Can selection to escape nectar thieving force plants to portion nectar in many flowers?

There are several hypotheses which try to explain why particular plants produce just the number of flowers that they do. These hypotheses include: compromises between the attraction of pollinators and avoiding self-pollination by geitonogamy; optimal nectar production as a result of diminishing gains of nectar production; opportunity for selective abortion; or different bet-hedging strategies. In this paper, I present a hypothesis which interferes with the others concerning flower numbers in plants: nectar thieving can apply a selection force in plants which result in a portioning of nectar in many flowers. Energy saved by reducing the quantity of nectar in each flower can be directed into greater flower numbers.     

Hummingbird responses to gender-biased nectar production: are nectar biases maintained by natural or sexual selection?

Pollinators mediate the evolution of secondary floral traits through both natural and sexual selection. Gender-biased nectar, for example, could be maintained by one or both, depending on the interactions between plants and pollinators. Here, I investigate pollinator responses to gender-biased nectar using the dichogamous herb Chrysothemis friedrichsthaliana (Gesneriaceae) which produces more nectar during the male floral phase. Previous research showed that the hummingbird pollinator Phaethornis striigularis visited male-phase flowers more often than female-phase flowers, and multiple visits benefited male more than female fecundity. If sexual selection maintains male-biased rewards, hummingbirds should prefer more-rewarding flowers independent of floral gender. If, however, differential rewards are partially maintained through natural selection, hummingbirds should respond to asymmetry with visits that reduce geitonogamy, i.e. selfing and pollen discounting. In plants with male biases, these visit types include single-flower visits and movements from low to high rewards. To test these predictions, I manipulated nectar asymmetry between pairs of real or artificial flowers on plants and recorded foraging behaviour. I also assessed maternal costs of selfing using hand pollinations. For plants with real flowers, hummingbirds preferred more-rewarding flowers and male-phase morphology, the latter possibly owing to previous experience. At artificial arrays, hummingbirds responded to extreme reward asymmetry with increased single-flower visits; however, they moved from high to low rewards more often than low to high. Finally, selfed flowers did not produce inferior seeds. In summary, sexual selection, more so than geitonogamy avoidance, maintains nectar biases in C. friedrichsthaliana, in one of the clearest examples of sexual selection in plants, to date.     

Does spatial aggregation of total nectar production lead to genetic structure?

Spatial aggregation of plants of high nectar production, receiving an enhanced pollinator service is known to occur in Echium vulgare. Moreover, an emanating effect of nectar production on pollinator visits may occur, i.e. many pollinator visits may be observed around high nectar patches. Consequently, gene flow within patches of plants of high nectar production and their close neighbours may result in genetic structure. In this study, we investigated whether aggregation of total nectar production (nectar production per flower×number of flowers) and its emanating effect resulted in genetic structure in a natural E. vulgare population. We compared the spatial structure of total nectar production, pollinator visits and microsatellite markers using spatial autocorrelation analysis. Increased geitonogamy, caused by longer boutlengths in plants of high nectar production may generate genetic structure. We estimated selfing rates of plants of the highest and lowest total nectar production. Spatial aggregation of total nectar production occurred on a relatively small scale up to 2.83 m. A significant emanating correlation between total nectar production and pollinator visits was observed on a relatively large scale up to a 4.24 m. Thus, around patches of high nectar production numbers of pollinator visits were relatively high, while few visits were observed around patches of low nectar production. Weak genetic structure was present on a small scale up to 2.20 m. This corresponded with the scale of aggregation of total nectar production. High gene flow around the patches of high nectar production seems to weaken genetic structure. This is supported by the relatively low selfing rates. The average selfing rate of the plants of highest nectar production was 8.8% and that of the plants of lowest nectar production 5.0%. Low gene flow within and around low nectar patches sustain a weak genetic structure or, conversely, may have caused it in the first instance. Results indicate the importance of spatial structure of nectar production for pollinator movement.     

Why do ants shift their foraging from extrafloral nectar to aphid honeydew?

