Postpollination nectar reabsorption in the African epiphyte Aerangis verdickii (Orchidaceae)

Nectar reabsorption in flowering plants seems to be a rare phenomenon and to our knowledge it has not been reported previously in the Orchidaceae. In this study we present data that show statistically significant differences in nectar sugar concentrations before and after pollination. Virgin flowers of the African epiphytic orchid Aerangis verdickii showed mean sugar concentrations in the nectar of 14.4 and of 18.3% in 1994 and 1995, respectively (sucrose mass by mass equivalent). Sugar concentration in nectar from naturally pollinated flowers had a mean of 3.2 and of 8.7% for the same period. In an experiment in the wild 60 flowers were protected against nectar theft by the ants. Half of those flowers were hand-pollinated. Control (virgin) flowers had nectar with a mean value of 13.8%, while flowers measured 48 h after pollination had a mean sugar concentration of 4.3%. The mean volume of nectar based on measurements of 23 virgin flowers from nine different plants was 19 uL. We estimated an average energy value for the nectar produced of 684 J per plant per season. At the study site, over 60% of the unmanipulated flowers were robbed of all their nectar by arboreal ants, Polyrachis spp. (Formicinae). The adaptive significance of nectar reabsorption for A. verdickii is probably a function of the environmental stresses to which it has been exposed and the relative costs of nectar production.     

Extranuptial nectaries in flowers: ants increase the reproductive success of the ant‐plant Miconia tococa (Melastomataceae)

• Although the production of extranuptial nectar is a common strategy of indirect defence against herbivores among tropical plants, the presence of extranuptial nectaries in reproductive structures is rare, especially in ant‐plants. This is because the presence of ants in reproductive organs can generate conflicts between the partners, as ants can inhibit the activity of pollinators or even castrate their host plants. Here we evaluate the hypothesis that the ant‐plant Miconia tococa produces nectar in its petals which attracts ants and affects fruit set.
• Floral buds were analysed using anatomical and histochemical techniques. The frequency and behaviour of floral visitors were recorded in field observations. Finally, an ant exclusion experiment was conducted to evaluate the effect of ant presence on fruit production.
• The petals of M. tococa have a secretory epidermis that produces sugary compounds. Nectar production occurred during the floral bud stage and attracted 17 species of non‐obligate ants (i.e. have a facultative association with ant‐plants). Ants foraged only on floral buds, and thus did not affect the activity of pollinators in the neighbouring open flowers. The presence of ants in the inflorescences increased fruit production by 15%.
• To our knowledge, the production of extranuptial nectar in the reproductive structures of a myrmecophyte is very rare, with few records in the literature. Although studies show conflicts between the partners in the ant–plant interaction, ants that forage on M. tococa floral buds protect the plant against floral herbivores without affecting bee pollination.

     

Effects of nectar-robbing on plant reproduction and evolution

The relationship between plant and pollinator is considered as the mutualism because plant benefits from the pollinator's transport of male gametes and pollinator benefits from plant's reward.Nectar robbers are frequently described as cheaters in the plant-pollinator mutualism,because it is assumed that they obtain a reward (nectar) without providing a service (pollination).Nectar robbers are birds,insects,or other flower visitors that remove nectar from flowers through a hole pierced or bitten in the corolla.Nectar robbing represents a complex relationship between animals and plants.Whether plants benefit from the relationship is always a controversial issue in earlier studies.This paper is a review of the recent literatures on nectar robbing and attempts to acquire an expanded understanding of the ecological and evolutionary roles that robbers play.Understanding the effects of nectar robbers on the plants that they visited and other flower visitors is especially important when one considers the high rates of robbing that a plant population may experience and the high percentage of all flower visitors that nectar robbers make to some species.There are two standpoints in explaining why animals forage on flowers and steal nectar in an illegitimate behavior.One is that animals can only get food in illegitimate way because of the mismatch of the morphologies of animals'mouthparts and floral structure.The other point of view argues that nectar robbing is a relatively more efficient,thus more energy-saving way for animals to get nectar from flowers.This is probably associated with the difficulty of changing attitudes that have been held for a long time.In the case of positive effect,the bodies of nectar robbers frequently touch the sex organs of plants during their visiting to the flowers and causing pollination.The neutral effect,nectar robbers' behavior may destruct the corollas of flowers,but they neither touch the sex organs nor destroy the ovules.Their behavior does not affect the fruit sets or seed sets of the hosting plant.Besides the direct impacts on plants,nectar robbers may also have an indirect effect on the behavior of the legitimate pollinators.Under some circumstances,the change in pollinator behavior could result in improved reproductive fitness of plants through increased pollen flow and out-crossing.     

