Amy L. Parachnowitsch

Covariation and phenotypic integration in chemical communication displays: biosynthetic constraints and eco-evolutionary implications

Chemical communication is ubiquitous. The identification of conserved structural elements in visual and acoustic communication is well established, but comparable information on chemical communication displays (CCDs) is lacking. We assessed the phenotypic integration of CCDs in a meta-analysis to characterize patterns of covariation in CCDs and identified functional or biosynthetically constrained modules. Poorly integrated plant CCDs (i.e. low covariation between scent compounds) support the notion that plants often utilize one or few key compounds to repel antagonists or to attract pollinators and enemies of herbivores. Animal CCDs (mostly insect pheromones) were usually more integrated than those of plants (i.e. stronger covariation), suggesting that animals communicate via fixed proportions among compounds. Both plant and animal CCDs were composed of modules, which are groups of strongly covarying compounds. Biosynthetic similarity of compounds revealed biosynthetic constraints in the covariation patterns of plant CCDs. We provide a novel perspective on chemical communication and a basis for future investigations on structural properties of CCDs. This will facilitate identifying modules and biosynthetic constraints that may affect the outcome of selection and thus provide a predictive framework for evolutionary trajectories of CCDs in plants and animals.

Spatiotemporal floral scent variation of Penstemon digitalis

Variability in floral volatile emissions can occur temporally through floral development, during diel cycles, as well as spatially within a flower. These spatiotemporal patterns are hypothesized to provide additional information to floral visitors, but they are rarely measured, and their attendant hypotheses are even more rarely tested. In Penstemon digitalis, a plant whose floral scent has been shown to be under strong phenotypic selection for seed fitness, we investigated spatiotemporal variation in floral scent by using dynamic headspace collection, respectively solid-phase microextraction, and analyzed the volatile samples by combined gas chromatography–mass spectrometry. Total volatile emission was greatest during flowering and peak pollinator activity hours, suggesting its importance in mediating ecological interactions. We also detected tissue and reward-specific compounds, consistent with the hypothesis that complexity in floral scent composition reflects several ecological functions. In particular, we found tissue-specific scents for the stigma, stamens, and staminode (a modified sterile stamen common to all Penstemons). Our findings emphasize the dynamic nature of floral scents and highlight a need for greater understanding of ecological and physiological mechanisms driving spatiotemporal patterns in scent production.

Natural selection on floral volatile production in Penstemon digitalis: highlighting the role of linalool.

Natural selection is thought to have shaped the evolution of floral scent; however, unlike other floral characters, we have a rudimentary knowledge of how phenotypic selection acts on scent. We found that floral scent was under stronger selection than corolla traits such as flower size and flower color in weakly scented Penstemon digitalis. Our results suggest that to understand evolution in floral phenotypes, including scent in floral selection, studies are crucial. For P. digitalis, linalool was the direct target of selection in the scent bouquet. Therefore, we determined the enantiomeric configuration of linalool because interacting insects may perceive the enantiomers differentially. We found that P. digitalis produces only (S)-(+)-linalool and, more interestingly, it is also taken up into the nectar. Because the nectar is scented and flavored with (S)-(+)-linalool, it may be an important cue for pollinators visiting P. digitalis flowers.

Direct and pollinator-mediated effects of herbivory on strawberry and the potential for improved resistance

The global decline in pollinators has partly been blamed on pesticides, leading some to propose pesticide-free farming as an option to improve pollination. However, herbivores are likely to be more prevalent in pesticide-free environments, requiring knowledge of their effects on pollinators, and alternative crop protection strategies to mitigate any potential pollination reduction. Strawberry leaf beetles (SLB) Galerucella spp. are important strawberry pests in Northern Europe and Russia. Given that SLB attack both leaf and flower tissue, we hypothesized pollinators would discriminate against SLB-damaged strawberry plants (Fragaria vesca, cultivar ‘Rügen’), leading to lower pollination success and yield. In addition we screened the most common commercial cultivar ‘Rügen’ and wild Swedish F. vesca genotypes for SLB resistance to assess the potential for inverse breeding to restore high SLB resistance in cultivated strawberry. Behavioral observations in a controlled experiment revealed that the local pollinator fauna avoided strawberry flowers with SLB-damaged petals. Low pollination, in turn, resulted in smaller more deformed fruits. Furthermore, SLB-damaged flowers produced smaller fruits even when they were hand pollinated, showing herbivore damage also had direct effects on yield, independent of indirect effects on pollination. We found variable resistance in wild woodland strawberry to SLB and more resistant plant genotypes than the cultivar ‘Rügen’ were identified. Efficient integrated pest management strategies should be employed to mitigate both direct and indirect effects of herbivory for cultivated strawberry, including high intrinsic plant resistance.

Adapting with the enemy : local adaptation in plant-herbivore interactions

Local adaptation by natural selection is a fundamental process in population differentiation and speciation. To determine if populations are adapted to local conditions, researchers use reciprocal transplant experiments: individuals are moved among populations to compare their performance in familiar (local) and foreign (nonlocal) conditions. These experiments are meant to evaluate whether adaption to one environment comes at a cost (via fitness trade-off) to performing well in another (nonlocal) environment (Kawecki & Ebert, 2004). Multiple meta-analyses of these experiments confirm that local adaptation can be a common phenomenon (e.g. Lajeunesse & Forbes, 2002; Leimu & Fischer, 2008; Hoeksema & Forde, 2008; Hereford, 2009), but individual experiments often cannot distinguish which aspect of the environment (abiotic or biotic) populations are adapted to. In research exploring local adaptation to biotic factors, such as antagonist interactions between plants and herbivores, coevolutionary theory plays an important role in forming predictions of when local adaptation should be observed (e.g. Gandon, 2002). One prediction is that populations will vary in who is ahead in the co-evolutionary ‘arms race’, and that this race is the primary driver of local adaptation. However, until Garrido et al. (pp. 445–453) in this issue of New Phytologist, no study had simultaneously examined local adaption in both host plants and their herbivores, while controlling for the potential abiotic factors that mediate local adaptation.

