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Nectar is a
sugar Sugar is the generic name for sweet-tasting, soluble carbohydrates, many of which are used in food. Simple sugars, also called monosaccharides, include glucose, fructose, and galactose. Compound sugars, also called disaccharides or double ...
-rich liquid produced by plants in glands called nectaries or nectarines, either within the
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...
s with which it attracts
pollinating Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, a ...
animals, or by extrafloral nectaries, which provide a nutrient source to animal
mutualists Mutualism describes the ecological Biological interaction, interaction between two or more species where each species has a net benefit. Mutualism is a common type of ecological interaction. Prominent examples include most vascular plants engag ...
, which in turn provide herbivore protection. Common nectar-consuming
pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are the maj ...
s include
mosquito Mosquitoes (or mosquitos) are members of a group of almost 3,600 species of small flies within the family Culicidae (from the Latin ''culex'' meaning " gnat"). The word "mosquito" (formed by ''mosca'' and diminutive ''-ito'') is Spanish for "li ...
es,
hoverflies Hover flies, also called flower flies or syrphid flies, make up the insect family Syrphidae. As their common name suggests, they are often seen hovering or nectaring at flowers; the adults of many species feed mainly on nectar and pollen, while ...
,
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. Th ...
s, bees,
butterflies Butterflies are insects in the macrolepidopteran clade Rhopalocera from the Order (biology), order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The ...
and
moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...
s,
hummingbird Hummingbirds are birds native to the Americas and comprise the biological family Trochilidae. With about 361 species and 113 genera, they occur from Alaska to Tierra del Fuego, but the vast majority of the species are found in the tropics aro ...
s,
honeyeater The honeyeaters are a large and diverse family (biology), family, Meliphagidae, of small to medium-sized birds. The family includes the Epthianura, Australian chats, myzomelas, friarbirds, wattlebirds, Manorina, miners and melidectes. They are ...
s and
bats Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bi ...
. Nectar plays a crucial role in the foraging economics and evolution of nectar-eating species; for example, nectar foraging behavior is largely responsible for the divergent evolution of the African honey bee, ''A. m. scutellata'' and the
western honey bee The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for "bee", and ''mellifera'' is the Latin for "honey-bearing" or "honey carrying", ...
. Nectar is an economically important substance as it is the sugar source for
honey Honey is a sweet and viscous substance made by several bees, the best-known of which are honey bees. Honey is made and stored to nourish bee colonies. Bees produce honey by gathering and then refining the sugary secretions of plants (primar ...
. It is also useful in
agriculture Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people to ...
and
horticulture Horticulture is the branch of agriculture that deals with the art, science, technology, and business of plant cultivation. It includes the cultivation of fruits, vegetables, nuts, seeds, herbs, sprouts, mushrooms, algae, flowers, seaweeds and no ...
because the adult stages of some predatory insects feed on nectar. For example, a number of
parasitoid wasps Parasitoid wasps are a large group of hymenopteran superfamilies, with all but the wood wasps (Orussoidea) being in the wasp-waisted Apocrita. As parasitoids, they lay their eggs on or in the bodies of other arthropods, sooner or later causi ...
(e.g. the social wasp species ''
Apoica flavissima ''Apoica flavissima'' is a paper wasp found primarily in South America. The species is distinguishable by its light coloring, unique single comb nests, and nocturnal nature. A notable feature of this species is the size dimorphism between queens ...
'') rely on nectar as a primary food source. In turn, these wasps then hunt agricultural pest insects as food for their young.


Etymology

''Nectar'' is derived from
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
''νεκταρ'', 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 nectarine is floral tissue found in different locations in the flower, and is one of several secretory floral structures, including elaiophores and osmophores, producing nectar, oil and scent respectively. The function of these structures is to attract potential
pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are the maj ...
s, which may include insects, including bees and
moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...
s, and vertebrates such as
hummingbird Hummingbirds are birds native to the Americas and comprise the biological family Trochilidae. With about 361 species and 113 genera, they occur from Alaska to Tierra del Fuego, but the vast majority of the species are found in the tropics aro ...
s and
bat Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most ...
s. 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) *
hypanthium In angiosperms, a hypanthium or floral cup is a structure where basal portions of the calyx, the corolla, and the stamens form a cup-shaped tube. It is sometimes called a floral tube, a term that is also used for corolla tube and calyx tube. It ...
