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Brassicaceae () or (the older) Cruciferae () is a medium-sized and economically important
family 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. Idea ...
of
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 commonly known as the mustards, the crucifers, or the cabbage family. Most are
herbaceous plant Herbaceous plants are vascular plants that have no persistent wood, woody stems above ground. This broad category of plants includes many perennial plant, perennials, and nearly all Annual plant, annuals and Biennial plant, biennials. Definition ...
s, while some are shrubs. The leaves are simple (although are sometimes deeply incised), lack
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, and appear alternately on stems or in rosettes. The
inflorescence An inflorescence is a group or cluster of flowers arranged on a Plant stem, stem that is composed of a main branch or a complicated arrangement of branches. Morphology (biology), Morphologically, it is the modified part of the shoot of sperma ...
s are terminal and lack bracts. The flowers have four free
sepal 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 coine ...
s, four free alternating petals, two shorter free stamens and four longer free stamens. The
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 ...
has seeds in rows, divided by a thin wall (or septum). The family contains 372 genera and 4,060 accepted
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
. The largest genera are ''
Draba ''Draba'' is a large genus of flowering plants in the family Brassicaceae, commonly known as whitlow-grasses (though they are not related to the true grasses). Species There are over 400 species: *'' Draba abajoensis'' Windham & Al-Shehbaz *'' ...
'' (440 species), ''
Erysimum ''Erysimum'', or wallflower, is a genus of flowering plants in the cabbage family, Brassicaceae. It includes more than 150 species, both popular garden plants and many wild forms. The genus ''Cheiranthus'' is sometimes included here in whole o ...
'' (261 species), ''
Lepidium ''Lepidium'' is a genus of plants in the mustard/cabbage family, Brassicaceae. The genus is widely distributed in the Americas, Africa, Asia, Europe, and Australia.Cardamine ''Cardamine'' is a large genus of flowering plants in the mustard family, Brassicaceae, known as bittercresses and toothworts. It contains more than 200 species of annuals and perennials. Species in this genus can be found worldwide, except the ...
'' (233 species), and ''
Alyssum ''Alyssum'' is a genus of over a hundred species of flowering plants in the family Brassicaceae, native to Europe, Asia, and northern Africa, with the highest species diversity in the Mediterranean region. The genus comprises annual and perenni ...
'' (207 species). The family contains the
cruciferous vegetables Cruciferous vegetables are vegetables of the family Brassicaceae (also called Cruciferae) with many genera, species, and cultivars being raised for food production such as cauliflower, cabbage, kale, garden cress, bok choy, broccoli, Brusse ...
, including species such as ''
Brassica oleracea ''Brassica oleracea'' is a plant species from family Brassicaceae that includes many common cultivars used as vegetables, such as cabbage, broccoli, cauliflower, kale, Brussels sprouts, collard greens, Savoy cabbage, kohlrabi, and gai lan. ...
'' (cultivated as cabbage, kale, cauliflower,
broccoli Broccoli (''Brassica oleracea'' var. ''italica'') is an edible green plant in the cabbage family (family Brassicaceae, genus ''Brassica'') whose large flowering head, stalk and small associated leaves are eaten as a vegetable. Broccoli is cla ...
and
collards Collard is a group of certain loose-leafed cultivars of '' Brassica oleracea'', the same species as many common vegetables including cabbage ( Capitata group) and broccoli ( Italica group). Collard is a member of the Viridis group of ''Brassic ...
), ''
Brassica rapa ''Brassica rapa'' is a plant species growing in various widely cultivated forms including the turnip (a root vegetable); napa cabbage, bomdong, bok choy, and rapini. ''Brassica rapa'' subsp. ''oleifera'' is an oilseed which has many common n ...
'' (
turnip The turnip or white turnip ('' Brassica rapa'' subsp. ''rapa'') is a root vegetable commonly grown in temperate climates worldwide for its white, fleshy taproot. The word ''turnip'' is a compound of ''turn'' as in turned/rounded on a lathe and ...
,
Chinese cabbage Chinese cabbage (''Brassica rapa'', subspecies ''pekinensis'' and ''chinensis'') can refer to two cultivar groups of leaf vegetables often used in Chinese cuisine: the Pekinensis Group (napa cabbage) and the Chinensis Group (bok choy). These ...
, etc.), '' Brassica napus'' (
rapeseed Rapeseed (''Brassica napus ''subsp.'' napus''), also known as rape, or oilseed rape, is a bright-yellow flowering member of the family Brassicaceae (mustard or cabbage family), cultivated mainly for its oil-rich seed, which naturally contains a ...
, etc.), ''
Raphanus sativus The radish (''Raphanus raphanistrum'' subsp. ''sativus'') is an edible root vegetable of the family Brassicaceae that was domesticated in Asia prior to Roman times. Radishes are grown and consumed throughout the world, being mostly eaten raw ...
'' (common
radish The radish (''Raphanus raphanistrum'' subsp. ''sativus'') is an edible root vegetable of the family Brassicaceae that was domesticated in Asia prior to Roman times. Radishes are grown and consumed throughout the world, being mostly eaten raw ...
), ''
Armoracia rusticana Horseradish (''Armoracia rusticana'', syn. ''Cochlearia armoracia'') is a perennial plant of the family Brassicaceae (which also includes mustard, wasabi, broccoli, cabbage, and radish). It is a root vegetable, cultivated and used worldwide a ...
'' (
horseradish Horseradish (''Armoracia rusticana'', syn. ''Cochlearia armoracia'') is a perennial plant of the family Brassicaceae (which also includes mustard, wasabi, broccoli, cabbage, and radish). It is a root vegetable, cultivated and used worldwi ...
), but also a cut-flower ''
Matthiola ''Matthiola'' () is a genus of flowering plant in the mustard family Brassicaceae. It is named after Italian naturalist Pietro Andrea Mattioli (1501–1577). The genus contains about 50 species of annual, biennial and perennial herbaceous plants ...
'' (stock) and the model organism '' Arabidopsis thaliana'' (thale cress). ''
Pieris rapae ''Pieris rapae'' is a small- to medium-sized butterfly species of the whites-and-yellows family Pieridae. It is known in Europe as the small white, in North America as the cabbage white or cabbage butterfly, on several continents as the small c ...
'' and other butterflies of the family
Pieridae The Pieridae are a large family of butterflies with about 76 genera containing about 1,100 species, mostly from tropical Africa and tropical Asia with some varieties in the more northern regions of North America and Eurasia.DeVries P. J. in Le ...
are some of the best-known pests of Brassicaceae species planted as commercial crops. ''Trichoplusia ni'' (
cabbage looper The cabbage looper (''Trichoplusia ni'') is a medium-sized moth in the family Noctuidae, a family commonly referred to as owlet moths. Its common name comes from its preferred host plants and distinctive crawling behavior. Cruciferous vegetable ...
) moth is also becoming increasingly problematic for crucifers due to its resistance to commonly used
pest control Pest control is the regulation or management of a species defined as a pest; any animal, plant or fungus that impacts adversely on human activities or environment. The human response depends on the importance of the damage done and will range ...
methods. Some rarer ''Pieris'' butterflies, such as '' P. virginiensis'', depend upon native mustards for their survival in their native habitats. Some non-native mustards such as ''
Alliaria petiolata ''Alliaria petiolata'', or garlic mustard, is a biennial flowering plant in the mustard family (Brassicaceae). It is native to Europe, western and central Asia, north-western Africa, Morocco, Iberia and the British Isles, north to northern S ...
'' (garlic mustard), an extremely invasive species in the United States, can be toxic to their
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
e.


