It sounds incredibly simple on one level — much like rewarding a dog with a treat after it obeys a command. Flowering plants will optimise the characteristics of their nectar in order to influence the foraging behaviours of pollinators and ultimately improve their reproductive fitness. The characteristics of the nectar not only determine which pollinators are attracted and when they come, but how frequently they visit and how long they stay. Suddenly one realises that there is an extremely complex system of regulatory mechanisms behind nectar secretion, which have not only influenced the evolution of flowering plants, but of the pollinators themselves.
Red Admiral butterfly Vanessa atalanta drinking nectar. There is no denying, however, that carbohydrates — sugars such as glucose, sucrose and fructose — are usually the main constituent of nectar.
Other sugars might also be present in small amounts as well as sugar alcohols, such as sorbitol. It is these sugars that are the primary energy source for nectar consumers. Amino acids and proteins are the next most abundant solute in nectar after the sugars. There are essential and non-essential amino acids, which are the building blocks for proteins and there are some non-protein amino acids that are constituents of enzymes and preservatives.
It is thought that the amino acid and protein content of nectar may play a role in the taste preferences of insects [1], presumably related to their nutritional needs. The water content of nectar may also be an important reward for pollinators, particularly in dry habitats. Nectar also contains important ions, such as potassium, as well as antioxidants, trace amounts of lipids and some secondary compounds that seem to be associated with resistance to herbivory.
Many species have also been shown to have antimicrobial compounds in their nectar, which prevents microbes from growing in the nectar as well as inhibiting florally transmitted diseases [2].
Terpinoids, which are the volatile organic compounds that give flowers their scent, also accumulate in the nectar. Flowers frequented by hummingbirds, for example, generally produce nectar in small amounts with high sugar content, while those frequented by more generalist passerine birds produce dilute nectar in large quantities. The stamen, which is the male part of the flower, includes an anther that holds pollen grains as they form and a filament that supports the anther.
Pollen is a fine, powdery dust, usually yellow. Petals attract insects with the promise of nectar, the sugary liquid found inside the flower carpal at the base of the petals. When insects crawl inside to get to the nectar, pollen grains brush onto the insects, who take it with them to the next flower that offers nectar.
The pollen sticks to the stigma of the next flower, and a tube develops. When the tube reaches the young seeds, or ovules, inside the ovary, sperm in the pollen enters the tube and fertilizes the seeds.
This process only happens when the pollen is spread to flowers of the same kind. Some nectar contains toxins that act as defense mechanisms. This nectar is produced in extrafloral nectaries, usually on the stems or edges of leaves. Some toxins protect flowers from fungus while other nectar contains toxins to repel organisms that take the nectar without helping in reproduction.
Most flowers open in the morning and close in the afternoon so nectar was not available all day. Nectar-sugar concentration and sugar value h increasing temperature. High nectar-foraging activity by honeybees coincided with peak nectar-sugar production.
A nectary is a nectar-secreting gland found in different locations in the flower. The different types of floral nectaries include 'septal nectaries' found on the sepal, 'petal nectaries', 'staminal nectaries' found on the stamen, and 'gynoecial nectaries' found on the ovary tissue. Nectaries can also be categorized as structural or non-structural.
Pollinators feed on the nectar and, depending on the location of the nectary, the pollinator assists in fertilization and outcrossing of the plant as they brush against the reproductive organs, the stamen and pistil, of the plant and pick up or deposit pollen.
Nectar from floral nectaries is sometimes used as a reward to insects, such as ants, that protect the plant from predators. Many floral families have evolved a nectar spur.
These spurs are projections of various lengths formed from different tissues, such as the petals or sepals. They allow for pollinators to land on the elongated tissue and more easily reach the nectaries and obtain the nectar reward. Extra-floral nectaries are nectar-producing glands physically apart from the flower located on leaf laminae, petioles, rachids, bracts, stipules, pedicels, fruit, etc.
Their size, shape and secretions vary with plant species. Extra-floral nectar content differs from floral nectar and may or may not flow in a daily pattern. Two functions for the extra-floral nectar have been hypothesized: 1 as an excretory organ for the plant to rid itself of metabolic wastes or 2 to attract beneficial insects for plant defense. The nectar attracts predatory insects that consume both the nectar and plant-eating arthropods, functioning as bodyguards. Nectar-seeking ants expel herbivores and enhance the reproductive success of plants with extra-floral nectaries.
The greater the importance of extra-floral nectar to the ants, the better for the plants, as this increases the ants' aggressiveness toward herbivores. The actual process of transforming the flower nectar into honey requires teamwork. First, older forager worker bees fly out from the hive in search of nectar-rich flowers. Using its straw-like proboscis, a forager bee drinks the liquid nectar from a flower and stores it in a special organ called the honey stomach. The bee continues to forage until its honey stomach is full, visiting 50 to flowers per trip from the hive.
Extrafloral nectar at the plant-insect interface: a spotlight on chemical ecology, phenotypic plasticity, and food webs. Manipulators live better, but are they always parasites? Partner manipulation stabilises a horizontally transmitted mutualism. Protective ant-plant interactions as model systems in ecological and evolutionary research. Science , — Hendriksma, H. Amino acid and carbohydrate tradeoffs by honey bee nectar foragers and their implications for plant—pollinator interactions. Insect Physiol.