(hypanthial) *
tepals A tepal is one of the outer parts of a flower (collectively the perianth). The term is used when these parts cannot easily be classified as either sepals or petals. This may be because the parts of the perianth are undifferentiated (i.e. of very ...
(perigonal, tepal) *
sepals A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 The term ''sepalum'' was coined b ...
(sepal) *
petal Petals are modified Leaf, leaves that surround the reproductive parts of flowers. They are often advertising coloration, brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''c ...
(petal, corolla) *
stamen The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...
(staminal, androecial: filament, anther, staminodal) *
pistil Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ''pistils'' ...
(gynoecial: stigmatic, stylar) ** pistillodes (pistillodal, carpellodial) **
ovaries The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. T ...
(ovarian: non-septal, septal, gynopleural) Most members of
Lamiaceae The Lamiaceae ( ) or Labiatae are a family of flowering plants commonly known as the mint, deadnettle or sage family. Many of the plants are aromatic in all parts and include widely used culinary herbs like basil, mint, rosemary, sage, savory ...
have a nectariferous disc which surrounds the ovary base and derived from developing ovarian tissue. In most
Brassicaceae Brassicaceae () or (the older) Cruciferae () is a medium-sized and economically important family of flowering plants commonly known as the mustards, the crucifers, or the cabbage family. Most are herbaceous plants, while some are shrubs. The le ...
the nectary is at the base of the stamen filament. Many
monocotyledons Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of t ...
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
coevolved In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...
depending on the pollinator that feeds on the plant's nectar. Nectar is secreted from
epidermal The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water relea ...
cells of the nectaries, which have a dense
cytoplasm In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. The ...
, by means of
trichomes Trichomes (); ) are fine outgrowths or appendages on plants, algae, lichens, and certain protists. They are of diverse structure and function. Examples are hairs, glandular hairs, scales, and papillae. A covering of any kind of hair on a plant ...
or modified
stomata In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is bor ...
. Adjacent vascular tissue conducts
phloem Phloem (, ) is the living biological tissue, tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This tran ...
bringing
sugars Sugar is the generic name for sweet-tasting, soluble carbohydrates, many of which are used in food. Simple sugars, also called monosaccharides, include glucose, fructose, and galactose. Compound sugars, also called disaccharides or double s ...
to the secretory region, where it is secreted from the cells through
vesicle Vesicle may refer to: ; In cellular biology or chemistry * Vesicle (biology and chemistry), a supramolecular assembly of lipid molecules, like a cell membrane * Synaptic vesicle ; In human embryology * Vesicle (embryology), bulge-like features o ...
s packaged by the
endoplasmic reticulum The endoplasmic reticulum (ER) is, in essence, the transportation system of the eukaryotic cell, and has many other important functions such as protein folding. It is a type of organelle made up of two subunits – rough endoplasmic reticulum ( ...
. The adjacent subepidermal cells may also be secretory. Flowers that have longer nectaries sometimes have a
vascular The blood vessels are the components of the circulatory system that transport blood throughout the human body. These vessels transport blood cells, nutrients, and oxygen to the tissues of the body. They also take waste and carbon dioxide away f ...
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
fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Proce ...
and
outcrossing Out-crossing or out-breeding is the technique of crossing between different breeds. This is the practice of introducing distantly related genetic material into a breeding line, thereby increasing genetic diversity. Outcrossing can be a useful ...
of the plant as they brush against the reproductive organs, the
stamen The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...
and
pistil Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ''pistils'' ...