Description

Species belonging to the Brassicaceae are mostly
annual Annual may refer to: *Annual publication, periodical publications appearing regularly once per year ** Yearbook ** Literary annual *Annual plant *Annual report *Annual giving *Annual, Morocco, a settlement in northeastern Morocco *Annuals (band), ...
,
biennial Biennial means (an event) lasting for two years or occurring every two years. The related term biennium is used in reference to a period of two years. In particular, it can refer to: * Biennial plant, a plant which blooms in its second year and th ...
, or
perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wid ...
herbaceous plant Herbaceous plants are vascular plants that have no persistent wood, woody stems above ground. This broad category of plants includes many perennial plant, perennials, and nearly all Annual plant, annuals and Biennial plant, biennials. Definition ...
s, some are dwarf shrubs or shrubs, and very few
vine A vine (Latin ''vīnea'' "grapevine", "vineyard", from ''vīnum'' "wine") is any plant with a growth habit of trailing or scandent (that is, climbing) stems, lianas or runners. The word ''vine'' can also refer to such stems or runners themsel ...
s. Although generally terrestrial, a few species such as water awlwort live submerged in fresh water. They may have a
taproot A taproot is a large, central, and dominant root from which other roots sprout laterally. Typically a taproot is somewhat straight and very thick, is tapering in shape, and grows directly downward. In some plants, such as the carrot, the taproo ...
or a sometimes woody caudex that may have few or many branches, some have thin or tuberous rhizomes, or rarely develop runners. Few species have multi-cellular glands.
Hairs Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and f ...
consist of one cell and occur in many forms: from simple to forked, star-, tree- or T-shaped, rarely taking the form of a shield or scale. They are never topped by a gland. The stems may be upright, rise up towards the tip, or lie flat, are mostly herbaceous but sometimes woody. Stems carry leaves or the stems may be leafless (in ''
Caulanthus ''Caulanthus'' is a genus of plants in the family Brassicaceae. Plants of this genus may be known as jewelflowers. They are also often referred to as wild cabbage, although this common name usually refers to wild variants of ''Brassica oleraceae' ...
''), and some species lack stems altogether. The leaves do not have
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, but there may be a pair of glands at base of leafstalks and flowerstalks. The leaf may be seated or have a leafstalk. The
leaf blade 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 ...
is usually simple, entire or
dissected Dissection (from Latin ' "to cut to pieces"; also called anatomization) is the dismembering of the body of a deceased animal or plant to study its anatomical structure. Autopsy is used in pathology and forensic medicine to determine the cause ...
, rarely
trifoliolate The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular o ...
or pinnately compound. A leaf rosette at the base may be present or absent. The leaves along the stem are almost always alternately arranged, rarely apparently opposite. The stomata are of the anisocytic type. The
genome size Genome size is the total amount of DNA contained within one copy of a single complete genome. It is typically measured in terms of mass in picograms (trillionths (10−12) of a gram, abbreviated pg) or less frequently in daltons, or as the total ...
of Brassicaceae compared to that of other Angiosperm families is very small to small (less than 3.425 million base pairs per cell), varying from 150 Mbp in '' Arabidopsis thaliana'' and '' Sphaerocardamum'' spp., to 2375 Mbp '' Bunias orientalis''. The number of homologous chromosome sets varies from four (n=4) in some ''
Physaria ''Physaria'' is a genus of flowering plants in the family Brassicaceae. Many species are known generally as twinpods, bladderpods, or lesquerella. They are native to the Americas, with many species endemic to western North America. They are dense ...
'' and '' Stenopetalum'' species, five (n=5) in other ''Physaria'' and ''Stenopetalum'' species, ''Arabidopsis thaliana'' and a '' Mathiola'' species, to seventeen (n=17). About 35% of the species in which chromosomes have been counted have eight sets (n=8). Due to