Induced Floral and Extrafloral Nectar Production Affect Ant‐pollinator Interactions and Plant Fitness

Thousands of plant species throughout tropical and temperate zones secrete extrafloral nectar to attract ants, whose presence provides an indirect defense against herbivores. Extrafloral nectaries are located close to flowers and may modify competition between ants and pollinators. Here, we used Lima bean (Phaseolus lunatus L.) to study the plants interaction between ants and flower visitors and its consequences for plant fitness. To test these objectives, we carried out two field experiments in which we manipulated the presence of ants and nectar production via induction with jasmonic acid (JA). We then measured floral and extrafloral nectar production, the number of patrolling ants and flower visitors as well as specific plant fitness traits. Lima bean plants under JA induction produced more nectar in both extrafloral nectaries and flowers, attracted more ants and produced more flowers and seeds than non‐induced plants. Despite an increase in floral nectar in JA plants, application of this hormone had no significant effects on flower visitor attraction. Finally, ant presence did not result in a decrease in the number of visits, but our results suggest that ants could negatively affect pollination efficiency. In particular, JA‐induced plants without ants produced a greater number of seeds compared with the JA‐treated plants with ants.     

Introducing plants for bee-keeping at any cost? – Assessment of Phacelia tanacetifolia as nectar source plant under xeric Mediterranean conditions

 We assessed nectar and honey potential of Phacelia tanacetifolia, an American plant, under Mediterranean conditions. Its flowering occurred during the major flowering season unless intensely irrigated, whereas duration and flower life span were shorter compared to continental conditions. Mediterranean climate limitations to nectar secretion were assessed on plants grown under natural conditions (xeric) vs. regular irrigation. May flowering xeric plants produced less nectar per flower than the irrigated ones, but had the same nectar potential per surface area. On the contrary, at the cost of intense irrigation, July flowering plants produced much higher nectar per flower and per surface area compared to xeric ones. In all flowering periods and sections honeybees were the most constant and numerous visitors, visiting the flowers mainly for nectar, whereas solitary bees were scarce. Based on our results, we suggest that although the plant may be a good nectar source for honeybees in some cases, we have serious reservations for a generalised use in the Mediterranean. Received September 3, 2002; accepted November 18, 2002 Published online: June 2, 2003     

Novel nectar robbing negatively affects reproduction in Digitalis purpurea

With many plant–pollinator interactions undergoing change as species’ distributions shift, we require a better understanding of how the addition of new interacting partners can affect plant reproduction. One such group of floral visitors, nectar robbers, can deplete plants of nectar rewards without contributing to pollination. The addition of nectar robbing to the floral visitor assemblage could therefore have costs to the plant´s reproductive output. We focus on a recent plant colonist, Digitalis purpurea, a plant that in its native range is rarely robbed, but experiences intense nectar robbing in areas it has been introduced to. Here, we test the costs to reproduction following experimental nectar robbing. To identify any changes in the behavior of the principal pollinators in response to nectar robbing, we measured visitation rates, visit duration, proportion of flowers visited, and rate of rejection of inflorescences. To find the effects of robbing on fitness, we used proxies for female and male components of reproductive output, by measuring the seeds produced per fruit and the pollen export, respectively. Nectar robbing significantly reduced the rate of visitation and lengths of visits by bumblebees. Additionally, bumblebees visited a lower proportion of flowers on an inflorescence that had robbed flowers. We found that flowers in the robbed treatment produced significantly fewer seeds per fruit on average but did not export fewer pollen grains. Our finding that robbing leads to reduced seed production could be due to fewer and shorter visits to flowers leading to less effective pollination. We discuss the potential consequences of new pollinator environments, such as exposure to nectar robbing, for plant reproduction.     

