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Nectar is a , -rich produced by in glands called nectaries, either within the with which it attracts animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists, which in turn provide herbivore protection. Common nectar-consuming include , , , , and , , and bats. Nectar is an economically important substance as it is the sugar source for . It is also useful in and because the adult stages of some predatory insects feed on nectar. For example, a number of predacious or (e.g., the social wasp species Apoica flavissima) rely on nectar as a primary food source. In turn, these wasps then hunt agricultural pest insects as food for their young.

Nectar is most often associated with flowering plants , but it is also produced by other groups, including .


Etymology
Nectar is derived from νέκταρ, the fabled drink of eternal life. Some derive the word from νε- or νη- "not" plus κτα- or κτεν- "kill", meaning "unkillable", thus "immortal". The common use of the word "nectar" to refer to the "sweet liquid in flowers", is first recorded in AD 1600.


Floral nectaries
A nectary or honey gland is floral tissue found in different locations in the flower and is one of several secretory floral structures, including and osmophores, producing nectar, oil and scent respectively. The function of these structures is to attract potential , which may include insects, including and , and vertebrates such as and . Nectaries can occur on any floral part, but they may also represent a modified part or a novel structure. The different types of floral nectaries include:
  • receptacle (receptacular: extrastaminal, intrastaminal, interstaminal)
  • (hypanthial)
  • (perigonal, tepal)
  • (sepal)
  • (petal, corolla)
  • (staminal, androecial: filament, anther, staminodal)
  • (gynoecial: stigmatic, stylar)
    • pistillodes (pistillodal, carpellodial)
    • ovaries (ovarian: non-septal, septal, gynopleural)
Most members of have a nectariferous disc which surrounds the ovary base and derived from developing ovarian tissue. In most , the nectary is at the base of the stamen filament. Many have septal nectaries, which are at the unfused margins of the carpels. These exude nectar from small pores on the surface of the gynoecium. Nectaries may also vary in color, number, and symmetry.Willmer, Pat. Pollination and floral ecology. Princeton University Press, 2011. Nectaries can also be categorized as structural or non-structural. Structural nectaries refer to specific areas of tissue that exude nectar, such as the types of floral nectaries previously listed. Non-structural nectaries secrete nectar infrequently from non-differentiated tissues. The different types of floral nectaries depending on the pollinator that feeds on the plant's nectar. Nectar is secreted from epidermal cells of the nectaries, which have a dense , by means of or modified . Adjacent vascular tissue conducts bringing to the secretory region, where it is secreted from the cells through vesicles packaged by the endoplasmic reticulum. The adjacent subepidermal cells may also be secretory. Flowers that have longer nectaries sometimes have a strand in the nectary to assist in transport over a longer distance.

Pollinators feed on the nectar and depending on the location of the nectary the pollinator assists in and of the plant as they brush against the reproductive organs, the and , of the plant and pick up or deposit . Nectar from floral nectaries is sometimes used as a reward to insects, such as , 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. Different characteristics of the spur, such as its length or position in the flower, may determine the type of pollinator that visits the flower.

Defense from is often one of the roles of extrafloral nectaries. Floral nectaries can also be involved in defense. In addition to the found in nectar, certain may also be found in nectar secreted by floral nectaries. In plants, these proteins have and properties and can be secreted to defend the from certain pathogens.

Floral nectaries have and diverged into the different types of nectaries due to the various pollinators that visit the flowers. In , different types of floral nectaries have evolved and been lost many times. Flowers that ancestrally produced nectar and had nectaries may have lost their ability to produce nectar due to a lack of nectar consumption by pollinators, such as certain species of . Instead they focused on energy allocation to production. Species of that have nectaries use the nectar to attract pollinators that consume the nectar, such as and . In , septal nectaries (a form of gynoecial nectary) are common in species that are insect or bird pollinated. In species that are wind pollinated, nectaries are often absent because there is no pollinator. In flowers that are generally pollinated by a long-tongued organism such as certain , , butterflies, and birds, nectaries in the ovaries are common because they are able to reach the nectar reward when pollinating. Sepal and petal nectaries are often more common in species that are pollinated by short-tongued insects that cannot reach so far into the flower.


Secretion
Nectar secretion increases as the flower is visited by pollinators. After pollination, the nectar is frequently reabsorbed into the plant. The amount of nectar in flowers at any given time is variable due to many factors, including flower age, plant location, and habitat management.


Extrafloral nectaries
Extrafloral nectaries (also known as extranuptial nectaries) are specialised nectar-secreting plant glands that develop outside of flowers and are not involved in , generally on the or petiole (foliar nectaries) and often in relation to the . They are highly diverse in form, location, size, and mechanism. They have been described in virtually all above-ground plant parts—including , , , and , among others. They range from single-celled to complex cup-like structures that may or may not be . Like floral nectaries, they consist of groups of glandular trichomes (e.g., spp.) or elongated secretory epidermal cells. The latter are often associated with underlying vascular tissue. They may be associated with specialised pockets (), pits or raised regions (e.g., ). The leaves of some tropical (e.g., ) and (e.g., ) possess pearl glands or bodies which are globular trichomes specialised to attract ants. They secrete matter that is particularly rich in carbohydrates, proteins and lipids.

