Addition Of Pollen Research Articles

Mating system, outcrossing distance effects and pollen availability in the wind-pollinated treeline species Polylepis australis BITT. (Rosaceae)

Abstract Fragmentation may negatively affect plant fitness through pollen limitation and increased levels of inbreeding. Effects of fragmentation may vary with regard to life form and breeding system, and few studies exist for wind-pollinated trees. We examined the effects of hand-selfing, varying outcrossing distances and pollen addition on seed mass and germination rate of Polylepis australis BITT. (Rosaceae), a wind-pollinated treeline species endemic to Argentina. We also investigated pollen germination on the stigma and pollen tube growth to determine compatibility resulting from selfing and outcrossing. Selfing reduced seed germination rates with significant differences between open pollination and outcrosses at 30 km. In addition, we found a tendency for pollen germination and pollen tube growth to decrease following selfing. Between-fragment crosses resulted in a trend of higher reproductive output than within-fragment crosses, whereas values were similar between open pollination and between-fragment crosses. Pollen addition did not increase reproductive success neither in small nor in larger fragments. Our results suggest that highly isolated P. australis forests have a potential for inbreeding depression through selfing and within-fragment crosses. However, the results also indicate that pollen flow between P. australis forest fragments is still effective at the current fragmentation level, counteracting negative effects on seed quality resulting from reproductive isolation.

Florivory affects pollinator visitation and female fitness in Nemophila menziesii

While herbivory has traditionally been studied as damage to leaves, florivory - herbivory to flowers prior to seed set - can also have large effects on plant fitness. Florivory can decrease fitness directly, either through the destruction of gametes or through alterations to plant physiology during fruit set, and can also change the appearance of a flower, deterring pollinators and reducing seed set. In order to distinguish between these hypotheses, it is necessary to both damage flowers and add pollen in excess to study the effects of damage on pollen limitation. Very few studies have used this technique over the lifetime of a plant. Here I describe a series of experiments showing the effects of natural and artificial damage on reproductive success in the annual plant Nemophila menziesii (Hydrophyllaceae, sensu lato). I show that natural and artificial petal damage decreased radial symmetry relative to controls and that both types of damage deterred pollinator activity. Both naturally damaged flowers and artificially damaged flowers in the field set fewer fruit or seed relative to undamaged control flowers. Finally, in an experiment crossing artificial petal damage with pollen addition, petal damage alone over the lifetime of this plant decreased female fitness, but only after a threshold of damage was reached. The fitness effect appeared to be direct because there was no detectable effect of pollen addition on the relationship between florivory and fitness. This result implies that both damaged and undamaged plants show similar amounts of pollen limitation and suggests that pollinator-mediated effects contributed little to the negative effects of florivory on female fitness. Florivores may thus be an under-appreciated agent of selection in certain plants, although more experimental manipulation of florivory is needed to determine if it is important over a range of taxa.

Pollen limitation and natural selection on floral characters in the yellow monkeyflower, Mimulus guttatus

In flowering plants, pollen limitation has been proposed to intensify selection on floral characters important in pollinator attraction, but may also select for traits that increase seed set through autonomous selfing. Here, a factorial design (+/- pollen addition, +/- pollinator removal) was used to investigate how the pollination environment affects selection on floral morphology via female fitness in a mixed-mating population of the yellow monkeyflower, Mimulus guttatus (Phrymaceae). Female fitness was strongly pollen-limited, with supplementally pollinated plants setting 37% more seeds than open-pollinated individuals. Strong positive selection was found on flower length, weak positive selection on flower width : length ratio and no selection on stigma-anther distance in both open-pollinated and supplementally pollinated treatments. By contrast, flowers with relatively narrow corollas and low stigma-anther distances were favored in the pollinator exclusion treatment. These results provide mixed support for the idea that pollen limitation intensifies selection on floral characters. Despite strong phenotypic selection, natural pollen limitation did not mediate selection on characters associated with either pollinator attraction or self-fertilization. However, the novel pattern of selection on severely pollen-limited plants suggests that reproductive assurance against pollinator loss may have been directly involved in the floral evolution of closely related selfing taxa.

