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The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10


Morphology and terminology

A stamen typically consists of a stalk called the filament and an anther which contains microsporangia. Most commonly, anthers are two-lobed (each lobe is termed a locule) and are attached to the filament either at the base or in the middle area of the anther. The sterile (i.e. nonreproductive) tissue between the lobes is called the connective, an extension of the filament containing conducting strands. It can be seen as an extension on the dorsal side of the anther. A
pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
grain develops from a
microspore Microspores are land plant spores that develop into male gametophytes, whereas megaspores develop into female gametophytes. The male gametophyte gives rise to sperm cells, which are used for fertilization of an egg cell to form a zygote. Megaspo ...
in the microsporangium and contains the male
gametophyte A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the se ...
. The size of anthers differs greatly, from a tiny fraction of a millimeter in ''Wolfia'' spp up to five inches (13 centimeters) in '' Canna iridiflora'' and '' Strelitzia nicolai''. The stamens in a flower are collectively called the androecium. The androecium can consist of as few as one-half stamen (i.e. a single locule) as in '' Canna'' species or as many as 3,482 stamens which have been counted in the saguaro cactus (''Carnegiea gigantea''). The androecium in various species of plants forms a great variety of patterns, some of them highly complex. It generally surrounds the
gynoecium Gynoecium (; ; : gynoecia) 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 (botany), whorl of a flower; it consists ...
and is surrounded by the
perianth The perianth (perigonium, perigon or perigone in monocots) is the non-reproductive part of the flower. It is a structure that forms an envelope surrounding the sexual organs, consisting of the calyx (sepals) and the corolla (petals) or tepal ...
. A few members of the family Triuridaceae, particularly '' Lacandonia schismatica'' and '' Lacandonia brasiliana'', along with a few species of '' Trithuria'' (family Hydatellaceae) are exceptional in that their gynoecia surround their androecia.


Etymology

* Stamen is the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
word meaning "thread" (originally thread of the warp, in weaving). * Filament derives from
classical Latin Classical Latin is the form of Literary Latin recognized as a Literary language, literary standard language, standard by writers of the late Roman Republic and early Roman Empire. It formed parallel to Vulgar Latin around 75 BC out of Old Latin ...
filum, meaning "thread"Lewis, C.T. & Short, C. (1879). ''A Latin dictionary founded on Andrews' edition of Freund's Latin dictionary.'' Oxford: Clarendon Press. * Anther derives from French anthère,Klein, E. (1971). ''A comprehensive etymological dictionary of the English language. Dealing with the origin of words and their sense development thus illustration the history of civilization and culture.'' Amsterdam: Elsevier Science B.V. from classical Latin anthera, meaning "medicine extracted from the flower"Siebenhaar, F.J. (1850). ''Terminologisches Wörterbuch der medicinischen Wissenschaften.'' (Zweite Auflage). Leipzig: Arnoldische Buchhandlung.Saalfeld, G.A.E.A. (1884). ''Tensaurus Italograecus. Ausführliches historisch-kritisches Wörterbuch der Griechischen Lehn- und Fremdwörter im Lateinischen.'' Wien: Druck und Verlag von Carl Gerold's Sohn, Buchhändler der Kaiserl. Akademie der Wissenschaften. in turn from
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
ἀνθηρά (), feminine of ἀνθηρός () meaning "flowery",Liddell, H.G. & Scott, R. (1940). ''A Greek-English Lexicon. revised and augmented throughout by Sir Henry Stuart Jones. with the assistance of. Roderick McKenzie.'' Oxford: Clarendon Press. from ἄνθος () meaning "flower" * Androecium (: androecia) derives from
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
ἀνήρ () meaning "man", and οἶκος () meaning "house" or "chamber/room".


