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Plants produce an array of defensive compounds with toxic or deterrent effects on insect herbivores. Pollen can contain relatively high concentrations of such defense compounds, but the causes and consequences of this enigmatic phenomenon remain mostly unknown. These compounds could potentially protect pollen against antagonists but could also reduce flower attractiveness to pollinators. We combined field observations of the pollen-rewarding Lupinus argenteus with chemical analysis and laboratory assays to test three hypotheses for the presence of pollen defense compounds: (1) these compounds are the result of spillover from adjacent tissues, (2) they protect against pollen thieves, and (3) they act as antimicrobial compounds. We also tested whether pollen defense compounds affect pollinator behavior. We found a positive relationship between alkaloid concentrations in pollen and petals, supporting the idea that pollen defense compounds partly originate from spillover. However, pollen and petals exhibited quantitatively (but not qualitatively) distinct alkaloid profiles, suggesting that plants can adjust pollen alkaloid composition independently from that of adjacent tissues. We found no relationship between pollen alkaloid concentration and the abundance of pollen thieves in Lupinus flowers. However, pollen alkaloids were negatively associated with bacterial abundance. Finally, plants with more alkaloids in their pollen received more pollinator visits, but these visits were shorter, resulting in no change in the overall number of flowers visited. We propose that pollen defense compounds are partly the result of spillover from other tissues, while they also play an antimicrobial role. The absence of negative effects of these compounds on pollinator visitation likely allows their maintenance in pollen at relatively high concentrations. Taken together, our results suggest that pollen alkaloids affect and are mediated by the interplay of multiple interactions., (© 2024 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Specialist insect herbivores make up a substantial fraction of Earth's biodiversity; however, they exploit a minority of plant lineages. For instance, in the eastern United States and Canada, ~25% of bee species are pollen specialists, but they are hosted by a small fraction of the native, animal-pollinated angiosperms in the region: Only 6% of plant genera and 3% of families support pollen-specialist bees. It is unclear why some plant lineages host specialist bees while others do not. We know that at least some specialist bees use plant taxa that are avoided by generalists, suggesting that specialist bees favor plants with low-quality pollen, potentially as a strategy to escape competition or obtain protection from natural enemies. There is also evidence that specialist bees prefer superabundant host plants. Here we investigate whether pollen quality and plant abundance predict patterns of host use by specialist bees in eastern North America. Through field observations, we find that plants hosting specialist bees are frequent sources of pollen for generalists, suggesting that their pollen is not generally avoided by bees due to poor pollen quality. In addition, our analysis of a large citizen-science data set shows that regional abundance strongly predicts which plant genera in the eastern United States host pollen-specialist bees. Our results show that bees specialize on regionally abundant-but not necessarily low-quality-plant lineages. These plant lineages may provide more opportunities for the evolution of specialists and lower likelihood of specialist extinction., (© 2023 The Ecological Society of America.)

Many bee species are dietary specialists and restrict their pollen foraging to a subset of the available flowers. However, the reasons for specialization-and the reasons certain plant taxa support numerous specialists-are often unclear. Many bees specialize on the plant family Asteraceae, despite evidence its pollen is a poor food for non-specialists. Here, we studied six mason bee (Osmia) species, including three Asteraceae specialists, to test whether observed pollen-usage patterns reflect larval nutritional requirements, to investigate what aspects of Asteraceae pollen make it unsuitable for non-specialists, and to understand how Asteraceae specialists tolerate their seemingly low-quality diet. We reared larval bees on host and nonhost pollen and found that Asteraceae specialists could develop on nonhost provisions, but that other bees could not survive on Asteraceae provisions. These effects did not seem related to nutritional deficiencies, since Asteraceae provisions were not amino acid deficient, and we found no consistent differences in digestive efficiency among pollen types. However, Asteraceae specialists completed more foraging flights per larva, generally collected relatively larger provisions, and produced more frass (waste) than the other species, suggesting quantitative compensation for low food quality. Toxins, deficiencies in unmeasured nutrients, or aspects of pollen grain structure might explain poor survival of non-specialists on Asteraceae provisions. Our results suggest that floral host selection by specialist bees is not related to optimizing larval nutrition. We recommend further investigation of host-selection behaviour in adult bees and of pollen digestion in larvae to better understand the evolution of bee-flower associations.