Relationships between the ant Brachymyrmex obscurior (Hymenoptera, Formicidae) and Acacia pennatula (Fabaceae)

Summary. In central Mexico, the ant Brachymyrmex obscurior Forel feeds on nectar produced by extrafloral nectaries of Acacia pennatula (Schlecht. & Cham.) Benth. However, no studies have determined whether the ants visitation is related to plant nectar availability and whether ants protect A. pennatula from herbivory. The objectives of this 2-yr study (2000–2001) were to assess whether seasonal changes in ant visitation coincide with extrafloral nectar productivity in A. pennatula and to determine whether ants protect the plant. At the end of the dry season (April–June) B. obscurior was the only ant species on A. pennatula and extrafloral nectar production is limited to this period. Exclusion experiments, performed at the end of the dry season showed that A. pennatula did not receive a protective benefit when visited by ants. Branches with ants and branches where ants are excluded had similar numbers of the nonmyrmecophile leafhopper Sibovia sp. which was the only herbivore observed under natural conditions.Received 24 March 2004; revised 4 September 2004; accepted 8 September 2004.     

Unpredictability of nectar nicotine promotes outcrossing by hummingbirds in Nicotiana attenuata

Many plants use sophisticated strategies to maximize their reproductive success via outcrossing. Nicotiana attenuata flowers produce nectar with nicotine at concentrations that are repellent to hummingbirds, increasing the number of flowers visited per plant. In choice tests using native hummingbirds, we show that these important pollinators learn to tolerate high‐nicotine nectar but prefer low‐nicotine nectar, and show no signs of nicotine addiction. Nectar nicotine concentrations, unlike those of other vegetative tissues, are unpredictably variable among flowers, not only among populations, but also within populations, and even among flowers within an inflorescence. To evaluate whether variations in nectar nicotine concentrations increase outcrossing, polymorphic microsatellite markers, optimized to evaluate paternity in native N. attenuata populations, were used to compare outcrossing in plants silenced for expression of a biosynthetic gene for nicotine production (Napmt1/2) and in control empty vector plants, which were antherectomized and transplanted into native populations. When only exposed to hummingbird pollinators, seeds produced by flowers with nicotine in their nectar had a greater number of genetically different sires, compared to seeds from nicotine‐free flowers. As the variation in nectar nicotine levels among flowers in an inflorescence decreased in N. attenuata plants silenced in various combinations of three Dicer‐like (DCL) proteins, small RNAs are probably involved in the unpredictable variation in nectar nicotine levels within a plant.     

Do ant visitors to extrafloral nectaries of plants repel pollinators and cause an indirect cost of mutualism?

Plants and ants have widespread relationships that are commonly mediated by the offer of extrafloral nectar (EFN) to ants that protect plants against herbivores. However, these ant–plant interactions are highly facultative and vary in time and space, mainly depending on the characteristics of the ant species, such as density and aggressiveness. In general, the outcomes of these relationships are positive, but in some cases, the presence of ants is neutral or negative to plants. Some studies suggest that aggressive attacks or merely the presence of ants might reduce the visitation rate of insect pollinators, such as bees, to flowers. We used experimental manipulation in natural conditions to test the hypothesis that ants on flowers of EFN-bearing plants might be recognized as a danger by pollinators (bees) and reduce the plant fitness (fruit-set). Our results show that the avoidance that ant bodyguard species feeding on EFNs of the Malpighiaceae Heteropterys pteropetala cause in pollinators, is not enough to decrease plant fruit-set. However, ants were indeed identified as a danger to pollinators as hypothesized and as suggested for other plant–pollinator relationships: flowers with plastic ants placed on the petals produced significantly fewer fruits than other treatments (using instead a plastic circle) or the control (natural condition). Indirect costs of facultative mutualisms are the focus of few studies and have been performed only rarely in the Neotropics; our results show that mutualism must be considered in multitrophic interactions studies for a better understanding of the functioning of the system.     