Do Plants Eavesdrop on Floral Scent Signals

Plants emit a diverse array of volatile organic compounds that can function as cues to other plants. Plants can use volatiles emitted by neighbors to gain information about their environment, and respond by adjusting their phenotype. Less is known about whether the many different volatile signals that plants emit are all equally likely to function as cues to other plants. We review evidence for the function of floral volatile signals and conclude that plants are as likely to perceive and respond to floral volatiles as to other, better-studied volatiles. We propose that eavesdropping on floral volatile cues is particularly likely to be adaptive because plants can respond to these cues by adjusting traits that directly affect pollination and mating.

Female Salix viminalis are more severely infected by Melampsora spp. but neither sex experiences associational effects

Abstract Associational effects of plant genotype or species on plant biotic interactions are common, not least for disease spread, but associational effects of plant sex on interactions have largely been ignored. Sex in dioecious plants can affect biotic interactions with herbivores and pollinators; however, its effects on plant–pathogen interactions are understudied and associational effects are unknown. In a replicated field experiment, we assessed Melampsora spp. leaf rust infection in monosexual and mixed sex plots of dioecious Salix viminalis L. to determine whether plant sex has either direct or associational effects on infection severity. We found no differences in Melampsora spp. infection severity among sexual monocultures and mixtures in our field experiment. However, female plants were overall more severely infected. In addition, we surveyed previous studies of infection in S. viminalis clones and reevaluated the studies after we assigned sex to the clones. We found that females were generally more severely infected, as in our field study. Similarly, in a survey of studies on sex‐biased infection in dioecious plants, we found more female‐biased infections in plant–pathogen pairs. We conclude that there was no evidence for associational plant sex effects of neighboring conspecifics for either females or males on infection severity. Instead, plant sex effects on infection act at an individual plant level. Our findings also suggest that female plants may in general be more severely affected by fungal pathogens than males.

Bacteria colonising Penstemon digitalis show volatile and tissue-specific responses to a natural concentration range of the floral volatile linalool

Bacteria on floral tissue can have negative effects by consuming resources and affecting nectar quality, which subsequently could reduce pollinator visitation and plant fitness. Plants however can employ chemical defences to reduce bacteria density. In North American, bee-pollinated Penstemon digitalis, the nectar volatile S-(+)-linalool can influence plant fitness, and terpenes such as linalool are known for their antimicrobial properties suggesting that it may also play a role in plant–microbe interactions. Therefore, we hypothesized linalool could affect bacterial growth on P. digitalis plants/flowers. Because P. digitalis emits linalool from nectar and nectary tissue but not petals, we hypothesised that the effects of linalool could depend on tissue of origin due to varying exposure. We isolated bacteria from nectary tissue, petals and leaves, and compared their growth relative to control using two volatile concentrations representing the natural emission range of linalool. To assess whether effects were specific to linalool, we compared results with the co-occurring nectar volatile, methyl nicotinate. We show that response to floral volatiles can be substance and tissue-origin specific. Because linalool could slow growth rate of bacteria across the P. digitalis phyllosphere, floral emission of linalool could play a role in mediating plant–bacteria interactions in this system.

New Synthesis: The Evolutionary Ecology of Floral Volatiles

Natural selection is a key process of population to species divergence and diversification. Since Lande and Arnold’s 1983 seminal paper “The measurement of selection on correlated characters”, hundreds of evolutionary biologists have measured phenotypic selection in wild populations (Kingsolver et al. 2012). These microevolutionary studies aim to understand the process of adaptive evolution, by regressing variation in putatively adaptive traits against measurements of fitness. The LandeArnold approach can compare selection strength, direction, and shape for different traits, making it a powerful tool for studying contemporary evolution. Although well over 50 studies have measured selection on floral characters, until recently none have incorporated chemical features of floral phenotypes, such as scent. Differing traditions may in part explain the scarcity of selection studies on floral scent; while evolutionary ecologists have focused on visual and mechanical aspects of floral display, floral scents have been more commonly studied in functional and ecological contexts. However, integration of floral characters into a unified perspective is critical to understand both function and evolution of floral diversity.

Bringing ecology blogging into the scientific fold: measuring reach and impact of science community blogs

The popularity of science blogging has increased in recent years, but the number of academic scientists who maintain regular blogs is limited. The role and impact of science communication blogs aimed at general audiences is often discussed, but the value of science community blogs aimed at the academic community has largely been overlooked. Here, we focus on our own experiences as bloggers to argue that science community blogs are valuable to the academic community. We use data from our own blogs (n = 7) to illustrate some of the factors influencing reach and impact of science community blogs. We then discuss the value of blogs as a standalone medium, where rapid communication of scholarly ideas, opinions and short observational notes can enhance scientific discourse, and discussion of personal experiences can provide indirect mentorship for junior researchers and scientists from underrepresented groups. Finally, we argue that science community blogs can be treated as a primary source and provide some key points to consider when citing blogs in peer-reviewed literature.