Herrera, C. Yeasts in floral nectar: a quantitative survey. Hoeksma, J. Bronstein Oxford: Oxford University Press , — Hojo, M. Lycaenid caterpillar secretions manipulate attendant ant behavior.
The Ants. Cambridge: Harvard University Press. Hughes, D. Hughes, J. Brodeur, and F. Thomas Oxford: Oxford University Press , — Inouye, D. Bentley and E.
Irwin, R. Nectar robbing: ecological and evolutionary perspectives. Mechanisms and evolution of deceptive pollination in orchids. Johnson, S. Batesian mimicry in the non-rewarding orchid Disa pulchra, and its consequences for pollinator behavior. Keasar, T. Variability in nectar production and standing crop, and their relation to pollinator visits in a Mediterranean shrub. Plant Interact. Keeler, K.
The extrafloral nectaries of Ipomoea carnea Convolvulaceae. Kessler, D. Making sense of nectar scents: the effects of nectar secondary metabolites on floral visitors of Nicotiana attenuata. Plant J. Unpredictability of nectar nicotine promotes outcrossing by hummingbirds in Nicotiana attenuata. Honeybees and nectar nicotine: deterrence and reduced survival versus potential health benefits. Koptur, S. Nectar secretion on fern fronds associated with lower levels of herbivore damage: field experiments with a widespread epiphyte of Mexican cloud forest remnants.
Labandeira, C. Lange, D. Variation in extrafloral nectary productivity influences the ant foraging. PLoS One e Influence of extrafloral nectary phenology on ant-plant mutualistic networks in a neotropical savanna. Leadbeater, E. Social transmission of nectar-robbing behaviour in bumble-bees.
B , — Leiss, K. Spatial distribution of nectar production in a natural Echium vulgare population: implications for pollinator behaviour. Lenaerts, M. Impact of microbial communities on floral nectar chemistry: potential implications for biological control of pest insects. Basic Appl. Maloof, J. Are nectar robbers cheaters or mutualists? Ecology 81, — Manson, J. Dose-dependent effects of nectar alkaloids in a montane plant-pollinator community.
Marazzi, B. The diversity, ecology and evolution of extrafloral nectaries: current perspectives and future challenges. Menzel, F. Parabiotic ants: the costs and benefits of symbiosis. Amino acids in nectar enhance butterfly fecundity: a long-awaited link.
Nepi, M. Beyond nectar sweetness: the hidden ecological role of non-protein amino acids in nectar. New perspective in nectar evolution and ecology: simple alimentary reward or complex multiorganism interaction? Acta Agrobot. Phylogenetic and functional signals in gymnosperm ovular secretions. The complexity of nectar: secretion and resorption dynamically regulate nectar features. Naturwissenschaften 95, — Nectar and pollination drops: how different are they? Ness, J. Catalpa bignonioides alters extrafloral nectar production after herbivory and attracts ant bodyguards.
Lach, C. Parr, and K. For ant-protected plants, the best defense is a hungry offense. Ecology 90, — Nicolson, S. Pacini Dordrecht: Springer , — Extraflorale Zuckerausscheidungen und Ameisenschutz. Ollerton, J. Pollinator diversity: distribution, ecological function, and conservation. How many flowering plants are pollinated by animals? Oikos , — Pacini, E. Palmer, T. Synergy of multiple partners, including freeloaders, increases host fitness in a multispecies mutualism.
Peakall, R. The significance of ant and plant traits for ant pollination in Leporella fimbriata. Oecologia 84, — Petanidou, T. Pozo, M. Pyke, G. Optimal foraging theory: a critical review. What does it cost a plant to produce floral nectar? Nature , 58— Floral nectar: pollinator attraction or manipulation?
Trends Ecol. Raguso, R. Why are some floral nectars scented? Ecology 85, — Floral scent in a whole-plant context: moving beyond pollinator attraction. Real, L. Individual variation in nectar production and its effect on fitness in Kalmia latifolia. Ecology 72, — Rico-Gray, V. Extrafloral nectar from cotton Gossypium hirsutum as a food source for parasitic wasps. Roy, R. Nectar biology: from molecules to ecosystems. Rusman, Q. Dealing with mutualists and antagonists: specificity of plant-mediated interactions between herbivores and flower visitors, and consequences for plant fitness.
Sachs, J. Bronstein Oxford: Oxford University Press , 93— Sampson, B. Nectar robbery by bees Xylocopa virginica and Apis mellifera contributes to the pollination of rabbiteye blueberry.
Santos, G. The seasonal dynamic of ant-flower networks in a semi-arid tropical environment. Sanz-Veiga, P. Pericarpial nectary-visiting ants do not provide fruit protection against pre-dispersal seed predators regardless of ant species composition and resource availability. Scheiner, R. Sucrose responsiveness and behavioral plasticity in honey bees Apis mellifera. Apidologie 35, — Schoonhoven, L. Insect—Plant Biology. Oxford: Oxford University Press. Schuettpelz, E.
Ranker and C. Haufler Cambridge: Cambridge University Press , — Simcock, N. Appetitive olfactory learning and memory in the honeybee depend on sugar reward identity. Single amino acids in sucrose rewards modulate feeding and associative learning in the honeybee.
Singh, V. Nectar robbing positively influences the reproductive success of Tecomella undulata Bignoniaceae.
0コメント