polyploidy Polyploidy is a condition in which the biological cell, cells of an organism have more than one pair of (Homologous chromosome, homologous) chromosomes. Most species whose cells have Cell nucleus, nuclei (eukaryotes) are diploid, meaning they ha ...
, some species may have up to 256 individual chromosomes, with some very high counts in the North American species of ''Cardamine'', such as '' C. diphylla''. Hybridisation is not unusual in Brassicaceae, especially in ''
Arabis ''Arabis'' ,''Sunset Western Garden Book,'' 1995:606–607 or rockcress, is a genus of flowering plants, within the family Brassicaceae. Description The species are herbaceous, annual or perennial plants, growing to 10–80 cm tall, usua ...
'', ''
Rorippa ''Rorippa'' is a genus of flowering plants in the family Brassicaceae, native to Europe through central Asia, Africa, and North America. ''Rorippa'' species are annual to perennial herbs, usually with yellow flowers and a peppery flavour. They a ...
'', ''
Cardamine ''Cardamine'' is a large genus of flowering plants in the mustard family, Brassicaceae, known as bittercresses and toothworts. It contains more than 200 species of annuals and perennials. Species in this genus can be found worldwide, except the ...
'' and ''
Boechera ''Boechera'' (rockcress) is a genus of the family Brassicaceae. It was named after the Danish botanist Tyge W. Böcher (1909–1983), who was known for his research in alpine plants, including the mustards ''Draba'' and ''Boechera holboellii''. ...
''. Hybridisation between species originating in Africa and California, and subsequent polyploidisation is surmised for ''
Lepidium ''Lepidium'' is a genus of plants in the mustard/cabbage family, Brassicaceae. The genus is widely distributed in the Americas, Africa, Asia, Europe, and Australia.racemes, panicles, or
corymb Corymb is a botanical term for an inflorescence with the flowers growing in such a fashion that the outermost are borne on longer pedicels than the inner, bringing all flowers up to a common level. A corymb has a flattish top with a superficial ...
s, with pedicels sometimes in the axil of a bract, and few species have flowers that sit individually on flower stems that spring from the axils of rosette leaves. The orientation of the pedicels when fruits are ripe varies dependent on the species. The flowers are bisexual, star symmetrical (zygomorphic in ''
Iberis ''Iberis'' , commonly called candytuft, is a genus of flowering plants belonging to the family Brassicaceae. It comprises annuals, evergreen perennials and subshrubs native to the Old World. The name "candytuft" is not related to candy, but der ...
'' and '' Teesdalia'') and the ovary positioned above the other floral parts. Each flower has four free or seldom merged
sepal 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 coine ...
s, the lateral two sometimes with a shallow spur, which are mostly shed after flowering, rarely persistent, may be reflexed, spreading, ascending, or erect, together forming a tube-, bell- or urn-shaped calyx. Each flower has four petals, set alternating with the sepals, although in some species these are rudimentary or absent. They may be differentiated into a
blade A blade is the portion of a tool, weapon, or machine with an edge that is designed to puncture, chop, slice or scrape surfaces or materials. Blades are typically made from materials that are harder than those they are to be used on. Histor ...
and a
claw A claw is a curved, pointed appendage found at the end of a toe or finger in most amniotes (mammals, reptiles, birds). Some invertebrates such as beetles and spiders have somewhat similar fine, hooked structures at the end of the leg or tarsus ...
or not, and consistently lack basal appendages. The blade is entire or has an indent at the tip, and may sometimes be much smaller than the claws. The mostly six stamens are set in two whorls: usually the two lateral, outer ones are shorter than the four inner stamens, but very rarely the stamens can all have the same length, and very rarely species have different numbers of stamens such as sixteen to twenty four in '' Megacarpaea'', four in '' Cardamine hirsuta'', and two in '' Coronopus''. The filaments are slender and not fused, while the anthers consist of two pollen producing cavities, and open with longitudinal slits. The pollen grains are tricolpate. The receptacle carries a variable number of nectaries, but these are always present opposite the base of the lateral stamens.