Effects of nectar-robbing on plant reproduction and evolution

The relationship between plant and pollinator is considered as the mutualism because plant benefits from the pollinator’s transport of male gametes and pollinator benefits from plant’s reward. Nectar robbers are frequently described as cheaters in the plant-pollinator mutualism, because it is assumed that they obtain a reward (nectar) without providing a service (pollination). Nectar robbers are birds, insects, or other flower visitors that remove nectar from flowers through a hole pierced or bitten in the corolla. Nectar robbing represents a complex relationship between animals and plants. Whether plants benefit from the relationship is always a controversial issue in earlier studies. This paper is a review of the recent literatures on nectar robbing and attempts to acquire an expanded understanding of the ecological and evolutionary roles that robbers play. Understanding the effects of nectar robbers on the plants that they visited and other flower visitors is especially important when one considers the high rates of robbing that a plant population may experience and the high percentage of all flower visitors that nectar robbers make to some species. There are two standpoints in explaining why animals forage on flowers and steal nectar in an illegitimate behavior. One is that animals can only get food in illegitimate way because of the mismatch of the morphologies of animals’ mouthparts and floral structure. The other point of view argues that nectar robbing is a relatively more efficient, thus more energy-saving way for animals to get nectar from flowers. This is probably associated with the difficulty of changing attitudes that have been held for a long time. In the case of positive effect, the bodies of nectar robbers frequently touch the sex organs of plants during their visiting to the flowers and causing pollination. The neutral effect, nectar robbers’ behavior may destruct the corollas of flowers, but they neither touch the sex organs nor destroy the ovules. Their behavior does not affect the fruit sets or seed sets of the hosting plant. Besides the direct impacts on plants, nectar robbers may also have an indirect effect on the behavior of the legitimate pollinators. Under some circumstances, the change in pollinator behavior could result in improved reproductive fitness of plants through increased pollen flow and out-crossing. __________ Translated from Acta phytoecologiaca Sinica, 2006, 30(4): 695–702 [译自: 植物生态学报]     

Flowers respond to pollinator sound within minutes by increasing nectar sugar concentration

Can plants sense natural airborne sounds and respond to them rapidly? We show that Oenothera drummondii flowers, exposed to playback sound of a flying bee or to synthetic sound signals at similar frequencies, produce sweeter nectar within 3 min, potentially increasing the chances of cross pollination. We found that the flowers vibrated mechanically in response to these sounds, suggesting a plausible mechanism where the flower serves as an auditory sensory organ. Both the vibration and the nectar response were frequency‐specific: the flowers responded and vibrated to pollinator sounds, but not to higher frequency sound. Our results document for the first time that plants can rapidly respond to pollinator sounds in an ecologically relevant way. Potential implications include plant resource allocation, the evolution of flower shape and the evolution of pollinators sound. Finally, our results suggest that plants may be affected by other sounds as well, including anthropogenic ones.     

The reproductive biology ofJaborosa integrifolia (Solanaceae): Why its fruits are so rare?

Jaborosa integrifolia exhibits stigma-height polymorphism. There are individuals with flowers where anthers and stigma are at the same height but the rule is variable herkogamy, the most common type (75%) being that with an exerted stigma. Self- and cross-tubes did not differ in their capability to reach the ovary (t = –0.67,P < 0.53); they had a high growth rate (6.95 ± 2.28 mm h^–1). There is not autogamy but mostly self-incompatibility. Fruits from controlled cross-pollination showed the highest seed set and seed viability. The nectar sugar is characterized by a similar amount of glucose and fructose, and by the absence of sucrose. Although nectar secretion was continuous throughout the life of the flower, most nectar was secreted during the first 24 h after flower opening. Nectar production costs appear to be lower than in other species since nectar secretion is neither inhibited after a removal (i.e. a pollinator visit) nor reabsorbed as the flower ages. Sphingids visit the flowers mainly after midnight. They insert their proboscis down to the base of the corolla tube to reach the nectar. The upper limit to fruit production is set by pollinator visits. Fruits produced from open-pollinated flowers are often predated by numerous larvae (mainly lepidopteran ones). Considering that this species is mostly self-incompatible and pollination is limited, that each plant displays only a low number of flowers throughout the flowering season, and that there is a high rate of fruit predation, it is not surprising that fruits ofJ. integrifolia are so rare.     

