Support for the predictions of the pollinator-mediated stabilizing selection hypothesis

Aims Floral traits are frequently used in traditional plant systematics because of their assumed constancy. One potential reason for the apparent constancy of flower size is that effective pollen transfer between flowers depends on the accuracy of the physical fit between the flower and pollinator. Therefore, flowers are likely to be under stronger stabilizing selection for uniform size than vegetative plant parts. Moreover, as predicted by the pollinator-mediated stabilizing selection (PMSS) hypothesis, an accurate fit between flowers and their pollinators is likely to be more important for specialized pollination systems as found in many species with bilaterally symmetric (zygomorphic) flowers than for species with radially symmetric (actinomorphic) flowers.Methods In a comparative study of 15 zygomorphic and 13 actinomorphic species in Switzerland, we tested whether variation in flower size, among and within individuals, is smaller than variation in leaf size and whether variation in flower size is smaller in zygomorphic compared to actinomorphic species.Important findings Indeed, variation in leaf length was significantly larger than variation in flower length and width. Within-individual variation in flower and leaf sizes did not differ significantly between zygomorphic and actinomorphic species. In line with the predictions of the PMSS, among-individual variation in flower length and flower width was significantly smaller for zygomorphic species than for actinomorphic species, while the two groups did not differ in leaf length variation. This suggests that plants with zygomorphic flowers have undergone stronger selection for uniform flowers than plants with actinomorphic flowers. This supports that the relative uniformity of flowers compared to vegetative structures within species, as already observed in traditional plant systematics, is, at least in part, a consequence of the requirement for effective pollination.     

Exploring the 'Most Effective Pollinator Principle' with Complex Flowers: Bumblebees and Ipomopsis aggregata

The ‘most effective pollinator principle’ impliesthat floral characteristics often reflect adaptation to thepollinator that transfers the most pollen, through a combinationof high rate of visitation to flowers and effective depositionof pollen during each visit. We looked for the expected positivecorrelation between quantity and quality of visits in Ipomopsisaggregata, whose red, tubular flowers are considered to be adaptedto hummingbirds. Hummingbirds were indeed the most common floralvisitors in 5 years of observation. However, long-tongued bumblebeesdeposited on average three-times as much outcross pollen pervisit to virgin flowers, and elicited four-times as much seedproduction, as did hummingbirds. Hence visitors that are relativelyinfrequent, and unexpected given the ‘pollination syndrome’of the plant, can be surprisingly good pollinators. One interpretationof this observation is that natural selection favours a specializedfloral morphology that excludes all but a single type of visitor,but that there are constraints on achieving this outcome. Analternative is that selection favours some degree of floralgeneralization, but that flowers can retain features that adaptthem to a particular type of pollinator in spite of this generalization.Copyright 2001 Annals of Botany Company Bumblebees, floral evolution, hummingbirds, Ipomopsis aggregata, pollination syndromes, pollinator effectiveness, scarlet gilia, seed set, specialization, temporal variation     

大叶铁线莲访花昆虫调查及盗蜜昆虫行为研究

大叶铁线莲Clematis heracleifolia DC. 的花朵大多下垂,需要传粉昆虫为其传粉,目前尚无关于其访花昆虫研究的报道。 通过2年的野外观察研究,共观察到27种昆虫访问大叶铁线莲。 发现有盗蜜行为的昆虫7种, 其中1种同时具有初级盗蜜和次级盗蜜行为,2种具有初级盗蜜行为,4种具有次级盗蜜行为;黄胸木蜂Xylocopa appendiculata Smith是主要的盗蜜昆虫,其盗蜜行为影响了其它昆虫的访花行为,对大叶铁线莲的传粉造成一定的影响。 在其余20种访花昆虫中,双带弄蝶Lobocla bifasciata (Bremer et Grey)、贡尺蛾Gonodontis aurata Prout、熊蜂Bombus sp.和姬蜂虻Systropus sp.是优势种; 而小青花金龟Oxycetonia jucunda Faldermann和日本条螽Ducetia japonica (Thunberg)访花频率最低,且访花目的只是取食花朵。 通过对大叶铁线莲访花昆虫的调查和盗蜜昆虫行为的研究,为大叶铁线莲的传粉生物学和保护提供参考依据。     