Pollen plays two important roles in angiosperm reproduction, serving as a vehicle for the plant's male gametes, but also, in many species, as a lure for pollen-feeding animals. Despite being an important food source for many pollinators, pollen often contains compounds with known deterrent or toxic properties, as documented in a growing number of studies. Here we review these studies and discuss the role of pollen defensive compounds in the coevolutionary relationship between plants and bees, the preeminent consumers of pollen. Next, we evaluate three hypotheses that may explain the existence of defensive compounds in pollen. The pleiotropy hypothesis, which proposes that defensive compounds in pollen merely reflect physiological spillover from other plant tissues, is contradicted by evidence from several species. Although plants may experience selection to defend pollen against poor-quality pollinators, we also find only partial support for the protection-against-pollen-collection-hypothesis. Finally, pollen defences might protect pollen from colonisation by antagonistic microorganisms (antimicrobial hypothesis), although data to evaluate this idea are scarce. Further research on the effects of pollen defensive compounds on pollinators, pollen thieves, and pollen-colonising microbes will be needed to understand why many plants have chemically defended pollen, and the consequences of those defences for pollen consumers., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

Warm temperatures are required for insect flight. Consequently, warming could benefit many high-latitude and high-altitude insects by increasing opportunities for foraging or oviposition. However, warming can also alter species interactions, including interactions with natural enemies, making the net effect of rising temperatures on population growth rate difficult to predict. We investigated the temperature-dependence of nesting activity and lifetime reproductive output over 3 yr in subalpine populations of a pollen-specialist bee, Osmia iridis. Rates of nest provisioning increased with ambient temperatures and with availability of floral resources, as expected. However, warmer conditions did not increase lifetime reproductive output. Lifetime offspring production was best explained by rates of brood parasitism (by the wasp Sapyga), which increased with temperature. Direct observations of bee and parasite activity suggest that although activity of both species is favored by warmer temperatures, bees can be active at lower ambient temperatures, while wasps are active only at higher temperatures. Thus, direct benefits to the bees of warmer temperatures were nullified by indirect costs associated with increased parasite activity. To date, most studies of climate-change effects on pollinators have focused on changing interactions between pollinators and their floral host-plants (i.e., bottom-up processes). Our results suggest that natural enemies (i.e., top-down forces) can play a key role in pollinator population regulation and should not be overlooked in forecasts of pollinator responses to climate change., (© 2016 by the Ecological Society of America.)

Many specialist herbivores eat foods that are apparently low quality. The compensatory benefits of a poor diet may include protection from natural enemies. Several bee lineages specialize on pollen of the plant family Asteraceae, which is known to be a poor-quality food. Here we tested the hypothesis that specialization on Asteraceae pollen protects bees from parasitism. We compared rates of brood parasitism by Sapyga wasps on Asteraceae-specialist, Fabeae-specialist, and other species of Osmia bees in the field over several years and sites and found that Asteraceae-specialist species were parasitized significantly less frequently than other species. We then tested the effect of Asteraceae pollen on parasites by raising Sapyga larvae on three pollen mixtures: Asteraceae, Fabeae, and generalist (a mix of primarily non-Asteraceae pollens). Survival of parasite larvae was significantly reduced on Asteraceae provisions. Our results suggest that specialization on low-quality pollen may evolve because it helps protect bees from natural enemies.

Plants produce an array of chemical and mechanical defences that provide protection against many herbivores and pathogens. Putatively defensive compounds and structures can even occur in floral rewards: for example, the pollen of some plant taxa contains toxic compounds or possesses conspicuous spines. Yet little is known about whether pollen defences restrict host-plant use by bees. In other words, do bees, like other insect herbivores, tolerate the defences of their specific host plants while being harmed by non-host defences? To answer this question, we compared the effects of a chemical defence from Lupinus (Fabaceae) pollen and a putative mechanical defence (pollen spines) from Asteraceae pollen on larval survival of nine bee species in the tribe Osmiini (Megachilidae) varying in their pollen-host use. We found that both types of pollen defences reduce larval survival rate in some bee species. These detrimental effects were, however, mediated by host-plant associations, with bees being more tolerant of the pollen defences of their hosts, relative to the defences of plant taxa exploited by other species. This pattern strongly suggests that bees are adapted to the pollen defences of their hosts, and that host-plant use by bees is constrained by their ability to tolerate such defences. [ABSTRACT FROM AUTHOR]

Species exhibit a range of specialisation in diet and other niche axes, with specialists typically thought to be more efficient in resource use but more vulnerable to extinction than generalists. Among herbivorous insects, dietary specialists seem more likely to lack acceptable host plants during the insect's feeding stage, owing to fluctuations in host‐plant abundance or phenology. Like other herbivores, bee species vary in host breadth from pollen specialisation (oligolecty) to generalisation (polylecty).Several studies have shown greater interannual variation in flowering phenology for earlier‐flowering plants than later‐flowering plants, suggesting that early‐season bees may experience substantial year‐to‐year variation in the floral taxa available to them.It was therefore reasoned that, among bees, early phenology could be a more viable strategy for generalists, which can use resources from multiple floral taxa, than for specialists. Consequently, it was expected that the median dates of collection of adult specimens to be earlier for generalist species than for specialists. To test this, phenology data and pollen diet information on 67 North American species of the bee genus Osmia was obtained.Controlling for latitude and phylogeny, it was found that dietary generalisation is associated with significantly earlier phenology, with generalists active, on average, 11–14 days earlier than specialists.This result is consistent with the generalist strategy being more viable than the specialist strategy for species active in early spring, suggesting that dietary specialisation may constrain the evolution of bee phenology—or vice versa. [ABSTRACT FROM AUTHOR]

A correction is presented to article, "Pollen chemical and mechanical defences restrict host-plant use by bees," published in the previous issue of the periodical.

