, thus acting as flower plunderers. Combretum lanceolatum presents a high fruit set under natural conditions, which likely favours its spreading and becoming a weed species. Received July 11, 2000 Accepted November 18, 2000