Ovary, fruit and seed

There is one superior pistil that consists of two
carpel 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' ...
s that may either sit directly above the base of the stamens or on a stalk. It initially consists of only one cavity but during its further development a thin wall grows that divides the cavity, both placentas and separates the two valves (a so-called false septum). Rarely, there is only one cavity without a septum. The 2–600
ovule In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the '' integument'', forming its outer layer, the ''nucellus'' (or remnant of the megasporangium), and the ...
s are usually along the side margin of the carpels, or rarely at the top. Fruits are capsules that open with two valves, usually towards the top. These are called
silique A silique or siliqua (plural ''siliques'' or ''siliquae'') is a type of fruit (seed capsule) having two fused carpels with the length being more than three times the width. When the length is less than three times the width of the dried fruit i ...
if at least three times longer than wide, or
silicle A silique or siliqua (plural ''siliques'' or ''siliquae'') is a type of fruit (seed capsule) having two fused carpels with the length being more than three times the width. When the length is less than three times the width of the dried fruit i ...
if the length is less than three times the width. The fruit is very variable in its other traits. There may be one persistent
style Style is a manner of doing or presenting things and may refer to: * Architectural style, the features that make a building or structure historically identifiable * Design, the process of creating something * Fashion, a prevailing mode of clothing ...
that connects the ovary to the globular or conical stigma, which is undivided or has two spreading or connivent lobes. The variously shaped seeds are usually yellow or brown in color, and arranged in one or two rows in each cavity. The seed leaves are entire or have a notch at the tip. The seed does not contain endosperm.


Differences with similar families

Brassicaceae have a bisymmetical corolla (left is mirrored by right, stem-side by out-side, but each quarter is not symmetrical), a septum dividing the fruit, lack
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 and have simple (although sometimes deeply incised) leaves. The sister family
Cleomaceae The Cleomaceae are a small family of flowering plants in the order Brassicales, comprising about 300 species in 10 genera, or about 150 species in 17 genera.Cleomaceae, Zhang Mingli (张明理)1; Gordon C. Tucker2, Harvard.edu/ref> These genera ...
has bilateral symmetrical corollas (left is mirrored by right, but stem-side is different from out-side), stipules and mostly palmately divided leaves, and mostly no septum. Capparaceae generally have a
gynophore A gynophore is the stalk of certain flowers which supports the gynoecium (the ovule-producing part of a flower), elevating it above the branching points of other floral parts. Plant genera that have flowers with gynophores include '' Telopea'', '' ...
, sometimes an androgynophore, and a variable number of stamens.