The influence of floral symmetry,dependence on pollinators and pollination generalization on flower size variation

Background and Aims

The pollinator-mediated stabilizing selection hypothesis suggests that the specialized pollination system of zygomorphic flowers might cause stabilizing selection, reducing their flower size variation compared with actinomorphic flowers. However, the degree of ecological generalization and of dependence on pollinators varies greatly among species of both flower symmetry types and this may also affect flower size variation.

Methods

Data on 43 species from two contrasting communities (one alpine and one lowland community) were used to test the relationships and interactions between flower size phenotypic variation, floral symmetry, ecological pollination generalization and species'' dependence on pollinators.

Key Results

Contrary to what was expected, higher flower size variation was found in zygomorphic than in actinomorphic species in the lowland community, and no difference in flower size variation was found between symmetry types in the alpine community. The relationship between floral symmetry and flower size variation depended on ecological generalization and species'' dependence on pollinators, although the influence of ecological generalization was only detected in the alpine community. Zygomorphic species that were highly dependent on pollinators and that were ecologically specialized were less variable in flower size than ecologically generalist and selfing zygomorphic species, supporting the pollinator-mediated stabilizing selection hypothesis. However, these relationships were not found in actinomorphic species, probably because they are not dependent on any particular pollinator for efficient pollination and therefore their flower size always shows moderate levels of variation.

Conclusions

The study suggests that the relationship between flower size variation and floral symmetry may be influenced by population-dependent factors, such as ecological generalization and species'' dependence on pollinators.     

Do flowers wave to attract pollinators? A case study with Silene maritima

To answer the question whether flowers wave to attract pollinators, we determine: (1) the heritability of floral mobility; (2) whether wavy flowers attract more insects; (3) does the duration of pollination affect seed set; and (4) the relationship between seed set and floral mobility. The pollination ecology of Silene maritima was investigated. Flowers on stalks of different waviness were used to investigate the effect of floral movement on pollinator visits. There is heritable variation in both direct and indirect estimates of floral mobility. The highest insect total visitation times were associated with medium length thin stalks that were visited more frequently and by more insect species. Although mean individual visit durations were less than those of less mobile flowers, this was compensated for by increased visits. Observations of controlled pollinations show that when the visit times are low, so is seed set and therefore low and high mobility flowers might suffer from reduced fitness. Combining these observations provides a mechanism that could be driving stabilizing selection for flower stalk traits, with a trade-off applying between waving to attract pollinators and not being too mobile as to prevent effective pollination. Further evidence for stabilizing selection is provided by the relationship observed in the field between seed set and floral mobility where the highest levels of fitness was associated with intermediate levels of floral waving.     

Is plasticity across seasons adaptive in the annual cleistogamous plant Lamium amplexicaule?

Background and aims Many angiosperms exhibit cleistogamy, the production of both cleistogamous flowers (CL), which remain closed and obligately self-pollinated, and chasmogamous flowers (CH), which are potentially open-pollinated. The CH proportion can be plastic. Plasticity is adaptive if environmental changes can be reliably assessed and responded to with an appropriate phenotype and if plastic genotypes have higher fitness in variable environments than non-plastic ones.Methods We studied the plastic response of four natural populations from northern and southern France of an annual cleistogamous plant, Lamium amplexicaule, to predictable seasonal variation. Plants were grown in a semi-controlled environment in spring and in autumn. We assessed the variation in flower number, phenology and cleistogamy-related traits, which were all plastic with respect to season. The CH proportion was higher in spring than in autumn in all four populations.Key Results We showed significant stabilizing selection for cleistogamy traits, with higher optimal CH proportions and more pronounced stabilizing selection in spring than in autumn. Observed CH proportions were close to the predicted optimal CH proportions in each season except in autumn for southern populations, which do not experience the autumnal growing season in nature.Conclusions These results are consistent with adaptive plasticity across seasons of cleistogamy in L. amplexicaule. We propose that adaptive plasticity of cleistogamy could be driven by pollination environment variation, with CL flowers providing reproductive assurance when pollinators are scarce and CH flowers reducing the inbreeding depression in offspring when pollinators are abundant.     