, of the plant and pick up or deposit
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
. Nectar from floral nectaries is sometimes used as a reward to insects, such as
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
s, that protect the plant from predators. Many floral families have evolved a nectar
spur A spur is a metal tool designed to be worn in pairs on the heels of riding boots for the purpose of directing a horse or other animal to move forward or laterally while riding. It is usually used to refine the riding aids (commands) and to back ...
. 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
herbivory A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
is often one of the roles of extrafloral nectaries. Floral nectaries can also be involved in defense. In addition to the
sugars Sugar is the generic name for sweet-tasting, soluble carbohydrates, many of which are used in food. Simple sugars, also called monosaccharides, include glucose, fructose, and galactose. Compound sugars, also called disaccharides or double s ...
found in nectar, certain
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
s may also be found in nectar secreted by floral nectaries. In
tobacco Tobacco is the common name of several plants in the genus '' Nicotiana'' of the family Solanaceae, and the general term for any product prepared from the cured leaves of these plants. More than 70 species of tobacco are known, but the ...
plants, these proteins have
antimicrobial An antimicrobial is an agent that kills microorganisms or stops their growth. Antimicrobial medicines can be grouped according to the microorganisms they act primarily against. For example, antibiotics are used against bacteria, and antifungals ar ...
and
antifungal An antifungal medication, also known as an antimycotic medication, is a pharmaceutical fungicide or fungistatic used to treat and prevent mycosis such as athlete's foot, ringworm, candidiasis (thrush), serious systemic infections such as crypto ...
properties and can be secreted to defend the
gynoecium Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ''pistils'' ...
from certain pathogens. Floral nectaries have
evolved Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation t ...
and diverged into the different types of nectaries due to the various pollinators that visit the flowers. In
Melastomataceae Melastomataceae is a family of dicotyledonous flowering plants found mostly in the tropics (two-thirds of the genera are from the New World tropics) comprising c. 175 genera and c. 5115 known species. Melastomes are annual or perennial herbs, sh ...
, 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 bees. Instead they focused on energy allocation to
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
production. Species of
angiosperms Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
that have nectaries use the nectar to attract pollinators that consume the nectar, such as
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
s and
butterflies Butterflies are insects in the macrolepidopteran clade Rhopalocera from the Order (biology), order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The ...
. In
Bromeliaceae The Bromeliaceae (the bromeliads) are a family of monocot Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain o ...
, 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 to provide a reward for. In flowers that are generally pollinated by a long-tongued organism such as certain
flies Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
,
moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...
s, 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
pollination Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds ...
, generally on the
leaf A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...
or petiole (foliar nectaries) and often in relation to the
leaf venation A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ...
. They are highly diverse in form, location, size, and mechanism. They have been described in virtually all above-ground plant parts—including
stipule In botany, a stipule is an outgrowth typically borne on both sides (sometimes on just one side) of the base of a leafstalk (the petiole). Stipules are considered part of the anatomy of the leaf of a typical flowering plant, although in many speci ...
s,
cotyledon A cotyledon (; ; ; , gen. (), ) is a significant part of the embryo within the seed of a plant, and is defined as "the embryonic leaf in seed-bearing plants, one or more of which are the first to appear from a germinating seed." The numb ...
s,
fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
s, and
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
s, among others. They range from single-celled
trichome Trichomes (); ) are fine outgrowths or appendages on plants, algae, lichens, and certain protists. They are of diverse structure and function. Examples are hairs, glandular hairs, scales, and papillae. A covering of any kind of hair on a pla ...
s to complex cup-like structures that may or may not be
vascularized Angiogenesis is the physiological process through which new blood vessels form from pre-existing vessels, formed in the earlier stage of vasculogenesis. Angiogenesis continues the growth of the vasculature by processes of sprouting and splitting ...
. Like floral nectaries, they consist of groups of glandular trichomes (e.g. ''
Hibiscus ''Hibiscus'' is a genus of flowering plants in the mallow family, Malvaceae. The genus is quite large, comprising several hundred species that are native to warm temperate, subtropical and tropical regions throughout the world. Member species ...