Phytochemistry

Almost all Brassicaceae have
C3 carbon fixation carbon fixation is the most common of three metabolic pathways for carbon fixation in photosynthesis, along with C4 carbon fixation, and Crassulacean acid metabolism, CAM. This process converts carbon dioxide and ribulose bisphosphate (RuBP, a ...
. The only exceptions are a few '' Moricandia'' species, which have a hybrid system between C3 and C4 carbon fixation, C4 fixation being more efficient in drought, high temperature and low nitrate availability. Brassicaceae contain different cocktails of dozens of glucosinolates. They also contain enzymes called
myrosinase Myrosinase (, ''thioglucoside glucohydrolase'', ''sinigrinase'', and ''sinigrase'') is a family of enzymes involved in plant defense against herbivores, specifically the mustard oil bomb. The three-dimensional structure has been elucidated and is ...
s, that convert the glucosinolates into
isothiocyanate In organic chemistry, isothiocyanate is the functional group , formed by substituting the oxygen in the isocyanate group with a sulfur. Many natural isothiocyanates from plants are produced by enzymatic conversion of metabolites called glucosi ...
s,
thiocyanate Thiocyanate (also known as rhodanide) is the anion . It is the conjugate base of thiocyanic acid. Common derivatives include the colourless salts potassium thiocyanate and sodium thiocyanate. Mercury(II) thiocyanate was formerly used in pyr ...
s and nitriles, which are toxic to many organisms, and so help guard against herbivory.


Taxonomy

Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his Nobility#Ennoblement, ennoblement in 1761 as Carl von Linné#Blunt, Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalise ...
in 1753 regarded the Brassicaceae as a natural group, naming them "Klass" Tetradynamia. Alfred Barton Rendle placed the family in the order Rhoedales, while
George Bentham George Bentham (22 September 1800 – 10 September 1884) was an English botanist, described by the weed botanist Duane Isely as "the premier systematic botanist of the nineteenth century". Born into a distinguished family, he initially studie ...
and
Joseph Dalton Hooker Sir Joseph Dalton Hooker (30 June 1817 – 10 December 1911) was a British botanist and explorer in the 19th century. He was a founder of geographical botany and Charles Darwin's closest friend. For twenty years he served as director of ...
in their system published from 1862–1883, assigned it to their cohort Parietales (now the class Violales). Following Bentham and Hooker, John Hutchinson (botanist), John Hutchinson in 1948 and again in 1964 thought the Brassicaceae to stem from near the Papaveraceae. In 1994, a group of scientists including Walter Stephen Judd suggested to include the Capparaceae in the Brassicaceae. Early DNA-analysis showed that the Capparaceae—as defined at that moment—were paraphyletic, and it was suggested to assign the genera closest to the Brassicaceae to the
Cleomaceae The Cleomaceae are a small family of flowering plants in the order Brassicales, comprising about 300 species in 10 genera, or about 150 species in 17 genera.Cleomaceae, Zhang Mingli (张明理)1; Gordon C. Tucker2, Harvard.edu/ref> These genera ...
. The Cleomaceae and Brassicaceae diverged approximately 41 million years ago. All three families have consistently been placed in one order (variably called Capparales or Brassicales). The APG II system merged Cleomaceae and Brassicaceae. Other classifications have continued to recognize the Capparaceae, but with a more restricted circumscription, either including ''Cleome'' and its relatives in the Brassicaceae or recognizing them in the segregate family
Cleomaceae The Cleomaceae are a small family of flowering plants in the order Brassicales, comprising about 300 species in 10 genera, or about 150 species in 17 genera.Cleomaceae, Zhang Mingli (张明理)1; Gordon C. Tucker2, Harvard.edu/ref> These genera ...
. The APG III system has recently adopted this last solution, but this may change as a consensus arises on this point. Current insights in the relationships of the Brassicaceae, based on a 2012 DNA-analysis, are summarized in the following tree.


Relationships within the family

Early classifications depended on morphological comparison only, but because of extensive convergent evolution, these do not provide a reliable phylogeny. Although a substantial effort was made through Molecular phylogenetics, molecular phylogenetic studies, the relationships within the Brassicaceae have not always been well resolved yet. It has long been clear that the ''Aethionema'' are Sister group, sister of the remainder of the family. One analysis from 2014 represented the relation between 39 tribes with the following tree.


Genera

Version 1 of The Plant List lists List of Brassicaceae genera, 349 genera.


Etymology

The name ''Brassicaceae'' comes to international scientific vocabulary from New Latin, from ''Brassica'', the type genus, + ''wikt:-aceae#Suffix, -aceae'', a standardized suffix for plant family names in modern taxonomy. The genus name comes from the Classical Latin word ''wikt:brassica#Noun 2, brassica'', referring to cabbage and other
cruciferous vegetables Cruciferous vegetables are vegetables of the family Brassicaceae (also called Cruciferae) with many genera, species, and cultivars being raised for food production such as cauliflower, cabbage, kale, garden cress, bok choy, broccoli, Brusse ...
. The alternative older name, wikt:Cruciferae, Cruciferae, meaning "cross-bearing", describes the four petals of Mustard plant, mustard flowers, which resemble a cross. Cruciferae is one of eight plant family names, not derived from a genus name and without the suffix ''-aceae'' that are authorized alternative names.