Variation in the intensity of pollination inDrosera tracyi: Selection is strongest when resources are intermediate

Summary I propose that the strength of selection is greatest when resource levels are intermediate. This idea was partially tested using artificial character variation for petal size inDrosera tracyi. This plant opens one flower each morning, with stigmas becoming available and pollen being presented simultaneously throughout the population. Bees remove all the pollen that can be removed and deposit all the pollen that can be deposited in one or two visits. Visitation rates track blooming density, as do pollen removal and deposition. I measured selection as the difference in pollen removal and deposition between flowers with trimmed petals versus flowers with normal petals. In one meadow, I found a wide range of levels of pollination and corresponding levels of selection, and selection was strongest at intermediate levels of pollination. This might mean that organisms are adapted to conditions when resources are at intermediate levels even though they may experience those conditions relatively rarely.     

Within-carpel and among-carpel competition during seed development, and selection on carpel number, in the apocarpous perennial herb Helleborus foetidus L. (Ranunculaceae)

Apocarpous flowers share opportunities for post-fertilization ovule selection among more functional levels than syncarpous flowers, because the occurrence of a variable number of unfused carpels adds a new source of variation to the likelihood of successful female reproduction. The extent to which post-fertilization events might differ among these unfused carpels may promote variations in the reproductive strategies of plants. We report a study of the variation, within and among carpels and flowers, in seed production and mass in the apocarpous Helleborus foetidus (Ranunculaceae), in relation to the number of carpels per flower. Differences within and among carpels in female reproductive success were affected by carpel number and pollination environment. When analysing whole flowers as functional units we also found that the magnitude of the differences related to carpel number and pollination treatment actually depended on the “distribution” of pollen types within flowers. Thus, variable within-flower pollination environments, more likely to occur in apocarpous than in syncarpous flowers, may affect the strategies of resource allocation for fruit development at different stages of the reproductive process. Regarding seed production, we found that producing more flowers with four carpels was under directional; however, when mean diaspore mass was considered as a measure of fitness, directional selection was found on producing flowers with two and three carpels (the modal carpel number found in wild populations). We discuss ecological and developmental reasons which could explain the observed pattern, and conclude that selection on an optimum carpel number may be very variable across the species range, as the discussed reasons may impose constraints on eventual evolutionary response, thus contributing to the maintenance of the intra-individual variability in carpel number.     

Are deception-pollinated species more variable than those offering a reward?

In most pollination systems, animals transfer pollen among plants of a given species. Pollinator visitations do not come without cost, so plants usually offer a reward. However, the flowers of some plant species, mostly orchids, lack rewards and deceive animals into visiting their flowers. Deceptive species are thought to have high levels of variation in traits associated with advertisement and pollinator attraction, which have been attributed to genetic drift, or disruptive selection due to pollinator behavior. Rewarding species are assumed to have less variation due to stabilizing selection. We compared variability in floral morphology and fragrance composition between deceptive and rewarding species. Because both suites of traits are often linked with floral advertisement and pollinator attraction, we expected variation to be greater in species with deceptive pollination systems than in those offering rewards. We obtained floral morphology metrics for 20 deceptive species and 41 rewarding species native or naturalized in Puerto Rico, Venezuela, and Ecuador. Floral fragrances were sampled from eight deceptive species and four rewarding species. We found that the amplitude of variation in floral morphology and fragrance composition covaries significantly. Comparison of coefficients of variation for morphology indicated that, overall, deceptive species show significantly higher variation than rewarding species, and this pattern was also found among just orchids or just nonorchids. There were no statistical differences in morphological variation between orchids and nonorchids within a functional pollination group. Fragrance variation, measured by Jaccard distance, tended to be greater for deceptive species than for rewarding species. Although overlap in measures of variation occurs between the two groups, the data support the hypothesis that populations of deception-pollinated species are more variable than rewarding species in traits associated with pollinator attraction.     