Reproductive biology of the invasive exotic shrub, Rhododendron ponticum L. (Ericaceae)

The reproductive biology of an exotic species will affect its ability to become naturalized and invasive in non-native habitats. Rhododendron ponticum is an ecologically damaging exotic weed in the British Isles, which spreads predominantly by seed. I investigated how inbreeding and outcrossing affect seed production and germination in a wild population of this species in Ireland. Experimental manipulations revealed low fruit and seed set when insects were excluded from flowers, suggesting that this species has limited capability for spontaneous autogamy. Hand-pollination treatments showed that, although flowers are self-compatible (with self and same plant pollen), higher levels of seed set occur following outcrossing (xenogamy). There was no significant difference in rate of germination of seeds from inbred or outcrossed treatments. The addition of xenogamous pollen to open flowers did not increase fruit or seed set, suggesting that flowers in this population are not pollen limited: native generalist pollinators, mainly bumblebees (Bombus spp.), are providing an adequate pollinator service. This work demonstrates that outcrossing increases seed set and pollinators are required to facilitate this. Hence, generalist native pollinators can promote invasion by exotic plants. Native pollinators can clearly play an important part in alien species invasion. © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 155, 373–381.

Is Floral Longevity Influenced by Reproductive Costs and Pollination Success in Cohniella ascendens (Orchidaceae)?

Although studies have shown that pollen addition and/or removal decreases floral longevity, less attention has been paid to the relationship between reproductive costs and floral longevity. In addition, the influence of reproductive costs on floral longevity responses to pollen addition and/or removal has not yet been evaluated. Here, the orchid Cohniella ascendens is used to answer the following questions. (a) Does experimental removal of flower buds in C. ascendens increase flower longevity? (b) Does pollen addition and/or removal decrease floral longevity, and does this response depend on plant reproductive resource status? To study the effect of reproductive costs on floral longevity 21 plants were selected from which we removed 50 % of the developing flower buds on a marked inflorescence. Another 21 plants were not manipulated (controls). One month later, one of four flowers on each marked inflorescence received one of the following pollen manipulation treatments: control, pollinia removal, pollination without pollinia removal or pollination with pollinia removal. The response variable measured was the number of days each flower remained open (i.e. longevity). The results showed significant flower bud removal and pollen manipulation effects on floral longevity; the interaction between these two factors was not significant. Flowers on inflorescences with previously removed flower buds remained open significantly longer than flowers on control inflorescences. On the other hand, pollinated flowers closed much faster than control and removed-pollinia flowers, the latter not closing significantly faster than control flowers, although this result was marginal. The results emphasize the strong relationship between floral longevity and pollination in orchids, as well as the influence of reproductive costs on the former.

Lipid‐enhanced pollen and lipid‐reduced flour diets and their effect on the longevity of honey bees (Apis mellifera L.)

Abstract As eucalypt pollens contain low concentrations of lipid, enhancing pollen diets with fatty acids was hypothesised to improve honey bee longevity. Different concentrations of linoleic and oleic acid added to eucalypt pollen were trialled in small cages containing approximately 1400 bees each. Bees fed diets of redgum (Corymbia calophylla (Lindl.) Hill & Johnson, formerly Eucalyptus calophylla) pollen had the lowest mortality of 22 diets tested for 6 weeks and had life spans (50%) greater than 42 days. Linoleic acid mixed with a redgum diet in concentrations >6% corresponded to life spans of 24–25 days. Bee longevity appeared to be more sensitive to oleic acid as life spans decreased to 15–21 days when diets had concentrations >2%. The life spans of bees fed soya bean flour were 26 days on low (0.6% lipid) fat, 19 days on defatted and 20 days on full‐fat diets. Bees fed lupin flour had a life span of 23 days. Adding redgum pollen to lupin flour caused increased mortality, but addition of pollen to soya bean flour was beneficial. Thus, beekeepers who choose to utilise soya bean or lupin flours as protein substitutes to pollen will have bees with reduced longevity. Bees fed redgum pollen that had been dried, crushed, irradiated and hermetically stored in a cool room for several years had similar longevity to bees fed fresh‐collected and frozen redgum pollen.

Plasticity and Timing of Flower Closure in Response to Pollination in Chamerion angustifolium (Onagraceae)

Flower longevity often exhibits a plastic response to pollination and may vary adaptively to maximize pollen removal and deposition and minimize flower maintenance costs. Using Chamerion angustifolium (fireweed), we experimentally tested two predictions that follow from this hypothesis: (1) flower closure should be triggered early, before fertilization, if it is to reduce costs and adjust to maximize pollen receipt; and (2) flower closure should respond differently to different kinds (outcross, self) and amounts (none, low, high) of pollen. In C. angustifolium, flower closure was initiated 4 h after pollination, well before any ovules were fertilized (25 h). The addition of pollen to stigmas significantly shortened the time to closure compared with unpollinated controls. Flower closure times were reduced with increasing pollen loads, up to 700 pollen grains, as might be expected if closure was adaptive. Flower closure was slower in self‐pollinations than outcross pollinations at 24 and 48 h after pollinat...