Distribution

Brassicaceae can be found almost on the entire land surface of the planet, but the family is absent from Antarctica, and also absent from some areas in the tropics i.e. northeastern Brazil, the Congo basin, Maritime Southeast Asia and tropical Australasian realm, Australasia. The area of origin of the family is possibly the Irano-Turanian Region, where approximately 900 species occur in 150 different genera. About 530 of those 900 species are endemics. Next in abundance comes the Mediterranean Region, with around 630 species (290 of which are endemic) in 113 genera. The family is less prominent in the Saharo-Arabian Region—65 genera, 180 species of which 62 are endemic—and North America (comprising the North American Atlantic Region and the Rocky Mountain Floristic Region)—99 genera, 780 species of which 600 are endemic. South-America has 40 genera containing 340 native species, Southern Africa 15 genera with over 100 species, and Australia and New-Zealand have 19 genera with 114 species between them.


Ecology

Brassicaceae are almost exclusively Entomophily, pollinated by insects. A chemical mechanism in the pollen is active in many species to avoid self-pollination, selfing. Two notable exceptions are exclusive Cleistogamy, self-pollination in closed flowers in ''Cardamine chenopodifolia'', and wind pollination in ''Pringlea antiscorbutica''. Although it can be cross-pollinated, ''
Alliaria petiolata ''Alliaria petiolata'', or garlic mustard, is a biennial flowering plant in the mustard family (Brassicaceae). It is native to Europe, western and central Asia, north-western Africa, Morocco, Iberia and the British Isles, north to northern S ...
'' (garlic mustard) is self-fertile. Most species reproduce sexually through seed, but ''Cardamine bulbifera'' produces Gemma (botany), gemmae and in others, such as ''Cardamine pentaphyllos'', the coral-like roots easily break into segments, that will grow into separate plants. In some species, such as in the genus ''
Cardamine ''Cardamine'' is a large genus of flowering plants in the mustard family, Brassicaceae, known as bittercresses and toothworts. It contains more than 200 species of annuals and perennials. Species in this genus can be found worldwide, except the ...
'', seed pods open with force and so catapult the seeds quite far. Many of these have sticky seed coats, assisting long distance dispersal by animals, and this may also explain several intercontinental dispersal events in the genus, and its near global distribution. Brassicaceae are common on serpentine soil, serpentine and Dolomite (mineral), dolomite rich in magnesium. Over a hundred species in the family accumulate heavy metals, particularly zinc and nickel, which is a record percentage. Several ''Alyssum'' species can accumulate nickel up to 0.3% of their dry weight, and may be useful in soil remediation or even bio-mining. Brassicaceae contain glucosinolates as well as
myrosinase Myrosinase (, ''thioglucoside glucohydrolase'', ''sinigrinase'', and ''sinigrase'') is a family of enzymes involved in plant defense against herbivores, specifically the mustard oil bomb. The three-dimensional structure has been elucidated and is ...
s inside their cells. When the cell is damaged, the myrosinases Hydrolysis, hydrolise the glucosinolates, leading to the synthesis of
isothiocyanate In organic chemistry, isothiocyanate is the functional group , formed by substituting the oxygen in the isocyanate group with a sulfur. Many natural isothiocyanates from plants are produced by enzymatic conversion of metabolites called glucosi ...
s, which are Plant defense against herbivory, compounds toxic to most animals, fungi and bacteria. Some insect herbivores have developed counter adaptations such as rapid absorption of the glucosinates, quick alternative breakdown into non-toxic compounds and avoiding cell damage. In the whites family (Pieridae), one counter mechanism involves glucosinolate sulphatase, which changes the glucosinolate, so that it cannot be converted to isothiocyanate. A second is that the glucosinates are quickly broken down, forming nitriles. Differences between the mixtures of glucosinolates between species and even within species is large, and individual plants may produce in excess of fifty individual substances. The energy penalty for synthesising all these glucosinolates may be as high as 15% of the total needed to produce a leaf. ''Barbarea vulgaris'' (bittercress) also produces triterpenoid saponins. These adaptations and counter adaptations probably have led to extensive diversification in both the Brassicaceae and one of its major pests, the butterfly family