Floral specialization for different pollinators and divergent use of the same pollinator among co‐occurring Impatiens species (Balsaminaceae) from Southeast Asia

Floral variation among closely related species is thought to often reflect differences in pollination systems. Flowers of the large genus Impatiens are characterized by extensive variation in colour, shape and size and in anther and stigma positioning, but studies of their pollination ecology are scarce and most lack a comparative context. Consequently, the function of floral diversity in Impatiens remains enigmatic. This study documents floral variation and pollination of seven co‐occurring Impatiens spp. in the Southeast Asian diversity hotspot. To assess whether floral trait variation reflects specialization for different pollination systems, we tested whether species depend on pollinators for reproduction, identified animals that visit flowers, determined whether these visitors play a role in pollination and quantified and compared key floral traits, including floral dimensions and nectar characteristics. Experimental exclusion of insects decreased fruit and seed set significantly for all species except I. muscicola, which also received almost no visits from animals. Most species received visits from several animals, including bees, birds, butterflies and hawkmoths, only a subset of which were effective pollinators. Impatiens psittacina, I. kerriae, I. racemosa and I. daraneenae were pollinated by bees, primarily Bombus haemorrhoidalis. Impatiens chiangdaoensis and I. santisukii had bimodal pollination systems which combined bee and lepidopteran pollination. Floral traits differed significantly among species with different pollination systems. Autogamous flowers were small and spurless, and did not produce nectar; bee‐pollinated flowers had short spurs and large floral chambers with a wide entrance; and bimodally bee‐ and lepidopteran‐pollinated species had long spurs and a small floral chamber with a narrow entrance. Nectar‐producing species with different pollination systems did not differ in nectar volume and sugar concentration. Despite the high frequency of bee pollination in co‐occurring species, individuals with a morphology suggestive of hybrid origin were rare. Variation in floral architecture, including various forms of corolla asymmetry, facilitates distinct, species‐specific pollen‐placement on visiting bees. Our results show that floral morphological diversity among Impatiens spp. is associated with both differences in functional pollinator groups and divergent use of the same pollinator. Non‐homologous mechanisms of floral asymmetry are consistent with repeated independent evolution, suggesting that competitive interactions among species with the same pollination system have been an important driver of floral variation among Impatiens spp.     

Indirect costs counteract the effects of pollinator-mediated phenotypic selection on corolla size in the Mediterranean shrub Halimium atriplicifolium

Aims Larger corollas receive more pollinator visits but involve higher production and maintenance costs, especially under hot and dry conditions. This can result in indirect costs on reproductive output, which may counteract the effects of pollinator-mediated phenotypic selection on corolla size. In this study, I explored the relationship between corolla size and indirect costs and whether these costs counteract the effects of pollinator-mediated phenotypic selection on this trait in the Mediterranean shrub Halimium atriplicifolium. I hypothesized that (i) corolla production entails direct costs in dry mass, N and P, (ii) corollas entail significant indirect costs in terms of fruit and seed production, (iii) indirect costs increase with corolla size, (iv) this species may suffer pollen limitation to a certain degree and (v) indirect costs counteract the effects of pollinator-mediated selection on corolla size.Methods I compared fruit set and seed production of petal-removed flowers (R flowers) and unmanipulated control flowers (C flowers) and evaluated the influence of individual mean corolla size on relative fruit and seed gain of R compared to C flowers. I also estimated phenotypic selection on corolla size mediated by indirect costs and the combined effect of costs and pollinators (i.e. total selection).Important findings Corollas allocated sizeable amount of resources in terms of dry mass relative to the other floral structures. Fruit set and seed per fruit were significantly higher in R flowers, while individual mean corolla size showed a positive relationship with relative fruit gain. Phenotypic selection analysis revealed cost-mediated negative directional selection and absence of positive directional total selection on corolla size through fruit set. This translated into stabilizing total selection. These results suggest that Mediterranean environments can impose constraints on corolla size, counteracting advantages of larger corollas from the pollination point of view with increased indirect costs of such flowers.     