'' spp.) or elongated secretory epidermal cells. The latter are often associated with underlying vascular tissue. They may be associated with specialised pockets (
domatia A domatium (plural: domatia, from the Latin "domus", meaning home) is a tiny chamber that houses arthropods, produced by a plant. Ideally domatia differ from galls in that they are produced by the plant rather than being induced by their inhabi ...
), pits or raised regions (e.g.
Euphorbiaceae Euphorbiaceae, the spurge family, is a large family of flowering plants. In English, they are also commonly called euphorbias, which is also the name of a genus in the family. Most spurges, such as ''Euphorbia paralias'', are herbs, but some, e ...
). The leaves of some tropical
eudicots The eudicots, Eudicotidae, or eudicotyledons are a clade of flowering plants mainly characterized by having two seed leaves upon germination. The term derives from Dicotyledons. Traditionally they were called tricolpates or non-magnoliid dico ...
(e.g.
Fabaceae The Fabaceae or Leguminosae,International Code of Nomenc ...
) and
magnoliids Magnoliids (or Magnoliidae or Magnolianae) are a clade of flowering plants. With more than 10,000 species, including magnolias, nutmeg, bay laurel, cinnamon, avocado, black pepper, tulip tree and many others, it is the third-largest group of angio ...
(e.g.
Piperaceae The Piperaceae (), also known as the pepper family, are a large family of flowering plants. The group contains roughly 3,600 currently accepted species in 5 genera. The vast majority of species can be found within the two main genera: ''Piper'' ( ...
) 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 have a defensive function. The nectar attracts predatory insects which will eat both the nectar and any plant-eating insects around, thus functioning as 'bodyguards'. Foraging predatory insects show a preference for plants with extrafloral nectaries, particularly some species of
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
s and
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. Th ...
s, which have been observed to defend the plants bearing them. ''
Acacia ''Acacia'', commonly known as the wattles or acacias, is a large genus of shrubs and trees in the subfamily Mimosoideae of the pea family Fabaceae. Initially, it comprised a group of plant species native to Africa and Australasia. The genus na ...
'' is one example of a plant whose nectaries attract ants, which protect the plant from other insect
herbivores A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
. Among
passion flowers ''Passiflora'', known also as the passion flowers or passion vines, is a genus of about 550 species of flowering plants, the type genus of the family (biology), family Passifloraceae. They are mostly tendril, tendril-bearing vines, with some b ...
, for example, extrafloral nectaries prevent herbivores by attracting ants and deterring two species of butterflies from laying eggs. In many
carnivorous plant Carnivorous plants are plants that derive some or most of their nutrients from trapping and consuming animals or protozoans Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryot ...
s, extrafloral nectaries are also used to attract insect
prey Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
. Darwin 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
excretory Excretion is a process in which metabolic waste is eliminated from an organism. In vertebrates this is primarily carried out by the lungs, kidneys, and skin. This is in contrast with secretion, where the substance may have specific tasks aft ...
in nature (
hydathode A hydathode is a type of pore, commonly found in angiosperms, that secretes water through pores in the epidermis or leaf margin, typically at the tip of a marginal tooth or serration. Hydathodes occur in the leaves of submerged aquatic plants s ...
s). Their defensive functions were first recognized by the
Italian Italian(s) may refer to: * Anything of, from, or related to the people of Italy over the centuries ** Italians, an ethnic group or simply a citizen of the Italian Republic or Italian Kingdom ** Italian language, a Romance language *** Regional Ita ...
botanist Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek wo ...
Federico Delpino Giacomo Giuseppe Federico Delpino (27 December 1833 – 14 May 1905) was an Italian botanist who made early observations on floral biology, particularly the pollination of flowers by insects. Delpino introduced a very broad view of plant ecolog ...
in his important
monograph A monograph is a specialist work of writing (in contrast to reference works) or exhibition on a single subject or an aspect of a subject, often by a single author or artist, and usually on a scholarly subject. In library cataloging, ''monograph ...