When aphids parasitize plants with extrafloral nectaries (EFNs) and aphid colony size is small, ants frequently use EFNs but hardly tend aphids. However, as the aphid colony size increases, ants stop using EFNs and strengthen their associations with aphids. Although the shift in ant behavior is important for determining the dynamics of the ant–plant–aphid interaction, it is not known why this shift occurs. Here, we test two hypotheses to explain the mechanism responsible for this behavioral shift: (1) Extrafloral nectar secretion changes in response to aphid herbivory, or (2) plants do not change extrafloral nectar secretion, but the total reward to ants from aphids will exceed that from EFNs above a certain aphid colony size. To judge which mechanism is plausible, we investigated secretion patterns of extrafloral nectar produced by plants with and without aphids, compared the amount of sugar supplied by EFNs and aphids, and examined whether extrafloral nectar or honeydew was more attractive to ants. Our results show that there was no inducible extrafloral secretion in response to aphid herbivory, but the sugar concentration in extrafloral nectar was higher than in honeydew, and more ant workers were attracted to an artificial extrafloral nectar solution than to an artificial aphid honeydew solution. These results indicate that extrafloral nectar is a more attractive reward than aphid honeydew per unit volume. However, even an aphid colony containing only two individuals can supply a greater reward to ants than EFNs. This suggests that the ant behavioral shift may be explained by the second hypothesis.     

Is the nectar redox cycle a floral defense against microbial attack?

Many angiosperms use a remarkable reproductive strategy that relies on attracting animals (insect, avian or mammalian pollinators) to transfer pollen between plants. Relying on other organisms for sexual reproduction seems evolutionarily untenable, but the great diversity of angiosperms illustrates how highly successful this strategy is. To attract pollinators, plants offer a variety of rewards. Perhaps the primary floral reward is floral nectar. Plant nectar has long been considered a simple sugar solution but recent work has demonstrated that nectar is a complex biological fluid containing significant and important biochemistry with the potential function of inhibiting microbial growth. These results lead the way to novel insights into the mechanisms of floral defense and the co-evolution of angiosperms and their pollinators.     

Do extrafloral nectar resources, species abundances, and body sizes contribute to the structure of ant–plant mutualistic networks?

Recent research has shown that many mutualistic communities display non-random structures. While our understanding of the structural properties of mutualistic communities continues to improve, we know little of the biological variables resulting in them. Mutualistic communities include those formed between ants and extrafloral (EF) nectar-bearing plants. In this study, we examined the contributions of plant and ant abundance, plant and ant size, and plant EF nectar resources to the network structures of nestedness and interaction frequency of ant–plant networks across five sites within one geographic locality in the Sonoran Desert. Interactions between ant and plant species were largely symmetric. That is, ant and plant species exerted nearly equivalent quantitative interaction effects on one another, as measured by their frequency of interaction. The mutualistic ant–plant networks also showed nested patterns of structure, in which there was a central core of generalist ant and plant species interacting with one another and few specialist–specialist interactions. Abundance and plant size and ant body size were the best predictors of symmetric interactions between plants and ants, as well as nestedness. Despite interactions in these communities being ultimately mediated by EF nectar resources, the number of EF nectaries had a relatively weak ability to explain variation in symmetric interactions and nestedness. These results suggest that different mechanisms may contribute to structure of bipartite networks. Moreover, our results for ant–plant mutualistic networks support the general importance of species abundances for the structure of species interactions within biological communities.     

Are nectar guide colour changes a reliable signal to pollinators that enhances reproductive success?

Background: Ageing and post-pollination changes in floral colour occur widely in flowering plants, but it remains an open question as to whether or not colour changes in nectar guides are associated with the quantity of floral rewards that ultimately influence pollinator visitations and reproductive success.

Aims: To examine whether nectar guide changes should be considered as a reliable signal to pollinators and to assess the effects of nectar guide changes on reproductive success.

