Floral nectar: fifty years of new ecological perspectives beyond pollinator reward

Floral nectar is central to ecology, since it mediates interactions with pollinators, flower-visiting antagonists and microbes through its chemical composition. Here we review how historical assumptions about its ecological meaning were first challenged, then modified and expanded since the discovery of secondary metabolites in nectar. We then explore the origin of specific neuroactive nectar compounds known to act as important insect neurotransmitters, and how advances in the field of bee cognition and plant-microbe-animal interactions challenge such historical views. As all actors involved in the latter interactions are under simultaneous reciprocal selective pressures, their coexistence is characterized by conflicts and trade-offs, the evolutionary interpretation of which suggests exciting new perspectives in one of the longest studied aspects of plant-pollinator interactions

Secondary Metabolites in Nectar-Mediated Plant-Pollinator Relationships

n recent years, our understanding of the complex chemistry of floral nectar and its ecological implications for plant-pollinator relationships has certainly increased. Nectar is no longer considered merely a reward for pollinators but rather a plant interface for complex interactions with insects and other organisms. A particular class of compounds, i.e., nectar secondary compounds (NSCs), has contributed to this new perspective, framing nectar in a more comprehensive ecological context. The aim of this review is to draft an overview of our current knowledge of NSCs, including emerging aspects such as non-protein amino acids and biogenic amines, whose presence in nectar was highlighted quite recently. After considering the implications of the different classes of NSCs in the pollination scenario, we discuss hypotheses regarding the evolution of such complex nectar profiles and provide cues for future research on plant-pollinator relationships

Composition of sexual fluids in Cycas revoluta ovules during pollination and fertilization

The composition of fuids that mediate fertilization in cycads is described for the frst time. Using tandem mass spectrometry, proteomes of two stages of fuid pro‑ duction, megagametophyte fuid and archegonial chamber fuid production, are compared in Cycas revoluta. These were compared with the proteome of another sexual fuid produced by ovules, the pollination drop proteins. Cycad ovules pro‑ duce complex liquids immediately prior fertilization. Compared with the pollination drops that mainly had few proteins in classes involved in defense and carbohydrate modifcation, megagametophyte fuid and archegonial chamber fuid had larger pro‑ teomes with many more protein classes, e.g. proteins involved in programmed cell death. Using high-performance liquid chromatography, megagametophyte fuid and archegonial chamber fuid were shown to have elevated concentrations of smaller molecular weight molecules including glucose, pectin and glutamic acid. Compared to megagametophyte fuid, archegonial chamber fuid had elevated pH as well as higher osmolality

Nectar in plant-insect mutualistic relationships: from food reward to partner manipulation

It has been known for centuries that floral and extrafloral nectar secreted by plants attracts and rewards animals. Extrafloral nectar is involved in so-called indirect defence by attracting animals (generally ants) that prey on herbivores, or by discouraging herbivores from feeding on the plant. Floral nectar is presented inside the flower close to the reproductive organs and rewards animals that perform pollination while visiting the flower. In both cases nectar is a source of carbon and nitrogen compounds that feed animals, the most abundant solutes being sugars and amino acids. Plant-animal relationships involving the two types of nectar have therefore been used for a long time as text-book examples of symmetric mutualism: services provided by animals to plants in exchange for food provided by plants to animals. Cheating (or deception or exploitation), namely obtaining the reward/service without returning any counterpart, is however well known in mutualistic relationships, since the interacting partners have conflicting interests and selection may favour cheating strategies. A more subtle way of exploiting mutualism was recently highlighted. It implies the evolution of strategies to maximize the benefits obtained by one partner while still providing the reward/service to the other partner. Several substances other than sugars and amino acids have been found in nectar and some affect the foraging behaviour of insects and potentially increase the benefits to the plant. Such substances can be considered plant cues to exploit mutualism. Recent evidence motivated some authors to use the term “manipulation” of animals by plants in nectar-mediated mutualistic relationships. This review highlights the recent background of the “manipulation” hypothesis, discussing it in the framework of new ecological and evolutionary scenarios in plant-animal interactions, as a stimulus for future research

Extrafloral-nectar based partner manipulation in plant-ant relationship

Plant–ant interactions are generally considered as mutualisms, with both parties gaining benefits from the association. It has recently emerged that some of these mutualistic associations have, however, evolved towards other forms of relationships and, in particular, that plants may manipulate their partner ants to make reciprocation more beneficial, thereby stabilizing the mutualism. Focusing on plants bearing extrafloral nectaries, we review recent studies and address three key questions: (i) how can plants attract potential partners and maintain their services; (ii) are there compounds in extrafloral nectar that could mediate partner manipulation; and (iii) are ants susceptible to such compounds? After reviewing the current knowledge on plant–ant associations, we propose a possible scenario where plant-derived chemicals, such as secondary metabolites, known to have an impact on animal brain, could have evolved in plants to attract and manipulate ant behaviour. This new viewpoint would place plant–animal interaction in a different ecological context, opening new ecological and neurobiological perspectives of drug seeking and use

Nutraceutical Characteristics of Ancient Malus x domestica Borkh. Fruits Recovered across Siena in Tuscany