Does natural deposition of pine pollen affect the ovipositional behavior of Frankliniella occidentalis and Frankliniella fusca?

AbstractTree pollen, especially Pinus spp. (Pinaceae), is shed in large quantities every spring in North America. Pine pollen deposition onto leaves was found to significantly influence the ovipositional behaviors of certain thrips species (Thysanoptera: Thripidae) in peanut and tomato leaf choice and no‐choice tests. Pine pollen (Pinus elliottii Engelm.) increased the oviposition rate 2.9‐fold for Frankliniella occidentalis (Pergande) (western flower thrips) and 1.6‐fold for Frankliniella fusca (Hinds) (tobacco thrips) in choice tests averaged over both plant species. These results support the idea that pollen has a greater impact on F. occidentalis behavior than on F. fusca behavior. The most dramatic increase was in peanut, where F. occidentalis only oviposited on leaves dusted with pollen, suggesting that the addition of pollen stimulated this flower thrips to lay eggs on a poor host‐plant part. The impact of pollen on the rate of oviposition by thrips is important because it is the early‐instar nymphs that acquire tomato spotted wilt virus (TSWV), which these two thrips species vector. In a laboratory bioassay, the addition of pine pollen to TSWV‐infected peanut foliage increased the percentage of infected F. fusca after one generation.

Fruit abortion in the Chihuahuan-Desert endemic cactus Opuntia microdasys

Resource and pollen limitation, as well as pollen/ovule incompatibility, have been proposed as causes to explain fruit abortion. To assess whether abortion in Opuntia microdasys was due to resource and/or pollen limitation and could therefore be reversed fruit set and seed set were studied using controlled pollination experiments on 60 plants that had been randomly assigned a combination of watering and fertilization treatments. On the other hand, to test whether fruit abortion was irreversible, due to pollen/ovule incompatibility, we examined the reproductive biology of the species. This included observations on floral phenology, nectar production, flower visitors, numbers of pollen grains and ovules, and self-pollination experiments. Results showed that O. microdasys is a fully self-incompatible species and its floral biology and the activity of the main pollinator allow constant deposition of incompatible pollen onto stigmas, which may contribute to fruit abortion. Reproductive success was limited by nutrients and pollen, but the fruit set increased only by 58%, compared to 47% of the control, after the experimental addition of pollen, nutrients and water. The magnitude of pollen and resource limitation suggests that similar levels of abortion will be present in good as well as in bad years. Selfing as well as incompatibility between ramets from the same clone and between closely related plants seem plausible candidates to explain the large proportion of fruit abortion, and experimental cross pollination between genotypes identified through molecular markers are necessary to fully understand the considerable abortion rate that remains unexplained after pollen and resource addition. Interestingly, the possible reason why the abortion of energetically expensive fruits has not been eliminated by natural selection is that the aborted fruits are propagules able to root and produce new plants with the same genotype of the mother. Abortion would have a dramatic effect on cross-fertilized genotypes because they result in zero fitness, but it would have a positive effect on the fitness of the maternal genotype because a clonal offspring is produced. Evidently, the exact fitness consequences to the maternal plant will depend on the differences in survival and reproduction of these different offspring types.