Methods: We studied the process and adaptive value of colour changes in the nectar guides of Arnebia szechenyi whose flowers typically display conspicuous nectar guides at the onset of anthesis, after which they begin to fade, and disappear completely on the second day.

Results: Changes in nectar guide colour in A. szechenyi were intrinsic and age-dependent, although pollination somewhat accelerated the change. By the time that the nectar guides disappeared completely, floral rewards were reduced almost to zero. Artificial removal of nectar guides decreased both fruit set and pollen export. Flowers without nectar guides do not appear to increase the overall attractiveness of the plants.

Conclusions: Nectar guides and their changes represent reliable signals to pollinators and enhance both male and female reproductive success.     

Is nectar reabsorption restricted by the stalk cells of floral and extrafloral nectary trichomes?

Reabsorption is a phase of nectar dynamics that occurs concurrently with secretion; it has been described in floral nectaries that exude nectar through stomata or unicellular trichomes, but has not yet been recorded in extrafloral glands. Apparently, nectar reabsorption does not occur in multicellular secretory trichomes (MST) due to the presence of lipophilic impregnations – which resemble Casparian strips – in the anticlinal walls of the stalk cells. It has been assumed that these impregnations restrict solute movement within MST to occur unidirectionally and exclusively by the symplast, thereby preventing nectar reflux toward the underlying nectary tissues. We hypothesised that reabsorption is absent in nectaries possessing MST. The fluorochrome lucifer yellow (LYCH) was applied to standing nectar of two floral and extrafloral glands of distantly related species, and then emission spectra from nectary sections were systematically analysed using confocal microscopy. Passive uptake of LYCH via the stalk cells to the nectary tissues occurred in all MST examined. Moreover, we present evidence of nectar reabsorption in extrafloral nectaries, demonstrating that LYCH passed the stalk cells of MST, although it did not reach the deepest nectary tissues. Identical (control) experiments performed with neutral red (NR) demonstrated no uptake of this stain by actively secreting MST, whereas diffusion of NR did occur in plasmolysed MST of floral nectaries at the post‐secretory phase, indicating that nectar reabsorption by MST is governed by stalk cell physiology. Interestingly, non‐secretory trichomes failed to reabsorb nectar. The role of various nectary components is discussed in relation to the control of nectar reabsorption by secretory trichomes.     

Does acoustic priming ‘sweeten the pot’ of floral nectar?

A recent claim that evening primrose flowers adaptively secrete nectar in response to vibrations from hovering bees lacks supporting evidence. The authors fail to demonstrate that bees can access the concealed nectar and that their visits enhance plant fitness. Reanalysis of the authors’ data raises additional concerns about their conclusions.     

Feeding by the short‐tailed bat (Mystacina tuberculata) on fruit and possibly nectar

New Zealand's short‐tailed bat (Mystacina tuberculata Gray, 1843) feeds on fruit, insects, and possibly nectar in North Island kauri (Agathis australis) forest. Fruits eaten by members of a colony of 500 bats in May included those of Freycinetia baueriana (Pandanaceae), Collospermum hastatum, and C. microspermum (Liliaceae). Pollen analyses of bat guano, and of the stomach contents of 4 short‐tailed bats from Omahuta Forest (Lat. 35°10'S) and 3 from Stewart Island and adjacent islands (Lat. 47°15'S), showed that most of the pollen was from flowers of Metrosideros and Leptospermum (Myrtaceae), Knightia excelsa (Proteaceae), and Collospermum, and that spores of the tree fern Cyathea (Cyatheaceae) were present also. Both Metrosideros and Knightia have abundant nectar. The partially extensile tongue of Mystacina is tipped with a brush of fine papillae, possibly to extract nectar and pollen; but the pollen and spores in the bat stomachs and guano could have come from insects eaten by the bats. Transverse ridges on the tongue may assist removal of juice from ripe fruits. These bats may disperse the small seeds of Freycinetia baueriana. The anatomical modifications of Mystacina for terrestrial and arboreal locomotion may have evolved primarily in response to its frugivorous and suspected nectarivorous habits.     