''Funzione mirmecofila nel regno vegetale'' (1886). Delpino's study was inspired by a disagreement with
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended fr ...
, with whom he corresponded regularly. Extrafloral nectaries have been reported in over 3941 species of
vascular plant Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They al ...
s belonging to 745
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
and 108
families Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideall ...
, 99.7% of which belong to
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s (angiosperms), comprising 1.0 to 1.8% of all known species. They are most common among
eudicot The eudicots, Eudicotidae, or eudicotyledons are a clade of flowering plants mainly characterized by having two seed leaves upon germination. The term derives from Dicotyledons. Traditionally they were called tricolpates or non-magnoliid dicot ...
s, occurring in 3642 species (of 654 genera and 89 families), particularly among
rosids The rosids are members of a large clade (monophyletic group) of flowering plants, containing about 70,000 species, more than a quarter of all angiosperms. The clade is divided into 16 to 20 orders, depending upon circumscription and classificatio ...
which comprise more than half of the known occurrences. The families showing the most recorded occurrences of extrafloral nectaries are
Fabaceae The Fabaceae or Leguminosae,International Code of Nomenc ...
, with 1069 species,
Passifloraceae The Passifloraceae are a family of flowering plants, containing about 750 species classified in around 27 genera. They include trees, shrubs, lianas, and climbing plants, and are mostly found in tropical regions. The family takes its name from t ...
, with 438 species, and
Malvaceae Malvaceae, or the mallows, is a family of flowering plants estimated to contain 244 genera with 4225 known species. Well-known members of economic importance include okra, cotton, cacao and durian. There are also some genera containing familiar ...
, with 301 species. The genera with the most recorded occurrences are ''
Passiflora ''Passiflora'', known also as the passion flowers or passion vines, is a genus of about 550 species of flowering plants, the type genus of the family Passifloraceae. They are mostly tendril-bearing vines, with some being shrubs or trees. They ...
'' (322 species, Passifloraceae), ''
Inga ''Inga'' is a genus of small tropical, tough-leaved, nitrogen-fixing treesElkan, Daniel. "Slash-and-burn farming has become a major threat to the world's rainforest" ''The Guardian'' 21 April 2004 and shrubs, subfamily Mimosoideae. ''Inga''s l ...
'' (294 species, Fabaceae), and ''
Acacia ''Acacia'', commonly known as the wattles or acacias, is a large genus of shrubs and trees in the subfamily Mimosoideae of the pea family Fabaceae. Initially, it comprised a group of plant species native to Africa and Australasia. The genus na ...
'' (204 species, Fabaceae). Other genera with extrafloral nectaries include ''
Salix Willows, also called sallows and osiers, from the genus ''Salix'', comprise around 400 speciesMabberley, D.J. 1997. The Plant Book, Cambridge University Press #2: Cambridge. of typically deciduous trees and shrubs, found primarily on moist so ...
'' (
Salicaceae The Salicaceae is the willow family of flowering plants. The traditional family (Salicaceae ''sensu stricto'') included the willows, poplar, aspen, and cottonwoods. Genetic studies summarized by the Angiosperm Phylogeny Group (APG) have greatly ...
), ''
Prunus ''Prunus'' is a genus of trees and shrubs, which includes (among many others) the fruits plums, cherries, peaches, nectarines, apricots, and almonds. Native to the North American temperate regions, the neotropics of South America, and the p ...
'' (
Rosaceae Rosaceae (), the rose family, is a medium-sized family of flowering plants that includes 4,828 known species in 91 genera. The name is derived from the type genus ''Rosa''. Among the most species-rich genera are ''Alchemilla'' (270), ''Sorbus ...