EXPANDING THE LIMITS OF THE POLLEN-LIMITATION CONCEPT: EFFECTS OF POLLEN QUANTITY AND QUALITY

Pollination commonly limits seed production, as addition of pollen to stigmas often increases fecundity. This response is usually interpreted as evidence that plants' stigmas receive too few pollen grains to maximize ovule fertilization (quantity limitation); however, many genetic studies demonstrate that poor-quality pollen can also reduce seed production (quality limitation). We explore both aspects of pollen limitation theoretically with a dose-response model that incorporates a saturating negative-exponential relation of seed production to pollen receipt. This relation depends on aspects of ovule production, pollen import, pollen-pistil interactions and seed development, all of which can contribute to pollen limitation. Our model reveals that quantity limitation is restricted to the lowest range of pollen receipt, for which siring success per pollen grain is high, whereas quality limitation acts throughout the range of pollen receipt if plants do not import the highest-quality pollen. In addition to pollinator availability and efficiency, quantity limitation is governed by all post-pollination aspects of seed production. In contrast, quality limitation depends on the difference in survival of embryos sired by naturally delivered pollen vs. by pollen of maximal quality. We briefly illustrate the distinction between these two components of pollen limitation with results from the mistletoe Tristerix corymbosus. Our model also shows that the standard pollen-supplementation technique neither estimates the total intensity of pollen limitation nor distinguishes between its quantity and quality components. As an alternative, we propose a methodological protocol that requires both measurement of seed production following excess pollination with only outcross pollen and quantification of the dose-response relation of seed output to pollen receipt. This method estimates both the total extent of pollen limitation and its two components. Finally, we consider the influences on quantity and quality limitation, which reveals that quantity limitation probably occurs much less often than has been inferred from pollen-supplementation experiments. These interpretations suggest that an expanded perspective that recognizes the fecundity consequences of pollination with poor-quality pollen would promote ecological understanding of pollen limitation.

Pollen limitation meets resource allocation: towards a comprehensive methodology

The standard method of measuring pollen limitation is to add pollen to a number of flowers, preferably to a whole plant, and to compare fruit and seed set with that of naturally pollinated flowers on other plants. In 25 yr of research, this method has yielded valuable data, but it is difficult to use in large plants. This has caused a bias in the available data towards smaller, herbaceous plants with relatively few flowers. I argue that, in order to widen our knowledge of how pollen limitation affects plants, we should go beyond whole-plant pollen addition and change our concept of how a flowering plant functions. The traditional method does not take into account the variation in and dynamics of resource allocation and pollen availability. The concept of integrated physiological units (IPUs) does, but, although it has been applied to pollination biology, it has not received the attention it deserves. I use this article to present its merits again, to propose a step-by-step methodology for studying pollen limitation, and to examine factors influencing possible plant strategies.

Effect of Pine Pollen on Settling Behavior of <I>Frankliniella occidentalis</I> and <I>Frankliniella fusca</I> (Thysanoptera: Thripidae) on Tomato and Peanut

Pollen was found to significantly influence the host leaf selection and settling behaviors of certain thrips species (Thysanoptera: Thripidae). The effect of slash pine pollen (Pinus elliottii Engelm.) on the settling of Frankliniella occidentalis (Pergande) (western flower thrips) and Frankliniella fusca (Hinds) (tobacco thrips) was studied with leaf choice tests in peanut and tomato. In choice tests, F. occidentalis settling behavior was increased by seven-fold with the addition of pine pollen, but F. fusca was not significantly affected. In the flower thrips, the effect of pollen decreased over time without the renewal of pollen, suggesting thrips consumption of the pollen grains. These results support the idea that slash pine pollen has a greater impact on flower thrips (F. occidentalis) behavior than foliage thrips (F. fusca) behavior.

Temporal associations between daily counts of fungal spores and asthma exacerbations

Background: Outdoor aeroallergens are one of a number of environmental factors thought to precipitate asthma exacerbations. Aims: To investigate the short term associations between daily fungal spore concentrations and indicators...

Reproductive ecology of Thymelaea velutina (Thymelaeaceae)- Factors contributing to the maintenance of heterocarpy

Thymelaea velutina (Thymelaeaceae) is a dioecious shrub that presents a unique type of heterocarpy which consists of the simultaneous production of dry and fleshy fruits. It is endemic to the Balearic Islands (Western Mediterranean) and is found both in dunes and mountain areas. The goal of this study was to identify which factors influence the production of both fruit types, examining the variation of their effects at a spatio-temporal scale (comparing two localities in different years). Specifically, we investigated (1) whether pollen limitation influences the type of fruit produced, and (2) the possible differences in seed size, mass, dispersal capacity, seed predation, germination patterns and seedling survival between fruit types. We also examined if the production of fleshy fruits was modified with the application of gibberellins to reproductive branches. Although fleshy fruits were consistently more abundant than dry ones at both populations, their proportion was significantly higher at the site with greater precipitation. The addition of either pollen or gibberellins did not affect the proportion of each fruit type. Seeds in fleshy fruits are consistently larger, heavier and more likely to be dispersed than seeds in dry fruits, but germinability, germination rate and seedling survival was similar among fruit types. Heterocarpy in this species is currently maintained as there is no apparent factor that exerts any strong selective pressure on either fruit type. The two fruit types might even have different `functions', one serving especially for dispersal and population expansion and the other for producing a seed bank that ensures an eventual germination.

