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Brassicaceae () or (the older but equally valid) Cruciferae () is a medium-sized and economically important family of commonly known as the mustards, the crucifers, or the cabbage family. Most are , while some are . The are simple (although are sometimes deeply incised), lack , and appear alternately on stems or in rosettes. The are terminal and lack . The flowers have four free , four free alternating , two shorter free and four longer free stamens. The has seeds in rows, divided by a thin wall (or septum).

The family contains 372 and 4,060 accepted . The largest genera are (440 species), (261 species), (234 species), (233 species), and (207 species). , it was divided into two subfamilies, and .

The family contains the cruciferous vegetables, including species such as Brassica oleracea (cultivated as , , , and ), (, , etc.), (, etc.), (common ), Armoracia rusticana (), but also a cut-flower (stock) and the Arabidopsis thaliana (thale cress).

and other butterflies of the family are some of the best-known pests of Brassicaceae species planted as commercial crops. Trichoplusia ni () moth is also becoming increasingly problematic for crucifers due to its resistance to commonly used methods.Turini TA, Daugovish O, Koike ST, Natwick ET, Ploeg A, Dara SK, Fennimore SA, Joseph S, LeStrange M, Smith R, Subbarao KV, Westerdahl BB. Revised continuously. UC IPM Pest Management Guidelines Cole Crops. UC ANR Publication 3442. Oakland, CA. 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 (garlic mustard), an extremely invasive species in the United States, can be toxic to their .


Description
Species belonging to the Brassicaceae are mostly , , or , some are or , and very few . Although generally terrestrial, a few species such as live submerged in fresh water. They may have a or a sometimes woody that may have few or many branches, some have thin or tuberous , or rarely develop . Few species have multi-cellular glands. 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 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 ), and some species lack stems altogether. The leaves do not have , but there may be a pair of glands at base of leaf stalks and flower stalks. The leaf may be seated or have a leafstalk. The leaf blade is usually simple, entire or dissected, rarely trifoliolate 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 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 spp., to 2375 Mbp Bunias orientalis. The number of homologous chromosome sets varies from four (n=4) in some and species, five (n=5) in other Physaria and Stenopetalum species, Arabidopsis thaliana and a species, to seventeen (n=17). About 35% of the species in which chromosomes have been counted have eight sets (n=8). Due to , 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 , , and . Hybridisation between species originating in Africa and California, and subsequent polyploidisation is surmised for species native to Australia and New Zealand.


Inflorescence and flower
of a Brassicaceae ( Erysimum "Bowles' Mauve")]]Flowers may be arranged in , , or , 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 and ) and the ovary positioned above the other floral parts. Each flower has four free or seldom merged , 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 , set alternating with the sepals, although in some species these are rudimentary or absent. They may be differentiated into a blade and a claw 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 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 , four in Cardamine hirsuta, and two in . 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 . The receptacle carries a variable number of , 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 that may either sit directly above the base of the stamens or on a . 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 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 if at least three times longer than wide, or 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 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 are entire or have a notch at the tip. The seed does not contain .


Differences with similar families
Brassicaceae have a bisymmetrical corolla (left is mirrored by right, stem-side by out-side, but each quarter is not symmetrical), a septum dividing the fruit, lack and have simple (although sometimes deeply incised) leaves. The 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 , sometimes an , and a variable number of stamens.


Phytochemistry
Almost all Brassicaceae have C3 carbon fixation. The only exceptions are a few 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.
(2025). 9789400739123, Springer Science & Business Media. .
Brassicaceae contain different cocktails of dozens of . They also contain enzymes called , that convert the glucosinolates into , and , which are toxic to many organisms, and so help guard against herbivory.


Taxonomy
in 1753 regarded the Brassicaceae as a natural group, naming them "Klass" Tetradynamia. Alfred Barton Rendle placed the family in the order Rhoeadales, while and Joseph Dalton Hooker in their system published from 1862 to 1883, assigned it to their cohort (now the class ). Following Bentham and Hooker, John Hutchinson in 1948 and again in 1964 thought the Brassicaceae to stem from near the . In 1994, a group of scientists including Walter Stephen Judd suggested to include the in the Brassicaceae. Early DNA-analysis showed that the Capparaceae—as defined at that moment—were , and it was suggested to assign the genera closest to the Brassicaceae to the . The Cleomaceae and Brassicaceae diverged approximately 41 million years ago. All three families have consistently been placed in one order (variably called or ). The 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 . 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 . Although a substantial effort was made through molecular phylogenetic studies, the relationships within the Brassicaceae have not always been well resolved yet. It has long been clear that the are of the remainder of the family. One analysis from 2014 represented the relation between 39 tribes with the following tree.