While their function is not always clear, and may be related to regulation of sugars, in most cases they appear to facilitate plant insect relationships. In contrast to floral nectaries, nectar produced outside the flower generally has a defensive function. The nectar attracts predatory insects which will eat both the nectar and any insects around, thus functioning as "bodyguards". Plant-Provided Food for Carnivorous Insects – Cambridge University Press Foraging predatory insects show a preference for plants with extrafloral nectaries, particularly some species of and , which have been observed to defend the plants bearing them. is one example of a plant whose nectaries attract ants, which protect the plant from other insect . Among , for example, extrafloral nectaries prevent herbivores by attracting ants and deterring two species of butterflies from laying eggs. In many carnivorous plants, extrafloral nectaries are also used to attract insect .Merbach, M. 2001. Nectaries in Nepenthes. In: C.M. Clarke Nepenthes of Sumatra and Peninsular Malaysia. Natural History Publications (Borneo), Kota Kinabalu.

understood that extrafloral nectar "though small in quantity, is greedily sought by insects" but believed that "their visits do not in any way benefit the plant". Instead, he believed that extrafloral nectaries were in nature (). Their defensive functions were first recognized by the in his important Funzione mirmecofila nel regno vegetale (1886). Delpino's study was inspired by a disagreement with Darwin, with whom he corresponded regularly.

Extrafloral nectaries have been reported in over 3941 species of belonging to 745 and 108 families, 99.7% of which belong to (angiosperms), comprising 1.0 to 1.8% of all known species. They are most common among , occurring in 3642 species (of 654 genera and 89 families), particularly among which comprise more than half of the known occurrences. The families showing the most recorded occurrences of extrafloral nectaries are , with 1069 species, , with 438 species, and , with 301 species. The genera with the most recorded occurrences are (322 species, Passifloraceae), (294 species, Fabaceae), and (204 species, Fabaceae). Other genera with extrafloral nectaries include (), () and ().

Foliar nectaries have also been observed in 101 species of belonging to eleven genera and six families, most of them belonging to (tree ferns) and . Fern nectaries appear to have evolved around 135 million years ago, nearly simultaneously with angiosperms. However, fern nectaries did not diversify remarkably until nearly 100 million years later, in the , with weak support for a role played by arthropod herbivore diversifications. They are absent in , , early , , and members of among the eudicots. studies and the wide distribution of extrafloral nectaries among vascular plants point to multiple independent origins of extrafloral nectaries in at least 457 independent lineages.


Components
The main ingredients in nectar are sugars in varying proportions of , , and . In addition, nectars have diverse other serving to both attract pollinators and discourage predators. , , and volatiles function to attract some species, whereas and appear to provide a protective function. The Nicotiana attenuata, a tobacco plant native to the US state of , uses several volatile aromas to attract pollinating birds and moths. The strongest such aroma is , but the plant also adds bitter , which is less aromatic, so may not be detected by the bird until after taking a drink. Researchers speculate the purpose of this addition is to discourage the forager after only a sip, motivating it to visit other plants, therefore maximizing the pollination efficiency gained by the plant for a minimum nectar output. such as are present in some nectars such as that of the California buckeye. C.Michael Hogan (2008) Aesculus californica, Globaltwitcher.com, ed. N. Stromberg Nectar contains water, , , , , and numerous other compounds.


Similar attractive substances
Some insect pollinated plants lack nectaries, but attract pollinators through other secretory structures. Elaiophores are similar to nectaries but are oil secreting. Osmophores are modified structural structures that produce volatile scents. In , these have qualities. Osmophores have thick domed or epidermis and dense cytoplasm. Platanthera bifolia produces a nocturnal scent from the labellum epidermis. labella have dome-shaped, papillate, dark-staining epidermal cells forming osmophores. Narcissus emit pollinator specific volatiles from the corona.


See also


Bibliography
Books
  • Baker, H.G. and Baker, I. (1975) Studies of nectar-constitution and pollinator-plant coevolution. In Coevolution of animals and plants. Gilbert, L.E. and Raven, P.H. ed. Univ. of Texas Press, Austin, 100–140.
  • Esau, K. (1977) Anatomy of seed plants. John Wiley & Sons, New York.
  • (2025). 9781402059377, Springer Publications. .
  • Roshchina, V.V. and Roshchina, V.D. (1993) The excretory function of higher plants. Springer-Verlag, Berlin.
  • (2025). 9781139459488, Cambridge University Press.
Articles
  • Baker, H.G. and Baker, I. (1981) Chemical constituents of nectar in relation to pollination mechanisms and phylogeny. In Biochemical aspects of evolutionary biology. 131–171.
  • Beutler, R. (1935) Nectar. Bee World 24:106–116, 128–136, 156–162.
  • Frey-Wyssling, A. (1955) The phloem supply to the nectaries. Acta Bot. Neerl. 4:358–369.
  • Vogel, S. (1969) Flowers offering fatty oil instead of nectar. Abstracts XIth Internatl. Bot. Congr. Seattle.
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