More than 87% of flowering plant species are animal-pollinated [1] and produce floral scents and other signals to attract pollinators. These floral cues may however also attract antagonistic visitors, including herbivores [2]. The dilemma is exacerbated when adult insects pollinate the same plant that their larvae consume. It remains largely unclear how plants maximize their fitness under these circumstances. Here we show that in the night-flowering wild tobacco Nicotiana attenuata, the emission of a sesquiterpene, (E)-α-bergamotene, in flowers increases adult Manduca sexta moth-mediated pollination success, while the same compound in leaves is known to mediate indirect defense against M. sexta larvae [3, 4]. Forward and reverse genetic analyses demonstrated that both herbivory-induced and floral (E)-α-bergamotene are regulated by the expression of a monoterpene-synthase-derived sesquiterpene synthase (NaTPS38). The expression pattern of NaTPS38 also accounts for variation in (E)-α-bergamotene emission among natural accessions. These results highlight that differential expression of a single gene that results in tissue-specific emission of one compound contributes to resolving the dilemma for plants when their pollinators are also herbivores. Furthermore, this study provides genetic evidence that pollinators and herbivores interactively shape the evolution of floral signals and plant defense.

Cost efficient foraging is of especial importance for animals like hawkmoths or hummingbirds that are feeding ‘on the wing', making their foraging energetically demanding. The economic decisions made by these animals have a strong influence on the plants they pollinate and floral volatiles are often guiding these decisions. Here we show that the hawkmoth Manduca sexta exhibits an innate preference for volatiles of those Nicotiana flowers, which match the length of the moth's proboscis. This preference becomes apparent already at the initial inflight encounter, with the odour plume. Free-flight respiration analyses combined with nectar calorimetry revealed a significant caloric gain per invested flight energy only for preferred—matching—flowers. Our data therefore support Darwin's initial hypothesis on the coevolution of flower length and moth proboscis. We demonstrate that this interaction is mediated by an adaptive and hardwired olfactory preference of the moth for flowers offering the highest net-energy reward.

Many plants emit diverse floral scents that mediate plant–environment interactions and attain reproductive success. However, how plants evolve novel and adaptive biosynthetic pathways for floral volatiles remains unclear. Here, we show that in the wild tobacco, Nicotiana attenuata, a dominant species-specific floral volatile (benzyl acetone, BA) that attracts pollinators and deters florivore is synthesized by phenylalanine ammonia-lyase 4 (NaPAL4), isoflavone reductase 3 (NaIFR3), and chalcone synthase 3 (NaCHAL3). Transient expression of NaFIR3 alone in N. attenuata leaves is sufficient and necessary for ectopic foliar BA emissions, and coexpressing NaIFR3 with NaPAL4 and NaCHAL3 increased the BA emission levels. Independent changes in transcription of NaPAL4 and NaCHAL3 contributed to intraspecific variations of floral BA emission. However, among species, the gain of expression of NaIFR3 resulted in the biosynthesis of BA, which was only found in N. attenuata. This study suggests that novel metabolic pathways associated with adaptation can arise via reconfigurations of gene expression.

No abstract

Benzyl acetone (4‐phenylbutan‐2‐one; BA), the dominant floral fragrance of the wild tobacco Nicotiana attenuata, is known to attract night‐time pollinators, such as Manduca sexta hawkmoths. For this pollinator, BA is not only essential for the pollen transport between conspecific plants, but also for the moth’s short‐distance handling of flowers at night. The emission of BA from the corolla limb, however, starts up to 4 hr before dusk, peaks at dusk and continues throughout the first half of the night. While the function of the nocturnal part of the emission is well studied, the function of the early part of the peak emission remained unknown. By using plants silenced in the expression of the biosynthetic gene of BA (Nachal1), we investigated the function of this early peak emission of BA in the plants´ native habitat in Southwestern Utah. By emitting BA before dusk, N. attenuata is able to prevent the establishment and resulting floral damage by a florivore, the cucumber beetle (Diabrotica undecimpunctata). Plants lacking BA emissions (CHAL) were not only more often colonized by florivores, but also suffered significantly more damage, than control empty vector plants (EV). Choice assays revealed that D. undecimpunctata feeding required the absence of BA. While feeding damage to CHAL flowers was higher than to EV flowers in the early night hours, the effect disappeared during the second half of the night, when flowers stop emitting BA. The close proximity of BA‐emitting flowers to non‐emitting CHAL flowers prevented D. undecimpunctata feeding. In the field, the emission of BA at dusk was sufficient to protect plants for the entire night, as beetles search for new hosts during the early evening hours and remain for the duration of the night, once a host is chosen. Floral BA emission before dusk may have evolved as a consequence of antagonistic interactions with florivores. A single floral volatile can thus simultaneously function as an important floral attractant for pollinators and as effective feeding deterrent against florivores in the same plant species. A plain language summary is available for this article.

Flowers are required for the Darwinian fitness of flowering plants, but flowers' advertisements for pollination services can attract florivores. Previous glasshouse work with Nicotiana attenuata revealed the role of jasmonate (JA) signaling in flower development, advertisement and defense. However, whether JA signaling mediates flowers' filtering of floral visitors in nature remained unknown. This field study revealed that silencing JA signaling resulted in flowers that produce less nectar and benzyl acetone, two pollinator-attractive traits. Meanwhile, flowers of defenseless plants were highly attacked by a suite of native herbivores, and damage to buds in native plants correlated negatively with their JA-Ile levels.

This paper describes ongoing research in the area of solar PV production forecasting intended to address a range of effects on the utility grid associated with high penetrations of PV. The ability to anticipate near-term -minutes ahead to hours ahead to day or multiple-day ahead--production of the variable solar resource will be key to successfully integrating ever larger PV capacities with minimal costs. A number of forecast methodologies are surveyed and a mechanism for validating their performance is described.

Most flowering plants depend on animal pollination for successful sexual reproduction. Floral signals such as color, shape, and odor are crucial in establishing this (often mutualistic) interaction. Plant and pollinator phenotypes can vary temporally but also spatially, thus creating mosaic-like patterns of local adaptations. Here, we investigated natural variation in floral morphology, flower volatile emission, and phenology in four accessions of a self-compatible wild tobacco, Nicotiana attenuata, to assess how these traits match the sensory perception of a known pollinator, the hawkmoth Manduca sexta. These accessions differ in floral traits and also in their habitat altitudes. Based on habitat temperatures, the accession occurring at the highest altitude (California) is less likely to be visited by M. sexta, while the others (Arizona, Utah 1, and Utah 2) are known to receive M. sexta pollinations. The accessions varied significantly in flower morphologies, volatile emissions, flower opening, and phenology, traits likely important for M. sexta perception and floral handling. In wind tunnel assays, we assessed the seed set of emasculated flowers after M. sexta visitation and of natural selfed and hand-pollinated selfed flowers. After moth visitations, plants of two accessions (Arizona and Utah 2) produced more capsules than the other two, consistent with predictions that accessions co-occurring with M. sexta would benefit more from the pollination services of this moth. We quantified flower and capsule production in four accessions in a glasshouse assay without pollinators to assess the potential for self-pollination. The two Utah accessions set significantly more seeds after pollen supplementation compared with those of autonomous selfing flowers, suggesting a greater opportunistic benefit from efficient pollinators than the other two. Moreover, emasculated flowers of the accession with the most exposed stigma (Utah 2) produced the greatest seed set after M. sexta visitation. This study reveals intraspecific variation in pollination syndromes that illuminate the potential of a plant species to adapt to local pollinator communities, changing environments, and altered pollination networks.