As of 2023 the Brassicaceae have been divided into two subfamilies -- the and the (containing only Aethionema) -- the former of which contains five supertribes, , , , , and .


Genera
As of October 2023 Plants of the World Online accepts 346 genera. Brassicaceae Burnett. Plants of the World Online. Retrieved 16 October 2023.


Etymology
The name Brassicaceae comes to international scientific vocabulary from , from , the , + , a standardized for plant family names in modern taxonomy. The genus name comes from the word , referring to and other cruciferous vegetables. The alternative older name, , meaning "cross-bearing", describes the four petals of flowers, which resemble a . 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 , Maritime Southeast Asia and tropical 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 . 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 . A chemical mechanism in the pollen is active in many species to avoid . Two notable exceptions are exclusive in Cardamine chenopodifolia, and wind pollination in Pringlea antiscorbutica. Although it can be cross-pollinated, Alliaria petiolata (garlic mustard) is . Most species reproduce sexually through seed, but Cardamine bulbifera produces 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 , 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 and dolomite rich in . Over a hundred species in the family accumulate , particularly and , which is a record percentage. Several Alyssum species can accumulate nickel up to 0.3% of their dry weight, and may be useful in or even .

Brassicaceae contain as well as inside their cells. When the cell is damaged, the myrosinases the glucosinolates, leading to the synthesis of , which are 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 . 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 ( Plutella xylostella). Like the Pieridae, it is capable of converting isothiocyanates into less problematic . 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 against caterpillars. 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 ( 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 in temperate North America due, in part, to its secretion of allelopathic chemicals. These inhibit the of most competing plants and kill beneficial soil needed by many plants, such as many tree species, to successfully see their seedlings grow to maturity. The 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 is also drastically reduced, particularly in terms of and . Research has found that removing 80% of the garlic mustard plants did not lead to a particularly significant recovery of that . Instead, it required around 100% removal. Given that not one of an estimated 76 that on the plant has been approved for biological control in and the variety of mechanisms the plant has to ensure its dominance without them (e.g. high seed production, self-fertility, , 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. Https://www.dnr.illinois.gov/grants/documents/wpfgrantreports/1998l06w.pdf< /ref> It is estimated that adequate control can be achieved with the introduction of two European , including one that is . The '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 butterflies such as , as they preferentially on it, although it is toxic to their .

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 , , , , , , , , and ( Brassica oleracea), , , , and ( ), ( ), ( ), ( Nasturtium officinale) and ( ) and a few spices like ( Armoracia rusticana), ( Eutrema japonicum), white, brown and black mustard ( , and respectively). Vegetable oil is produced from the seeds of several species such as (rapeseed oil), perhaps providing the largest volume of vegetable oils of any species. ( ) was used in the past to produce a blue textile dye (), but has largely been replaced by the same substance from unrelated tropical species like Indigofera tinctoria.

Pringlea antiscorbutica, commonly known as Kerguelen cabbage, is edible, containing high levels of . 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 , hence the species name's epithet antiscorbutica, which means "against scurvy" in . It was essential to the diets of the whalers on Kerguelen when pork, beef, or was used up.

The Brassicaceae also includes ornamentals, such as species of , , , , , , , , , Lobularia, , , and . Honesty ( ) is cultivated for the decorative value of the translucent remains of the fruits after drying.

(2025). 9780715334232, David & Charles.
It can be a pest species in areas where it is not native.

The small Eurasian weed Arabidopsis thaliana is widely used as in the study of the molecular biology of flowering plants ().

Some species are useful as food plants for , such as certain wild mustard and cress species, such as and Boechera laevigata that are utilized by several North American butterflies.

==Gallery==

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External links
  • BrassiBase, a collection of resources on Brassicaceae biology
  • BrassiToL app, an online Brassicaceae Tree of Life viewer and explorer


Further reading
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