Strong pollinator‐mediated selection for increased flower brightness and contrast in a deceptive orchid

Contrasting flower color patterns that putatively attract or direct pollinators toward a reward are common among angiosperms. In the deceptive orchid Anacamptis morio, the lower petal, which makes up most of the floral display, has a light central patch with dark markings. Within populations, there is pronounced variation in petal brightness, patch size, amount of dark markings, and contrast between patch and petal margin. We tested whether pollinators mediate selection on these color traits and on morphology (plant height, number of flowers, corolla size, spur length), and whether selection is consistent with facilitated or negative frequency‐dependent pollination. Pollinators mediated strong selection for increased petal brightness (Δβ_poll = 0.42) and contrast (Δβ_poll = 0.51). Pollinators also tended to mediate stabilizing selection on brightness (Δγ_poll = –0.27, n.s.) favoring the most common phenotype in the population. Selection for reduced petal brightness among hand‐pollinated plants indicated a fitness cost associated with brightness. The results demonstrate that flower color traits influence pollination success and seed production in A. morio, indicating that they affect attractiveness to pollinators, efficiency of pollen transfer, or both. The documented selection is consistent with facilitated pollination and selection for color convergence toward cooccurring rewarding species.     

Insect Choice and Floral Size Dimorphism: Sexual Selection or Natural Selection?

In considerations of sexual floral size dimorphism, there is a conflict between sexual selection theory, which predicts that larger floral displays attract more pollinators, and optimality theory—particularly the ideal free distribution—which predict that pollinators' visits should match nutritional rewards. As an alternate explanation of this dimorphism, Müller reported that pollinators tend to visit larger male flowers before visiting smaller female flowers, thereby promoting effective pollination. To investigate optimality predictions, I offered pollinators a choice between smaller, less numerous, but more rewarding flowers; and larger, more numerous, but less rewarding flowers. Foragers initially favored the larger and more numerous flowers, but rapidly shifted preferences to conform with the predictions of the ideal free distribution. To test Müller's hypothesis, I offered pollinators choices between larger and smaller corollas of equal caloric reward. Results showed that although pollinators tended to visit larger corollas first, they did not visit them more often. These experiments highlight the need for further investigation into the tradeoff between natural and sexual selection, and their respective influences in pollination ecology.     

A new pollination system: brood-site pollination by flower bugs in Macaranga (Euphorbiaceae)

Background and Aims Macaranga: (Euphorbiaceae) is a large genus of dioecious trees with approx.260 species. To date, only one pollination study of the genushas reported brood-site pollination by thrips in M. hullettii.In this study, the pollination system of Macaranga tanariusis reported. Methods: The study was conducted on Okinawa and Amami Islands, Japan.Flower visitors on M. tanarius were collected and their pollenload and behaviour on the flowers examined, as well as inflorescencestructure and reward for the pollinators. Key Results: The most abundant flower visitors found on the male and femaleinflorescences were Orius atratus (Anthocoridae, Hemiptera),followed by Decomioides schneirlai (Miridae, Hemiptera). Pollenload on O. atratus from flowering pistillate inflorescenceswas detected as well as from staminate flowers. Orius atratusand D. schneirlai are likely to use the enclosed chambers formedby floral bracts as breeding sites before and during floweranthesis, and feed on nectar on the adaxial surface of flowerbracts. The extrafloral nectary has a ball-shaped structureand the contained nectar is not exposed; the hemipterans piercethe ball to suck out the nectar. Conclusions: The results indicate that the plant is pollinated by flowerbugs breeding on the inflorescences. This study may be the firstreport of pollination systems in which flower bugs are the mainpollinators. Similarity of pollination systems between M. hullettiiand M. tanarius indicates that the two brood-site pollinationsystems have the same origin. The pollinator species belongsto a predacious group, whose major prey includes thrips. Thepollination system might represent a unique example of evolutionfrom predatory flower visitors feeding on the pollinators (thrips)to the main pollinators.     

