Brassicaceae
Brassicaceae or 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 leaves are simple, lack stipules, and appear alternately on stems or in rosettes. The inflorescences are terminal and lack bracts. The flowers have four free sepals, four free alternating petals, two shorter free stamens and four longer free stamens. The fruit has seeds in rows, divided by a thin wall., the family was divided into two subfamilies, Brassicoideae and Aethionemoideae, and as of 2025, it contained over 350 accepted genera.
Pieris rapae and other butterflies of the family Pieridae 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 pest control 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, an extremely invasive species in the United States, can be toxic to their larvae.
The family contains the cruciferous vegetables, including species such as Brassica oleracea, Brassica rapa, Brassica napus, Raphanus sativus, Armoracia rusticana, the cut-flower Matthiola and the model organism Arabidopsis thaliana.
Description
Species belonging to the Brassicaceae are mostly annual, biennial, or perennial herbaceous plants, some are dwarf shrubs or shrubs, and very few vines. Although generally terrestrial, a few species such as water awlwort live submerged in fresh water. They may have a taproot 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 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, and some species lack stems altogether. The leaves do not have stipules, 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 genome size of Brassicaceae compared to that of other Angiosperm families is very small to small, varying from 150 Mbp in Arabidopsis thaliana and Sphaerocardamum spp., to 2375 Mbp Bunias orientalis. The number of homologous chromosome sets varies from four in some Physaria and Stenopetalum species, five in other Physaria and Stenopetalum species, Arabidopsis thaliana and a Mathiola species, to seventeen. About 35% of the species in which chromosomes have been counted have eight sets. Due to polyploidy, 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, Rorippa, Cardamine and Boechera. Hybridisation between species originating in Africa and California, and subsequent polyploidisation is surmised for Lepidium species native to Australia and New Zealand.Inflorescence and flower
Flowers may be arranged in racemes, panicles, or corymbs, 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 and the ovary positioned above the other floral parts. Each flower has four free or seldom merged sepals, 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 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 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 carpels 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. Rarely, there is only one cavity without a septum. The 2–600 ovules 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 if at least three times longer than wide, or silicle 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 seed leaves are entire or have a notch at the tip. The seed does not contain endosperm.Differences from similar families
Brassicaceae have a bisymmetrical corolla, a septum dividing the fruit, lack stipules and have simple leaves. The sister family Cleomaceae has bilateral symmetrical corollas, stipules and mostly palmately divided leaves, and mostly no septum. Capparaceae generally have a gynophore, sometimes an androgynophore, and a variable number of stamens.Phytochemistry
Almost all Brassicaceae have C3 carbon fixation. 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 myrosinases, that convert the glucosinolates into isothiocyanates, thiocyanates and nitriles, 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 George Bentham and Joseph Dalton Hooker in their system published from 1862 to 1883, assigned it to their cohort Parietales. Following Bentham and Hooker, 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 and Brassicaceae diverged approximately 41 million years ago. All three families have consistently been placed in one order. 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 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 phylogenetic studies, the relationships within the Brassicaceae have not always been well resolved yet. It has long been clear that the Aethionema are sister 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 Brassicoideae and the Aethionemoideae —the former of which contains five supertribes, Arabodae, Brassicodae, Camelinodae, Heliophilodae, and Hesperodae.