Flower nectar of an autogamous perennialRorippa indica as an indirect defense mechanism against herbivorous insects

This report shows that one of the most important roles of the flower nectar of an autogamous perennialRorippa indica (L.) Hieron is as an attractant for employing some ant species as a defense against herbivorous insects. The plant has flowers from spring to early winter. Its flower nectar is frequently stolen by some ant species (hereafter cited as ants) which also feed on small herbivorous insects on the plant. Internations among the tritrophic levels (R. indica, herbivores, ants) were experimentally examined and the followings became clear. (1) Ants were attracted toR. indica in search of its flower nectar. (2) The gradual secretion of flower nectar seemed to detain ants on the plant. (3)Pieris butterfly lavae were the major herbivores onR. indica and were potentially harmful to the plant. (4) The presence of ants reduced the survival rate ofP. rapae larvae onR. indica. (5) The presence of ants reduced the feeding damage toR. indica. (6) The disadvantage of nectar use by ants seemed to be minimal for the plant since the ants did not disturb the other flower visitors. These facts suggest a mutualistic relationship betweenR. indica and ants. That is, the flower nectar serves as an indirect defense against herbivorous insects.     

Changing light conditions in pine rockland habitats affect the intensity and outcome of ant–plant interactions

Extrafloral nectar (EFN) mediates food‐for‐protection mutualisms between plants and ants. Such mutualisms exist within a complex web of biotic interactions, and in a framework provided by the abiotic environment. Both biotic and abiotic factors, therefore, affect the outcome of ant–plant interactions. We conducted an experiment to determine the effects of ant activity, and light intensity, on herbivory rates, growth, and reproductive fitness in Senna mexicana var. chapmanii, a perennial legume native to pine rockland habitats of south Florida. Forty plants were divided among four treatments in a factorial experimental design with two independent variables: ant activity and light intensity. Plants were divided equally between sunny and shady habitats, and ants were excluded from half of the plants in each habitat type. The presence of ants significantly reduced herbivory rates in S. chapmanii. In shaded habitats, the presence of ants had no effect on plant reproductive fitness, however, in sunny habitats plants with ants produced significantly more seeds over the duration of the 1‐yr study. Ants represent an important biotic defense against herbivores in S. chapmanii; however, their effects on plant fitness are dependent on light conditions. Pine rockland habitats in south Florida have been widely destroyed or mismanaged. In fragments that remain, suppression of fire has led to increased canopy closure and shading of the understory. These changes will likely negatively impact plants that rely on ants for defense. We highlight the importance of conservation efforts to preserve the pine rocklands and the fire regimes on which they rely.     

Floral nectar production: what cost to a plant?

Floral nectar production is central to plant pollination, and hence to human wellbeing. As floral nectar is essentially a solution in water of various sugars, it is likely a valuable plant resource, especially in terms of energy, with plants experiencing costs/trade-offs associated with its production or absorption and adopting mechanisms to regulate nectar in flowers. Possible costs of nectar production may also influence the evolution of nectar volume, concentration and composition, of pollination syndromes involving floral nectar, and the production of some crops. There has been frequent agreement that costs of floral nectar production are significant, but relevant evidence is scant and difficult to interpret. Convincing direct evidence comes from experimental studies that relate either enhanced nectar sugar production (through repeated nectar removal) to reduced ability to produce seeds, or increased sugar availability (through absorption of additional artificial nectar) to increased seed production. Proportions of available photosynthate allocated by plants to nectar production may also indicate nectar cost. However, such studies are rare, some do not include treatments of all (or almost all) flowers per plant, and all lack quantitative cost–benefit comparisons for nectar production. Additional circumstantial evidence of nectar cost is difficult to interpret and largely equivocal. Future research should repeat direct experimental approaches that relate reduced or enhanced nectar sugar availability for a plant with consequent ability to produce seeds. To avoid confounding effects of inter-flower resource transfer, each plant should experience a single treatment, with treatment of all or almost all flowers per plant. Resource allocation by plants, pathways used for resource transfer, and the locations of resource sources and sinks should also be investigated. Future research should also consider extension of nectar cost into other areas of biology. For example, evolutionary models of nectar production are rare but should be possible if plant fitness gains and costs associated with nectar production are expressed in the same currency, such as energy. It should then be possible to understand observed nectar production for different plant species and pollination syndromes involving floral nectar. In addition, potential economic benefits should be possible to assess if relationships between nectar production and crop value are evaluated.     