Prolongation of Embryo Sac Viability in Pear (Pyrus communis) Following Pollination or Treatment with Gibberellic Acid

Embryo sac development has been investigated in unpollinated,cross pollinated and gibberellic acid (GA₂) treated flowersof Pyrus communis L. While pollination and GA₃ treatments donot alter embryo sac development, they prolong embryo sac viability.In untreated unpollinated flowers, ovules degenerate between12 and 21 d after anthesis, while in cross pollinated and GA₃treated flowers this degeneration is postponed by about 10 d.Thus, in a cross pollinated flower this extends the period overwhich a successful fertilization can take place. This increasedperiod of viability is accompanied by an elongation of the embryosac itself. Elongation takes place two weeks prior to fertilizationin cross pollinated flowers. The extension of life span of embryo sacs following pollinationand treatment with gibberellic acid indicates that a stimulusinduced by ‘pollination’ could be mediated by GA₃Whatever its mechanism of operation, the prolongation of embryosac viability by pollination represents a selective advantage,in that the period at which the ovules are receptive to fertilizationmust be significantly extended. Embryo sac, gibberellic acid, Pyrus communis, pear, pollination     

Variation in style length and anther–stigma distance in Ixiolirion songaricum (Amaryllidaceae)

Variation in style length and anther–stigma distance (ASD) was investigated in 18 populations of Ixiolirion songaricum, an early-spring ephemeral perennial herb in northern Xinjiang. The effect of ASD on autonomous self-pollen deposition and seed set was assessed using bagging experiments. Seed production under autonomous self-pollination, cross-pollination and natural pollination was determined by manual pollination experiments. Stigmas of multiple flowers within an individual were sandwiched between two separate anther levels, and no individuals possessed approach or reverse herkogamy. Style length varied continuously among flowers, resulting in a wide range of variation in ASD. There were fewer flowers with a small ASD than with a large ASD in populations. ASD was negatively correlated with autonomous self-pollen deposition and seed set, but it was positively correlated with natural pollination seed set. Seed production under natural pollination was higher than that under autonomous self-pollination, but it was smaller than that under cross-pollination. Variation in style length and ASD among flowers was not caused by style or stamen elongation in I. songaricum, which differ from other species reported in literature that have continuous variation in style length and ASD.     

Nectar Sugar Composition in Relation to Pollination Syndromes in Sinningieae (Gesneriaceae)

A putative correlation between nectar sugar composition andpollination syndrome was evaluated in the tribe Sinningieae(Neotropical Gesneriaceae). Sucrose, fructose and glucose werequantified in the nectar of 45 species using high performanceanion-exchange chromatography. Representative species of thehummingbird, bee, bat and moth pollination syndromes were sampledin relation to their numeric importance in the tribe. In hummingbirdand bee flowers, which represent 95% of the species in Sinningieae,nectar was sucrose-dominant (ratio [sucrose]/[hexose] > 1).Sugar ratios below one were only found in the nectar of threespecies with moth and bat syndromes. Sugar concentration averaged23.9 ± 10.6% (wt/total wt) in hummingbird flowers and28.7 ± 10.6% in bee flowers, whereas diluted nectar (7.1± 3.4%) was restricted to bat flowers. Similarities inthe nectar of hummingbird and bee flowers contrast with thepresence of specific morphological traits associated with thesetwo syndromes, indicating that plant-pollinator relationshipsrely on flower display rather than on nectar characteristics.By contrast, distinct nectar chemistry is correlated with thebat syndrome in which a particularly low sucrose productionis responsible for hexose dominance. Copyright 2001 Annals ofBotany Company Nectar sugar composition, pollination syndrome, Sinningia, Gesneriaceae, Brazil     