Abstract: Evolutionary ecologists seek to explain the processes that maintain variation within populations. In plants, petal color variation can affect pollinator visitation, environmental tolerance, and herbivore deterrence. Variation in sexual organs may similarly affect plant performance. Within‐population variation in pollen color, as occurs in the eastern North American spring ephemeral Erythronium americanum, provides an excellent opportunity to investigate the maintenance of variation in this trait. Although the red/yellow pollen‐color polymorphism of E. americanum is widely recognized, it has been poorly documented. Our goals were thus (1) to determine the geographic distribution of the color morphs and (2) to test the effects of pollen color on components of pollen performance. Data provided by citizen scientists indicated that populations range from monomorphic red, to polymorphic, to monomorphic yellow, but there was no detectable geographic pattern in morph distribution, suggesting morph occurrence cannot be explained by a broad‐scale ecological cline. In field experiments, we found no effect of pollen color on the probability of predation by the pollen‐feeding beetle Asclera ruficollis, on the ability of pollen to tolerate UV‐B radiation, or on siring success (as measured by the fruit set of hand‐pollinated flowers). Pollinators, however, exhibited site‐specific pollen‐color preferences, suggesting they may act as agents of selection on this trait, and, depending on the constancy of their preferences, could contribute to the maintenance of variation. Collectively, our results eliminate some hypothesized ecological effects of pollen color in E. americanum, and identify effects of pollen color on pollinator attraction as a promising direction for future investigation. [ABSTRACT FROM AUTHOR]

Bilateral symmetry has evolved from radial symmetry in several floral lineages, and multiple hypotheses have been proposed to account for the success of this floral plan. One of these hypotheses posits that bilateral symmetry (or, more generally, a reduced number of planes of floral symmetry) allows for more precise pollen placement on pollinators. Greater precision would maximize the efficacy of pollen transfer to conspecifics, while minimizing reproductive interference amongst plant species. Despite the intuitiveness of this hypothesis, it has little experimental support. Here, we tested whether a reduction in the number of floral planes of symmetry (as in the transition from radial to bilateral symmetry) increases the potential precision of pollen placement. We analyzed video recordings of bumblebees (Bombus impatiens ) visiting artificial flowers to determine whether consistency in flower entry angle differed between radial (round) and disymmetric (rectangular) "flowers". We observed more consistent entry angles for disymmetric flowers than for radial flowers, with entry angles to radial flowers 43% more variable on average (standard deviations of 30° vs. 21°). Bees trained on flowers with an intermediate (square) morphology exhibited a slight, non-significant preference for radial symmetry over disymmetry. Our results show that disymmetry--an evolutionarily intermediate form of floral symmetry--has the potential to increase pollen transfer to conspecific stigmas, relative to radial symmetry. Thus, evolutionary reduction in the number of planes of floral symmetry likely provides benefits in terms of pollen delivery, as suggested by the pollen-placement-accuracy hypothesis. These findings offer insight into the evolution of floral symmetry. [ABSTRACT FROM AUTHOR]

Early-flowering species may be especially susceptible to occasional pollen limitation and, therefore, may benefit from a mixed-mating strategy that provides reproductive assurance. We studied cleistogamous (CL) and chasmogamous (CH) fruit set of spring-flowering Viola praemorsa Dougl. ex Lindl. along an elevational gradient in the Rocky Mountains, testing whether pollen limitation or allocation to CL reproduction covaried with timing of flowering onset, within and across sites. Contrary to predictions, we found no pollen limitation of reproduction at any site, and variation among sites in the pattern of allocation to cleistogamy was not related to growing season length. Differences in reproductive strategy between early- and late-flowering plants within sites were attributable to differences in plant size, with relative allocation to cleistogamy increasing with size. This pattern has been found in some other cleistogamous species, and may indicate a cost of large CH floral displays, perhaps associated with geitonogamy or herbivory. We found no experimental evidence for resource reallocation in response to CH reproductive output, although a weak negative relationship between CH and CL fruit set across a larger sample of unmanipulated plants suggests such a trade-off. The significance of cleistogamy may be clarified by studying how pollinator visitation, self-pollination, and herbivore damage vary temporally and with floral display size. [ABSTRACT FROM AUTHOR]