Variation in morphology

Depending on the species of plant, some or all of the stamens in a flower may be attached to the petals or to the floral axis. They also may be free-standing or fused to one another in many different ways, including fusion of some but not all stamens. The filaments may be fused and the anthers free, or the filaments free and the anthers fused. Rather than there being two locules, one locule of a stamen may fail to develop, or alternatively the two locules may merge late in development to give a single locule. Extreme cases of stamen fusion occur in some species of ''
Cyclanthera ''Cyclanthera'' is a genus of flowering plants in the family Cucurbitaceae. The name comes from the fact that some species show extreme cases of stamen The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of ...
'' in the family
Cucurbitaceae The Cucurbitaceae (), also called cucurbits or the gourd family, are a plant family (biology), family consisting of about 965 species in 101 genera.
and in section ''Cyclanthera'' of genus '' Phyllanthus'' (family
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 Euphorbia, the type genus of the family. Most spurges, such as ''Euphorbia paralias'', ar ...
) where the stamens form a ring around the gynoecium, with a single locule. Plants having a single stamen are referred to as "monandrous."


Pollen production

A typical anther contains four microsporangia. The ''microsporangia'' form sacs or pockets (''locules'') in the anther (anther sacs or pollen sacs). The two separate locules on each side of an anther may fuse into a single locule. Each microsporangium is lined with a nutritive tissue layer called the ''tapetum'' and initially contains diploid pollen mother cells. These undergo meiosis to form
haploid Ploidy () is the number of complete sets of chromosomes in a cell (biology), cell, and hence the number of possible alleles for Autosome, autosomal and Pseudoautosomal region, pseudoautosomal genes. Here ''sets of chromosomes'' refers to the num ...
spores. The spores may remain attached to each other in a tetrad or separate after meiosis. Each microspore then divides mitotically to form an immature microgametophyte called a pollen grain. The pollen is eventually released when the anther forms openings ( dehisces). These may consist of longitudinal slits, pores, as in the heath family (
Ericaceae The Ericaceae () are a Family (biology), family of flowering plants, commonly known as the heath or heather family, found most commonly in acidic and infertile growing conditions. The family is large, with about 4,250 known species spread acros ...
), or by valves, as in the barberry family ( Berberidaceae). In some plants, notably members of the
Orchidaceae Orchids are plants that belong to the family (biology), family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant. Orchids are cosmopolitan distribution, cosmopolitan plants that ...
and Asclepiadoideae families, the pollen remains in masses called pollinia, which are adapted to attach to particular pollinating agents such as birds or insects. More commonly, mature pollen grains separate and are dispensed by wind or water, pollinating insects, birds or other pollination vectors. Pollen of
angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed within a fruit ...
s must be transported to the stigma, the receptive surface of the
carpel Gynoecium (; ; : gynoecia) 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 ...
, of a compatible flower, for successful
pollination Pollination is the transfer of pollen from an anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds. Pollinating agents can be animals such as insects, for example bees, beetles or bu ...
to occur. After arriving, the pollen grain (an immature microgametophyte) typically completes its development. It may grow a pollen tube and undergo mitosis to produce two sperm nuclei.


Sexual reproduction in plants

In the typical flower (that is, in the majority of flowering plant species) each flower has both
carpel Gynoecium (; ; : gynoecia) 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 ...
s and stamens. In some species, however, the flowers are unisexual with only carpels or stamens. (
monoecious Monoecy (; adj. monoecious ) is a sexual system in seed plants where separate male and female cones or flowers are present on the same plant. It is a monomorphic sexual system comparable with gynomonoecy, andromonoecy and trimonoecy, and contras ...
= both types of flowers found on the same plant; dioecious = the two types of flower found only on different plants). A flower with only stamens is called androecious. A flower with only carpels is called gynoecious. A pistil consists of one or more carpels. A flower with functional stamens but no functional pistil is called a staminate flower, or (inaccurately) a male flower. A flower with a functional pistil but no functional stamens is called a pistillate flower, or (inaccurately) a female flower.Encyclopædia Britannica.com
/ref> An abortive or rudimentary stamen is called a staminodium or staminode, such as in '' Scrophularia nodosa''. The carpels and stamens of
orchid 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. Orchids are cosmopolitan plants that are found in almost every habitat on Eart ...
s are fused into a column. The top part of the column is formed by the anther, which is covered by an anther cap.