) and ''
Gossypium ''Gossypium'' () is a genus of flowering plants in the tribe Gossypieae of the mallow family, Malvaceae, from which cotton is harvested. It is native to tropical and subtropical regions of the Old and New Worlds. There are about 50 ''Gossypiu ...
'' (
Malvaceae Malvaceae, or the mallows, is a family of flowering plants estimated to contain 244 genera with 4225 known species. Well-known members of economic importance include okra, cotton, cacao and durian. There are also some genera containing familiar ...
). Foliar nectaries have also been observed in 39 species of
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except t ...
s belonging to seven genera and four families of
Cyatheales The order Cyatheales, which includes most tree ferns, is a taxonomic order of the fern class, Polypodiopsida. No clear morphological features characterize all of the Cyatheales, but DNA sequence data indicate the order is monophyletic. Some sp ...
and
Polypodiales The order Polypodiales encompasses the major lineages of polypod ferns, which comprise more than 80% of today's fern species. They are found in many parts of the world including tropical, semitropical and temperate areas. Description Polypodiale ...
. They are absent, however, in
bryophyte The Bryophyta s.l. are a proposed taxonomic division containing three groups of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. Bryophyta s.s. consists of the mosses only. They are characteristically limited in ...
s,
gymnosperm The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, ''Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμνό ...
s, early
angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s,
magnoliids Magnoliids (or Magnoliidae or Magnolianae) are a clade of flowering plants. With more than 10,000 species, including magnolias, nutmeg, bay laurel, cinnamon, avocado, black pepper, tulip tree and many others, it is the third-largest group of angio ...
, and members of
Apiales The Apiales are an order of flowering plants. The families are those recognized in the APG III system. This is typical of the newer classifications, though there is some slight variation and in particular, the Torriceliaceae may also be divide ...
among the eudicots.
Phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
studies and the wide distribution of extrafloral nectaries among vascular plants point to multiple independent
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
ary origins of extrafloral nectaries in at least 457 independent lineages.


Components

The main ingredients in nectar are sugars in varying proportions of
sucrose Sucrose, a disaccharide, is a sugar composed of glucose and fructose subunits. It is produced naturally in plants and is the main constituent of white sugar. It has the molecular formula . For human consumption, sucrose is extracted and refined ...
,
glucose Glucose is a simple sugar with the molecular formula . Glucose is overall the most abundant monosaccharide, a subcategory of carbohydrates. Glucose is mainly made by plants and most algae during photosynthesis from water and carbon dioxide, using ...
, and
fructose Fructose, or fruit sugar, is a Ketose, ketonic monosaccharide, simple sugar found in many plants, where it is often bonded to glucose to form the disaccharide sucrose. It is one of the three dietary monosaccharides, along with glucose and galacto ...
. In addition, nectars have diverse other
phytochemical Phytochemicals are chemical compounds produced by plants, generally to help them resist fungi, bacteria and plant virus infections, and also consumption by insects and other animals. The name comes . Some phytochemicals have been used as poisons ...
s serving to both attract pollinators and discourage predators.
Carbohydrate In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or ma ...
s,
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha am ...
s, and
volatiles Volatiles are the group of chemical elements and chemical compounds that can be readily vaporized. In contrast with volatiles, elements and compounds that are not readily vaporized are known as refractory substances. On planet Earth, the term ' ...
function to attract some species, whereas
alkaloid Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of similar ...
s and
polyphenol Polyphenols () are a large family of naturally occurring organic compounds characterized by multiples of phenol units. They are abundant in plants and structurally diverse. Polyphenols include flavonoids, tannic acid, and ellagitannin, some of ...
s appear to provide a protective function. The ''Nicotiana attenuata'', a tobacco plant native to the US state of
Utah Utah ( , ) is a state in the Mountain West subregion of the Western United States. Utah is a landlocked U.S. state bordered to its east by Colorado, to its northeast by Wyoming, to its north by Idaho, to its south by Arizona, and to it ...