Background: A diet rich in fruits and vegetables contributes to lowering the risk of chronic diseases. The fruits of Malus x domestica are a rich dietary source of bioactive compounds, namely vitamins and antioxidants, with recognized action on human health protection. Tuscany is known for its rich plant biodiversity, especially represented by ancient varieties of fruit trees. Particularly noteworthy are the many ancient Tuscan varieties of apple trees. Methods: Sugar quantification via HPLC and spectrophotometric assays to quantify the antioxidant power and total polyphenol content revealed interesting differences in 17 old varieties of Malus x domestica Borkh. recovered in Siena (Tuscany). Results: The quantification of antioxidants, polyphenols, and the main free sugars revealed that their content in the old fruits was often superior to the widespread commercial counterparts (‘Red Delicious’ and ‘Golden Delicious’). Such differences were, in certain cases, dramatic, with 8-fold higher values. Differences were also present for sugars and fibers (pectin). Most ancient fruits displayed low values of glucose and high contents of xylitol and pectin. Conclusions: The results reported here suggest the possible use of ancient apple varieties from Siena for nutraceutical purposes and draw attention to the valorization of local old varieties

Spatiotemporal variation in the pollination systems of a supergeneralist plant: is Angelica sylvestris (Apiaceae) locally adapted to its most effective pollinators?

Background and Aims: In terms of pollination systems, umbellifers (plants of the carrot family, Apiaceae) are regarded as generalists, since their (usually dichogamous) flowers are visited by a wide range of insects representing several taxonomic orders. However, recent analyses of insect effectiveness revealed that these plants may be pollinated effectively by a narrow assemblage of insect visitors. Of particular interest was whether populations of an umbellifer species varied in pollinator assemblages and whether this could lead to local specialization of the pollination system. We also explored whether variation in pollinator assemblages was associated with variation in floral traits, and whether this variation influences reproductive output. Methods: The focus was on Angelica sylvestris, a common European species visited by a taxonomically diverse insect assemblage. In three populations, located along an ~700-km transect, over three growth seasons insect visitors were identified, their effectiveness was assessed by surveying pollen loads present on the insect body, insect activity on umbels, nectar and scent composition was studied, and transplantation experiments were performed. Key Results: The populations investigated in this study differed in their nectar and scent profiles and, despite the similar taxonomic composition of insect visitor assemblages, were effectively pollinated by disparate pollinator morphogroups, i.e. flies and beetles. Although this suggested local adaptations to the most effective pollinators, analyses of body pollen loads and behaviour on umbels demonstrated functional equivalency of the visitor morphogroups, which is probably related to the fact that A. sylvestris bears few ovules per flower. The transplantation experiments confirmed that reproductive success was not related to the source of experimental plants and that the insects do not exhibit preferences towards local genotypes. Conclusions: Angelica sylvestris is morphologically well adapted to ecological generalization, and there is little evidence that the surveyed populations represent distinct pollination ecotypes. Most likely, the observed variation in floral characters can be interpreted as 'adaptive wandering'. Specialization in this family seems possible only under very special circumstances, for example when the pollinator community comprises insect visitor groups that clearly differ in their pollination capacity (e.g. due to differences in their functional morphology) and/or have different perceptional biases (e.g. for colour or scent). However, the barrier to the evolution of morphological adaptations resulting in the fine-tuning of the flower towards particular pollinator types may arise from the architectural constraints on the floral bauplan that make umbellifers so uniform in their floral displays and so successful in attracting large numbers of pollinators

Drought Stress Affects the Response of Italian Local Tomato (Solanum lycopersicum L.) Varieties in a Genotype-Dependent Manner

Drought stress is one of the most severe conditions for plants, especially in the face of the emerging problem of global warming. This issue is important when considering economically relevant crops, including the tomato. For these plants, a promising solution is the valorization of local agrobiodiversity as a source of genetic variability. In this paper we investigated how six Italian tomato varieties react to a prolonged period of water depletion. We used a multidisciplinary approach, from genetics to plant physiology and cytology, to provide a detailed overview of the response of plants to stress. The varieties analyzed, each characterized by a specific genetic profile, showed a genotype-specific response with the variety 'Fragola' being the most resistant and the variety 'Pisanello' the most susceptible. For all the parameters evaluated, 'Fragola' performed in a manner comparable to that of control plants. On the contrary, 'Pisanello' appeared to be more affected and showed an increase in the number of stomata and a drastic increase in antioxidants, a symptom of acute oxidative stress. Our work suggests the existence of a valuable reservoir of genetic biodiversity with more drought-tolerant tomato genotypes opening the way to further exploitation and use of local germplasm in breeding programs

Nectar sugars and bird visitation define a floral niche for basidiomycetous yeast on the Canary Islands

Studies on the diversity of yeasts in floral nectar were first carried out in the late 19th century. A narrow group of fermenting, osmophilous ascomycetes were regarded as exclusive specialists able to populate this unique and species poor environment. More recently, it became apparent that microorganisms might play an important role in the process of plant pollination. Despite the importance of these nectar dwelling yeasts, knowledge of the factors that drive their diversity and species composition is scarce