Terminology

;Stamen Stamens can also be adnate (fused or joined from more than one whorl): * epipetalous: adnate to the corolla * epiphyllous: adnate to undifferentiated tepals (as in many Liliaceae) They can have different lengths from each other: * didymous: two equal pairs * didynamous: occurring in two pairs, a long pair and a shorter pair * tetradynamous: occurring as a set of six stamens with four long and two shorter ones or respective to the rest of the flower (
perianth The perianth (perigonium, perigon or perigone in monocots) is the non-reproductive part of the flower. It is a structure that forms an envelope surrounding the sexual organs, consisting of the calyx (sepals) and the corolla (petals) or tepal ...
): * exserted: extending beyond the corolla * included: not extending beyond the corolla They may be arranged in one of two different patterns: * spiral; or * whorled: one or more discrete whorls (series) They may be arranged, with respect to the petals: * diplostemonous: in two whorls, the outer alternating with the petals, while the inner is opposite the petals. * haplostemenous: having a single series of stamens, equal in number to the proper number of petals and alternating with them * obdiplostemonous: in two whorls, with twice the number of stamens as petals, the outer opposite the petals, inner opposite the sepals, e.g. Simaroubaceae (''see
diagram A diagram is a symbolic Depiction, representation of information using Visualization (graphics), visualization techniques. Diagrams have been used since prehistoric times on Cave painting, walls of caves, but became more prevalent during the Age o ...
'') ;Connective Where the connective is very small, or imperceptible, the anther lobes are close together, and the connective is referred to as discrete, e.g. ''
Euphorbia ''Euphorbia'' is a large and diverse genus of flowering plants, commonly called spurge, in the family (biology), family Euphorbiaceae. Euphorbias range from tiny annual plants to large and long-lived trees, with perhaps the tallest being ''Eu ...
'' pp., '' Adhatoda zeylanica''. Where the connective separates the anther lobes, it is called divaricate, e.g. ''
Tilia ''Tilia'' is a genus of about 30 species of trees or bushes, native throughout most of the temperateness, temperate Northern Hemisphere. The tree is known as linden for the European species, and basswood for North American species. In Great Bri ...
'', '' Justicia gendarussa''. The connective may also be a long and stalk-like, crosswise on the filament, this is a distractile connective, e.g. ''
Salvia ''Salvia'' () is the largest genus of plants in the sage family Lamiaceae, with just under 1,000 species of shrubs, Herbaceous plant, herbaceous Perennial plant, perennials, and Annual plant, annuals. Within the Lamiaceae, ''Salvia'' is part o ...
''. The connective may also bear appendages, and is called appendiculate, e.g. '' Nerium odorum'' and some other species of
Apocynaceae Apocynaceae (, from '' Apocynum'', Greek for "dog-away") is a family of flowering plants that includes trees, shrubs, herbs, stem succulents, and vines, commonly known as the dogbane family, because some taxa were used as dog poison. Notable mem ...
. In ''Nerium'', the appendages are united as a staminal corona. ;Filament A column formed from the fusion of multiple filaments is known as an androphore. Stamens can be connate (fused or joined in the same whorl) as follows: * extrorse: anther dehiscence directed away from the centre of the flower. Cf. introrse, directed inwards, and latrorse towards the side. * monadelphous: fused into a single, compound structure * declinate: curving downwards, then up at the tip (also – declinate-descending) * diadelphous: joined partially into two androecial structures * pentadelphous: joined partially into five androecial structures * synandrous: only the anthers are connate (such as in the
Asteraceae Asteraceae () is a large family (biology), family of flowering plants that consists of over 32,000 known species in over 1,900 genera within the Order (biology), order Asterales. The number of species in Asteraceae is rivaled only by the Orchi ...
). The fused stamens are referred to as a synandrium. ;Anther Anther shapes are variously described by terms such as linear, rounded, sagittate, sinuous, or reniform. The anther can be attached to the filament's connective in two ways: * basifixed: attached at its base to the filament ** pseudobasifixed: a somewhat misnomer configuration where connective tissue extends in a tube around the filament tip * dorsifixed: attached at its center to the filament, usually versatile (able to move)


Gallery


References


Bibliography

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External links

{{Authority control Plant anatomy Plant morphology Plant sexuality Plant reproductive system Pollination