Biological control of cassava green mite with exotic and indigenous phytoseiid predators—Effects of intraguild predation and supplementary food

Abstract Intraguild predation, one aspect of predator–predator interaction, has received in recent years increasingly greater attention because of mounting evidence of the impact of predator–predator interactions on the structure of ecological communities and biological pest control. In laboratory experiments, we determined if intraguild predation occurs between Typhlodromalus manihoti and Euseius fustis , two phytoseiid mite predators of the cassava green mite Mononychellus tanajoa on cassava in Africa, and if the level of intraguild predation is affected by the availability of the primary prey, M. tanajoa , and maize pollen as alternative non-prey food. In the laboratory, the two predators attacked and killed heterospecific larvae but they did so only when prey mites and alternative food (maize pollen) were absent or scarce. On a diet of intraguild prey alone, the two predator species survived for several days but failed to reproduce. Addition of abundant levels of M. tanajoa and maize pollen increased survival and reproduction of the two predator species and reduced intraguild predation to very low levels. We then determined, on whole plants in pot experiments in a screenhouse, the effect of maize pollen (an alternative food) on the interactions between the two predator species and the impact of predator–predator interactions on suppression of M. tanajoa population densities. In single predator species treatments, both E. fustis and T. manihoti significantly reduced M. tanajoa densities; but surprisingly, E. fustis appeared to be superior to T. manihoti . The co-presence of the two predators on the same cassava plant resulted in similar suppression of M. tanajoa population densities, but greater suppression of M. tanajoa compared with T. manihoti alone, regardless of presence or absence of maize pollen. The presence of the two predator species together reduced their respective abundance compared with single predator species treatments. Addition of maize pollen, however, significantly increased densities of the generalist predator E. fustis , in both single and two predator species treatments; and tilted the balance of the interactions between the two predator species in favour of E. fustis . The findings of our research underscore the role of generalist predators like E. fustis in the suppression of M. tanajoa populations on cassava, and the potential role of alternative non-prey food in altering the interactions between co-occurring predators sharing the same primary prey (by favouring the generalist predators), and the effect of these interactions on suppression of population densities of the shared primary prey.

Style Persistence, Pollen Limitation, and Seed Set in the Common Prairie PlantEchinacea angustifolia(Asteraceae)

Pollen limitation of seed set in flowering plants has important ramifications for the population dynamics, evolution, and conservation of plant populations. I conducted a pollen addition and exclusion experiment demonstrating that style persistence signifies pollen limitation in the narrow-leaved purple coneflower Echinacea angustifolia, a species native to the North American prairie and plains. I developed a measure of style persistence, SP, a novel way to quantify pollen limitation in individual plants during the flowering season. Using this measure, I investigated the relationship between pollen limitation and seed set over two years in 19 and 27 natural remnant populations in an agricultural landscape. Population mean rates of seed set per plant varied from 0% to 54% in 1997 and from 0% to 63% in 1998. I found that pollen limitation reduced annual reproductive fitness within and among the populations studied. An analysis of the relationship between floret production and the rate of seed set provided no evidence that resource limitation influenced the rate of seed set. I estimated annual fecundity per plant as the product of the rate of seed set per floret, a pollen-limited process, and floret production per plant, likely a resource-limited process. Population means of individual annual fecundity ranged from 0 to 182 in 1997 and from 0 to 156 in 1998 and were predicted by population means of SP and the rate of seed set, but not by floret production. The effect of pollen limitation, as quantified by SP, overrides the strong, fundamental relationship between fecundity and floret production. This finding shows that populations consisting of large plants with large floral displays do not necessarily produce more seeds per plant.

Direct and indirect effects of pollinators and seed predators to selection on plant and floral traits

Although flowering traits are often assumed to be under strong selection by pollinators, significant variation in such traits remains the norm for most plant species. Thus, it is likely that the interactions among plants, mutualists, and other selective agents, such as antagonists, ultimately shape the evolution of floral and flowering traits. We examined the importance of pollination vs pre‐dispersal seed predation to selection on plant and floral characters via female plant‐reproductive success in Castilleja linariaefolia (Scrophulariaceae). C. linariaefolia is pollinated by hummingbirds and experiences high levels of pre‐dispersal seed predation by plume moth and fly larvae in the Rocky Mountains of Colorado, USA, where this work was conducted. We first examined whether female reproduction in C. linariaefolia was limited by pollination. Supplemental pollination only marginally increased components of female reproduction, likely because seed predation masked, in part, the beneficial effects of pollen addition. In unmanipulated populations, we measured calyx length, flower production, and plant height and used path analysis combined with structural equation modeling to quantify their importance to relative seed set through pathways involving pollination vs seed predation. We found that the strength of selection on calyx length, flower production, and plant height was greater for seed predation pathways than for pollination pathways, and one character, calyx length, experienced opposing selection via pollination vs seed predation. These results suggest that the remarkable intraspecific variation in plant and floral characters exhibited by some flowering plants is likely the result of selection driven, at least in part, by pollinators in concert with antagonists, such as pre‐dispersal seed predators. This work highlights the subtle but complex interactions that shape floral and vegetative design in natural ecosystems.