, uses several volatile aromas to attract pollinating birds and moths. The strongest such aroma is benzylacetone, but the plant also adds bitter
nicotine Nicotine is a natural product, naturally produced alkaloid in the nightshade family of plants (most predominantly in tobacco and ''Duboisia hopwoodii'') and is widely used recreational drug use, recreationally as a stimulant and anxiolytic. As ...
, 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. Neurotoxins such as
aesculin Aesculin, also called æsculin or esculin, is a coumarin glucoside that naturally occurs in the trees horse chestnut (''Aesculus hippocastanum''), California buckeye (''Aesculus californica''), prickly box (''Bursaria spinosa''), and daphnin ( ...
are present in some nectars such as that of the
California buckeye ''Aesculus californica'', commonly known as the California buckeye or California horse-chestnut, is a species of buckeye native to California and southwestern Oregon. Description It is a large deciduous shrub or small tree, up to tall, with ...
.C.Michael Hogan (2008) ''Aesculus californica'', Globaltwitcher.com, ed. N. Stromberg
/ref> Nectar contains water,
carbohydrate In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or ma ...
s,
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha am ...
s,
ion An ion () is an atom or molecule with a net electrical charge. The charge of an electron is considered to be negative by convention and this charge is equal and opposite to the charge of a proton, which is considered to be positive by conve ...
s and numerous other compounds.


Other floral secretory structures

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
orchids Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant. Along with the Asteraceae, they are one of the two largest families of flowering ...
these have
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
qualities. Osmophores have thick domed or
papillate This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary o ...
epidermis and dense cytoplasm. ''
Platanthera bifolia ''Platanthera bifolia'', commonly known as the lesser butterfly-orchid, is a species of orchid in the genus ''Platanthera'', having certain relations with the genus ''Orchis'', where it was previously included and also with the genus ''Habenari ...
'' produces a nocturnal scent from the labellum epidermis. ''
Ophrys The genus ''Ophrys'' is a large group of orchids from the alliance Orchis in the subtribe Orchidinae. They are widespread across much of Europe, North Africa, Caucasus, the Canary Islands, and the Middle East as far east as Turkmenistan. These p ...
'' labella have dome-shaped, papillate, dark-staining epidermal cells forming osmophores. '' Narcissus'' emit pollinator specific volatiles from the
corona Corona (from the Latin for 'crown') most commonly refers to: * Stellar corona, the outer atmosphere of the Sun or another star * Corona (beer), a Mexican beer * Corona, informal term for the coronavirus SARS-CoV-2, which causes the COVID-19 di ...
.


See also

*
Nectar guide Nectar guides are markings or patterns seen in flowers of some angiosperm species, that guide pollinators to their rewards. Rewards commonly take the form of nectar, pollen, or both, but various plants produce oil, resins, scents, or waxes. Such ...
*
Nectar source A nectar source is a flowering plant that produces nectar as part of its reproductive strategy. These plants create nectar, which attract pollinating insects and sometimes other animals such as birds. Nectar source plants are important for beekee ...
*
Nectarivore In zoology, a nectarivore is an animal which derives its energy and nutrient requirements from a diet consisting mainly or exclusively of the sugar-rich nectar produced by flowering plants. Nectar as a food source presents a number of benefits a ...
*
Northern American nectar sources for honey bees The nectar resource in a given area depends on the kinds of flowering plants present and their blooming periods. Which kinds grow in an area depends on soil texture, soil pH, soil drainage, daily maximum and minimum temperatures, precipitation, ...


References


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. * * Roshchina, V.V. and Roshchina, V.D. (1993) The excretory function of higher plants. Springer-Verlag, Berlin. * ;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. * * * * * Ferreres, F., Andrade, P., Gil, M.I. and Tomas Barberan, F.A. (1996) Floral nectar phenolics as biochemical markers for the botanical origin of heather honey. Zeitschrift für Lebensmittel Untersuchung und Forschung. 202:40–44. * 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. ;Websites *


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