Pollinators shift to nectar robbers when florivory occurs,with effects on reproductive success in Iris bulleyana (Iridaceae)

• Studies have indicated that florivory and nectar robbing may reduce reproductive success of host plants. However, whether and how these effects might interact when plants are simultaneously attacked by both florivores and nectar robbers still needs further investigation.
• We used Iris bulleyana to detect the interactions among florivory, nectar robbing and pollination, and moreover, their effects on plant reproductive success. Field investigations and hand‐pollination treatments were conducted on two experimental plots from a natural population, in which Experimental plot was protected from florivores and Control plot was not manipulated.
• The flower calyx was bitten by sawflies to consume the nectary, and three bumblebee species were pollinators. In addition, the short‐tongued pollinator, Bombus friseanus, was the only robber when there was a hole made by a sawfly. The bumblebee had significantly shortened flower handling time when robbing, as compared to legitimate visits. Pollinator visitation and seed production decreased significantly in damaged flowers. However, seed production per flower after supplementary hand‐pollination did not differ significantly between damaged and undamaged flowers. Compared to the Experimental plot, bumblebees visited fewer flowers per plant in a foraging bout in the Control plot.
• The flowers damaged by florivory allowed B. friseanus to shift to a nectar robber. Florivory and nectar robbing collectively decreased plant reproductive success by consuming nectar resources, which may reduce attractiveness to pollinators of the damaged flowers. However, the changes in pollinator behaviour might be beneficial to the plant by reducing the risk of geitonogamous mating.

     

Effects of nectar robbing on pollinator behavior and plant reproductive success of Pitcairnia angustifolia (Bromeliaceae)

Nectar robbers may have direct and indirect effects on plant reproductive success but the presence of nectar robbing is not proof of negative fitness effects. We combined census data and field experiments to disentangle the complex effects of nectar robbing on nectar production rates, pollinator behavior, pollen export, and female reproductive success of Pitcairnia angustifolia. Under natural conditions flowers were visited by four different animal species including a robber‐like pollinator and a secondary robber. Natural levels of nectar robbing ranged from 40 to 100%. Natural variation in nectar robbing was not associated with fruit set in any year whereas seed set was weakly positively associated for 1 year only. Artificial nectar robbing did not increase nectar production or concentration, did not affect the behavior of long‐billed hummingbirds, and when faced with artificially robbed flowers, these visitors behaved as secondary nectar robbers. The number of stigmas within a patch that received pollen dye analogs and the average distance traveled by these analogs were not significantly different between robbing treatments (robbed flowers versus unrobbed flowers), but the maximum distance traveled by these pollen analogs was higher when nectar robbing was not prevented. Overall, the proportion of robbed flowers on an inflorescence had a neutral effect to a weak positive effect on the reproduction of individual plants (i.e. positive association between nectar robbing and fruit set in 2002) even when it clearly changed the behavior of its most efficient pollinator potentially increasing the frequency of nectar robbing within a plant.     

Cheater or mutualist? Novel florivory interaction between nectar‐rich Crotalaria cunninghamii and small mammals