Pieridae The Pieridae are a large family of butterflies with about 76 genera containing about 1,100 species, mostly from tropical Africa and tropical Asia with some varieties in the more northern regions of North America and Eurasia.DeVries P. J. in Le ...
. A particular cocktail of volatile glucosinates triggers egg-laying in many species. Thus a particular crop can sometimes be protected by planting bittercress as a deadly bait, for the saponins kill the caterpillars, but the butterfly is still lured by the bittercress to lay its egg on the leaves. A moth that feeds on a range of Brassicaceae is the diamondback moth (''Plutella xylostella''). Like the Pieridae, it is capable of converting isothiocyanates into less problematic nitriles. Managing this pest in crops became more complicated after resistance developed against a toxin produced by ''Bacillus thuringiensis'', which is used as a wide spectrum biological plant protection against caterpillars. Parasitoid wasps that feed on such insect herbivores are attracted to the chemical compounds released by the plants, and thus are able to locate their prey. The cabbage aphid (''Brevicoryne brassicae'') stores glucosinolates and synthesises its own myrosinases, which may deter its potential predators. Since its introduction in the 19th century, ''Alliaria petiolata'' has been shown to be extremely successful as an invasive species in temperate North America due, in part, to its secretion of allelopathic chemicals. These inhibit the germination of most competing plants and kill beneficial soil fungi needed by many plants, such as many tree species, to successfully see their seedlings grow to maturity. The monoculture formation of an herb layer carpet by this plant has been shown to dramatically alter forests, making them wetter, having fewer and fewer trees, and having more vines such as poison ivy (''Toxicodendron radicans''). The overall herb layer biodiversity is also drastically reduced, particularly in terms of sedges and forbs. Research has found that removing 80% of the garlic mustard infestation plants did not lead to a particularly significant recovery of that Biodiversity, diversity. Instead, it required around 100% removal. Given that not one of an estimated 76
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
that prey on the plant has been approved for biological control in North America and the variety of mechanisms the plant has to ensure its dominance without them (e.g. high seed production, self-fertility, allelopathy, spring growth that occurs before nearly all native plants, roots that break easily when pulling attempts are made, a complete lack of palatability for herbivores at all life stages, etc.) it is unlikely that such a high level of control can be established and maintained on the whole.Becker, R., Gerber E., Hinz H., Katovich E., Panke B., Reardon R., Renz R., Van Riper L., 2013. Biology and Biological Control of Garlic Mustard. The Forest Technology Enterprise Team. https://www.fs.fed.us/foresthealth/technology/pdfs/FS_garlicmustard.pdfDriesche, F.V.; Blossey, B.; Hoodle, M.; Lyon, S.; Reardon, R., 2010. Biological Control of Invasive Plants in the Eastern United States. USDA Forest Service. Forest Health Technology Enterprise Team. http://wiki.bugwood.org/Archive:BCIPEUS It is estimated that adequate control can be achieved with the introduction of two European weevils, including one that is monophagous. The USDA's TAG group has blocked these introductions since 2004. In addition to being invasive, garlic mustard also is a threat to native North American ''Pieris (butterfly), Pieris'' butterflies such as ''Pieris oleracea, P. oleracea'', as they preferentially oviposit on it, although it is toxic to their
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
e. Invasive aggressive mustard species are known for being self-fertile, seeding very heavily with small seeds that have a lengthy lifespan coupled with a very high rate of viability and germination, and for being completely unpalatable to both herbivores and insects in areas to which they are not native. Garlic mustard is toxic to several rarer North American ''Pieris'' species.