POLLINATORS DISCRIMINATE AMONG FLORAL HEIGHTS OF A SEXUALLY DECEPTIVE ORCHID: IMPLICATIONS FOR SELECTION

Pollinators have influenced the evolution of many morphological floral traits, although few studies have shown that pollinators have influenced plant height. Chiloglottis trilabra is one of many Australian orchids that deceive and attract male pollinators by mimicking the sex pheromones and morphology of females insects. Orchids in this genus have unusually short flowers whose peduncle elongates dramatically after pollination to approximately twice the original height. In a series of choice experiments in the field, we show that pollinators of C. trilabra strongly discriminate among floral heights, preferring flowers presented at 15 cm-20 cm over flowers presented at lower and higher positions (ranging from 2 cm-100 cm). Our results suggested pollinators have the potential to mediate stabilizing selection for floral height when pollination is limiting. However, the natural height range of the orchid (mean = 10 cm, range 5 cm–15 cm) was lower than the experimentally determined optimum for visitation frequency. This difference may indicate that pollinator-mediated selection does not occur in this species, perhaps because seed set is not sufficiently limited. Alternatively, other life-history factors may counteract pollinator-mediated selection, yielding an evolutionary compromise in height.     

Effects of Temperature, Light, Nutrients and Carbon Dioxide on the Strength of the Self-Incompatibility Response in Detached Flowers of Lycopersicon peruvianum

The effects of temperature, light, nutrients and CO₂on the gametophyticself-incompatibility response in a clone of Lycopersicon peruvianumhave been quantified using fluorescence microscopy of stylesquashes prepared from detached flowers held under experimentalconditions for 48 h after self pollination. Self-incompatibilitywas significantly weakened by raising the temperature from 15°C to 22 or 25°C, and by incubating flowers withoutan energy source - i.e. in the dark and/or without externallysupplied sucrose. The sucrose effect was not duplicated by equimolarmannitol or a full mineral nutrient solution plus vitamins.CO₂ applied at 5% in air for 8 h after pollination had no detectableeffect on pollen tube arrest. Observed weakening effects werenot sufficient for use in production of selfed seed. They indicate,however, that self-incompatibility in this clone of L. peruvianumis a temperature-sensitive, energy-dependent process, and supportthe oppositional model of self-incompatibility in which incompatiblepollen tubes are actively inhibited. Lycopersicon, Solanaceae, carbon dioxide, light, nutrients, self-incompatibility, temperature     

小峰熊蜂访花偏爱性

传粉昆虫在访花时,通常会表现出对某一类型花的偏爱性。本研究利用人工制作的大小、颜色、形态和气味不同的9种类型的花来研究小峰熊蜂Bombus hypocrita的访花偏爱性。结果表明：当增加花的大小、形态、气味等附加特征数时,小峰熊蜂的偏爱性程度与花朵附加特征数有显著相关性（P<0.01）。当花朵颜色由2种增加到4种,熊蜂对紫色花的偏爱性程度降低,但花朵颜色的种类与小峰熊蜂的访花偏爱性没有相关性（P>0.05）,花颜色的种类对熊蜂访紫色花的偏爱性影响不大。大小为5 cm的紫花被访次数（108±9次）明显高于大小为3 cm的紫花被访次数（40±4次）（P<0.01）,说明熊蜂明显喜欢访大花瓣的紫花。完全盛开的紫花被访次数（129±13次）显著高于刚绽放的花被访次数（26±3次）（P<0.01）,说明熊蜂喜欢访盛开的紫花。柠檬味的紫花被访次数（63±8次）明显低于草莓味的紫色花被访次数（88±2次）（P<0.05）,说明熊蜂喜欢访草莓味的花朵。