Animals visit flowers to access resources and by moving pollen to conspecific individuals act as pollinators. While biotic pollinators can increase the seed set of plants, other flower visitors can reduce seed set directly by damaging vital reproductive organs and indirectly by affecting the way the plant interacts with subsequent flower visitors. It is, therefore, vital to understand the varied effects of all visitors and not only pollinators on plant fitness, including those visitors that are temporally or spatially rare. We document the first known case of flower visitation by small mammals to Crotalaria cunninghamii (Fabaceae), a plant species morphologically suited to bird pollination. During a rain‐driven resource pulse in the Simpson Desert in 2011, the rodents Mus musculus (Muridae) and Pseudomys hermannsburgensis (Muridae) visited flowers to remove nectar by puncturing the calyx. We investigated the effects of this novel interaction on the reproductive output of C. cunninghamii. Compared with another recent resource pulse in 2007, plants flowering during mammal visitation had five times as many inflorescences per plant, 90% more flowers per inflorescence, and two to three times more nectar per flower, but this nectar was 30% less sugar rich. Concurrently, rodent plagues were up to three times larger during this rain‐driven resource pulse than during a previous pulse in 2007. Up to 75% of flowers had evidence of small mammal florivory, but this was not necessarily destructive, as up to 90% of fruit had the remains of florivory. Through a series of exclusion experiments, we found that small mammal florivory did not directly reduce seed set. We conclude that rain‐driven resource pulses led to a unique combination of events that facilitated the novel florivory interaction. Our findings emphasize the dynamic nature of biotic interactions and the importance of testing the role of all visitors to pollination services.     

Variation of nectar production in relation to plant characteristics in protandrous Aconitum gymnandrum

Aims Floral nectar plays a vital role in plant reproductive success by attracting pollinators. Nectar traits of a flower can depend directly on plant characteristics other than environmental factors and exhibit extensive flower- and plant-level variations. Studies on nectar traits frequently focused on intraplant variation for dichogamous plants, but few have paid attention to both intra- and interplant nectar variations in relation to plant characteristics. Revealing within- and among-plant variation and its relative magnitude is important for our understanding of how pollinator-mediated selection can act on nectar traits and evolution of nectar traits.Methods Through investigating protandrous Aconitum gymnandrum populations at the Alpine Meadows and Wetland Ecosystems Research Station of Lanzhou University, we examined the relationships between nectar production per flower and plant characteristics (e.g. flower position within inflorescences, floral sexual phases, flowering time, inflorescence size and floral attractive traits).Important findings A. gymnandrum exhibited a declining gradient in the nectar volume along inflorescences, with more nectar in basal flowers than distal ones. Protandrous flowers of A. gymnandrum did not show gender-biased nectar production while the nectar volume varied with different stages of floral sexual phases. The significant correlation between the first flowering date of individuals and the mean nectar volume per flower was positive in 2013, but became negative in 2014, suggesting complex effects of biotic and abiotic factors. The mean nectar volume per flower was not related to inflorescence size (the number of total flowers per plant). Furthermore, nectar production was weakly associated with floral attractive traits (the petal width and the galea height), even if the effect of flowering time of individuals was removed, suggesting that the honesty of floral traits as signals of nectar reward for pollinators is not stable in this species.     

Increased Host Investment in Extrafloral Nectar (EFN) Improves the Efficiency of a Mutualistic Defensive Service

Extrafloral nectar (EFN) plays an important role as plant indirect defence through the attraction of defending ants. Like all rewards produced in the context of a mutualism, however, EFN is in danger of being exploited by non-ant consumers that do not defend the plant against herbivores. Here we asked whether plants, by investing more in EFN, can improve their indirect defence, or rather increase the risk of losing this investment to EFN thieves. We used the obligate plant-ant Acacia-Pseudomyrmex system and examined experimentally in the field during the dry and the rainy seasons how variations in EFN secretion are related to (i) ant activity, to (ii) the ant-mediated defence against herbivores and (iii) the exploitation of EFN by non-ant consumers. Extrafloral investment enhanced ant recruitment and was positively related to the ant mediated defence against herbivores. The ant-mediated protection from exploiters also increased in proportion to the nectar sugar concentration. Although the daily peak of EFN production coincided with the highest activity of EFN thieves, Pseudomyrmex ferrugineus ants protected this resource effectively from exploiters. Nevertheless, the defensive effects by ants differed among seasons. During the dry season, plants grew slower and secreted more EFN than in the rainy season, and thus, experienced a higher level of ant-mediated indirect defence. Our results show that an increased plant investment in an indirect defence trait can improve the resulting defensive service against both herbivores and exploiters. EFN secretion by obligate ant-plants represents a defensive trait for which the level of investment correlates positively with the beneficial effects obtained.