Floral morph composition and pollen limitation in the seed set of Nymphoides indica populations

We investigated the floral morph composition and seed set of the populations in a heterostylous aquatic plant, Nymphoides indica O. Kuntze (Menyanthaceae). The bias of floral morph ratio in the populations was negatively correlated with the seed set. In the populations with biased floral morph ratio, the pollen on the stigma of the common morphs were dominated by pollen of the same morph, resulting in reduced seed set. The addition of pollen of the opposite morph on the stigmas of the common morph resulted in an increase of fruit and seed set rates. The intermorph pollination rate and the fruit and seed set rates of Nymphoides indica were highest when two floral morphs were close in the populations. Various insect species including Lepidoptera, Diptera and Hymenoptera visited the flowers. However, geitonogamous pollination was common because of the short flying distance of the pollinators. These results indicate that the seed set of Nymphoides indica is limited by the shortage of compatible pollens and that the success of intermorph pollination is influenced by spatial distribution of the two floral morphs. As Nymphoides indica is endangered, conservation of the populations with high fecundity is proposed by us.

Life history characteristics of Frankliniella occidentalis on cucumber leaves with and without supplemental food

AbstractThe development time, fecundity, longevity, and resultant intrinsic growth rate of the western flower thrips Frankliniella occidentalis (Pergande) [Thysanoptera: Thripidae] encaged on a cucumber leaf were compared among seven types of food supplied: six pollen species and a mixture of milk powder and yeast. The rationale was to find a food source that offers the least benefit for thrips and could therefore be considered as a food source for the preventative introduction of thrips predators. With the exception of the mixture of milk powder and yeast, all the food sources tested offered a nutritional benefit for the thrips. The addition of pollen increased fecundity and reduced development time, mainly during the larval stage. Betula pubescens and Corylus avellana pollen also increased adult longevity. The nutritional benefit of Pinus sylvestris pollen was greater than that of the other five pollen species, as manifested by its significantly greater positive effect on fecundity. The other pollen species could not be ranked in terms of nutritional benefit to F. occidentalis. The negative effect of the milk powder plus yeast mix on the life‐table parameters of F. occidentalis probably only occurs in an encaged situation where thrips cannot escape from the unfavorable environment. The crude estimate of the intrinsic growth rate of F. occidentalis increased from 0.163 on the plain cucumber leaf to 0.240 when P. sylvestris pollen was added to the leaf. The differences in intrinsic growth rate mainly reflect the differences in fecundity among the food sources. Thus, the peak oviposition rate may be used as a measure of the nutritional benefit F. occidentalis can obtain by feeding on supplemental food sources. The positive effect of a supplemental food source on thrips does not necessarily mean it is unsuitable for the preventative introduction of thrips predators, because the supplemental food can also affect the population dynamics of the predator and the predator–prey interaction and, hence, the outcome of biological control.

HOW PLANTS BENEFIT FROM PROVIDING FOOD TO PREDATORS EVEN WHEN IT IS ALSO EDIBLE TO HERBIVORES

It is well established that plants provide alternative foods to predators of herbivorous arthropods. This provision may facilitate protection against herbivory. However, plants often cannot prevent other organisms from utilizing these foods as well. There are many examples of herbivorous arthropods that can feed on plant-provided foods such as extrafloral nectar and pollen. The question therefore arises whether individual plants still gain protection when not only the predators, but also the herbivores, can feed on these foods. We investigated this question using a mathematical model and experiments that assessed the impact of supplementary pollen on the dynamics of predatory mites (Iphiseius degenerans (Berlese)) and herbivorous thrips (Frankliniella occidentalis (Pergande)), two arthropods capable of using pollen for reproduction. Replicated greenhouse experiments showed that addition of pollen every two weeks to one young mature leaf of a male-sterile cucumber plant increased predator population growth...