How to use the paradigm of ischemic preconditioning to protect the heart?

Ischemic preconditioning affords the most powerful protection to a heart submitted to a prolonged ischemia-reperfusion. During the past decade, a huge amount of work allowed to better understand the features of this protective effect as well as the molecular mechanisms. Ischemic preconditioning reduces infarct size and improves functional recovery; its effects on arrhythmias remain debated. Triggering of the protection involves cell surface receptors that activate pro-survival pathways including protein kinase C, PI3-kinase, possibly Akt and ERK1/2, whose downstream targets remain to be determined. Much attention has been recently focused on the role of mitochondrial K(+)ATP channels and the permeability transition pore that seem to play a major role in the progression toward irreversible cellular injury. Based on these experimental studies attempts have been made to transfer preconditioning from bench to bedside. Human experimental models of ischemic preconditioning have been set up, including cardiac surgery, coronary angioplasty or treadmill exercise, to perform pathophysiological studies. Yet, protecting the heart of CAD (coronary artery disease) patients requires a pharmacological approach. The IONA trial has been an example of the clinical utility of preconditioning. It helped to demonstrate that chronic administration of nicorandil, a K(+)ATP opener that mimics ischemic preconditioning in experimental preparations, improves the cardiovascular prognosis in CAD patients. Recent experimental studies appear further encouraging. It appears that "postconditioning" the heart (i.e. performing brief episodes of ischemia-reperfusion at the time of reperfusion) is as protective as preconditioning. In other words, a therapeutic intervention performed as late as at the time of reflow can still significantly limit infarct size. Further work is needed to determine whether this may be transferred to the clinical practice.     

Do plants dynamically regulate nectar features through sugar sensing?

Nectar properties (volume, concentration, viscosity) change dynamically in time. As stated by Pedersen some decades ago (1958), “Nectar is not a static product remaining outside the plant once produced but is in close contact with the plant system.”¹ It is now evident that secretion may occur concomitantly with resorption and that the latter process sometimes continues after secretion has ended. The rate of the two processes may be modified dynamically by the plant in response to ecological and physiological constraints, maintaining a relatively constant nectar concentration to ensure pollinator visits (nectar homeostasis) and reallocating resources, especially during development of the ovules and pericarp after fertilization. We suspect that nectar resorption is under-estimated as a phenomenon, because it requires detailed information on the dynamics of nectar production throughout the life of the flower that is seldom available or taken into consideration. The cytological and molecular mechanisms involved in nectar resorption are almost completely unknown. Sugar sensing may have a fundamental role in nectar resorption and homeostasis. Due to direct contact with sugar solutions, nectaries may offer wide scope for insights into this phenomenon which has attracted interest as part of plant signalling systems.Key words: nectaries, nectar resorption, nectar homeostasis, nectar composition     

Ant–plant interaction in the Neotropical savanna: direct beneficial effects of extrafloral nectar on ant colony fitness

Current evidence suggests that ant–plant relationships may influence species composition, abundance, and interactions at the community scale. The main resource that plants offer to ants is extrafloral nectar (EFN) and the major part of published studies shown benefits from ants to plants possessing EFNs. However, the complementary question of whether and how ants benefit from EFNs is rarely addressed. Here, we present the results of a long-term study to demonstrate whether EFN has a positive effect on ant colony fitness. We quantified colony growth rate, survival and the final weight of individuals as measures of benefit derived from EFN. Our results provide clear evidence that EFN can have a significant positive impact on the survivorship, growth and reproduction of the Myrmicinae Cephalotes pusillus. In fact, a diet rich in EFN (providing at least 30 cal per day) resulted in five times more individuals per colony, greater body weights, and more eggs. These results have shed new light on the relationships between ants and EFN-bearing plants such as in tropical and temperate systems. The ant C. pusillus is the first case in which we have firm evidence that EFN improves colony growth and development, corroborating more than 100 years of experimental evidence of benefits to plants in these widespread relationships.