Uses

This family includes important agricultural crops, among which many vegetables such as cabbage,
broccoli Broccoli (''Brassica oleracea'' var. ''italica'') is an edible green plant in the cabbage family (family Brassicaceae, genus ''Brassica'') whose large flowering head, stalk and small associated leaves are eaten as a vegetable. Broccoli is cla ...
, cauliflower, kale, Brussels sprouts, collard greens, savoy cabbage, Savoy, kohlrabi, and gai lan (''
Brassica oleracea ''Brassica oleracea'' is a plant species from family Brassicaceae that includes many common cultivars used as vegetables, such as cabbage, broccoli, cauliflower, kale, Brussels sprouts, collard greens, Savoy cabbage, kohlrabi, and gai lan. ...
''),
turnip The turnip or white turnip ('' Brassica rapa'' subsp. ''rapa'') is a root vegetable commonly grown in temperate climates worldwide for its white, fleshy taproot. The word ''turnip'' is a compound of ''turn'' as in turned/rounded on a lathe and ...
, napa cabbage, bomdong, bok choy and rapini (''
Brassica rapa ''Brassica rapa'' is a plant species growing in various widely cultivated forms including the turnip (a root vegetable); napa cabbage, bomdong, bok choy, and rapini. ''Brassica rapa'' subsp. ''oleifera'' is an oilseed which has many common n ...
''), rocket salad, rocket salad/arugula (''Eruca sativa''), garden cress (''Lepidium sativum''), watercress (''Nasturtium officinale'') and
radish The radish (''Raphanus raphanistrum'' subsp. ''sativus'') is an edible root vegetable of the family Brassicaceae that was domesticated in Asia prior to Roman times. Radishes are grown and consumed throughout the world, being mostly eaten raw ...
(''Raphanus'') and a few spices like
horseradish Horseradish (''Armoracia rusticana'', syn. ''Cochlearia armoracia'') is a perennial plant of the family Brassicaceae (which also includes mustard, wasabi, broccoli, cabbage, and radish). It is a root vegetable, cultivated and used worldwi ...
(''
Armoracia rusticana Horseradish (''Armoracia rusticana'', syn. ''Cochlearia armoracia'') is a perennial plant of the family Brassicaceae (which also includes mustard, wasabi, broccoli, cabbage, and radish). It is a root vegetable, cultivated and used worldwide a ...
''), ''Brassica'', wasabi (''Eutrema japonicum''), white, Indian and black mustard (''Sinapis alba'', ''Brassica juncea'' and ''Brassica nigra, B. nigra'' respectively). Vegetable oil is produced from the seeds of several species such as '' Brassica napus'' (rapeseed oil), perhaps providing the largest volume of vegetable oils of any species. Woad (''Isatis tinctoria'') was used in the past to produce a blue textile dye (indigo), but has largely been replaced by the same substance from unrelated tropical species like ''Indigofera tinctoria''. Brassinosteroids are growing in agricultural and gardening importance. ''Pringlea antiscorbutica'', commonly known as Kerguelen cabbage, is edible, containing high levels of potassium. Its leaves contain a vitamin C-rich oil, a fact which, in the days of sailing ships, made it very attractive to sailors suffering from scurvy, hence the species name's epithet ''antiscorbutica'', which means "against scurvy" in Low Latin. It was essential to the diets of the whalers on Kerguelen Islands, Kerguelen when pork, beef, or seal meat was used up. The Brassicaceae also includes ornamentals, such as species of ''Aethionema'', ''
Alyssum ''Alyssum'' is a genus of over a hundred species of flowering plants in the family Brassicaceae, native to Europe, Asia, and northern Africa, with the highest species diversity in the Mediterranean region. The genus comprises annual and perenni ...
'', ''
Arabis ''Arabis'' ,''Sunset Western Garden Book,'' 1995:606–607 or rockcress, is a genus of flowering plants, within the family Brassicaceae. Description The species are herbaceous, annual or perennial plants, growing to 10–80 cm tall, usua ...
'', ''Aubrieta'', ''Aurinia'', ''Cheiranthus'', ''
Erysimum ''Erysimum'', or wallflower, is a genus of flowering plants in the cabbage family, Brassicaceae. It includes more than 150 species, both popular garden plants and many wild forms. The genus ''Cheiranthus'' is sometimes included here in whole o ...
'', ''Hesperis'', ''
Iberis ''Iberis'' , commonly called candytuft, is a genus of flowering plants belonging to the family Brassicaceae. It comprises annuals, evergreen perennials and subshrubs native to the Old World. The name "candytuft" is not related to candy, but der ...
'', ''Lobularia (plant), Lobularia'', ''Lunaria'', ''Malcolmia'', and ''
Matthiola ''Matthiola'' () is a genus of flowering plant in the mustard family Brassicaceae. It is named after Italian naturalist Pietro Andrea Mattioli (1501–1577). The genus contains about 50 species of annual, biennial and perennial herbaceous plants ...
''. Honesty (plant), Honesty (''Lunaria annua'') is cultivated for the decorative value of the translucent remains of the fruits after drying. It can be a pest species in areas where it is not native. The small Eurasian weed '' Arabidopsis thaliana'' is widely used as model organism in the study of the molecular biology of flowering plants (Angiospermae). Some species are useful as food plants for Lepidoptera, such as certain wild mustard and cress species, such as ''Turritis glabra'' and ''Boechera laevigata'' that are utilized by several North American butterflies.


Gallery

File:Brassicaceae_Coast_Sand_Loving_wallflower_erysimum_ammophilum.jpg, Coast/sand-loving wallflower ''Erysimum ammophilum'' File:Brassicaceae_Money_Plant_Honesty_Lunaria_annua.jpg, Honesty ''Lunaria annua'' File:Brassicaceae_Western_wallflower_erysimum_capitatum_var_capitatum.jpg, Western wallflower ''Erysimum capitatum var. capitatum''


References


External links


BrassiBase
a collection of resources on Brassicaceae biology


Further reading

* {{Authority control Brassicaceae, Brassicales families