Гүлді өсімдіктердің Brassicaceae тұқымдасы

Family of flowering plants

Brassicaceae (,/b//r//æ//s//ᵻ/'/k//eɪ//s//i://%//i://, // //s//i//%//aI/) or (the older) Cruciferae (/k//r//uː/'/s//ɪ//f//ər//i/) 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 (although are sometimes deeply incised), 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 (or septum). The family contains 372 genera and 4,060 accepted species. The largest genera are Draba (440 species), Erysimum (261 species), Lepidium (234 species), Cardamine (233 species), and Alyssum (207 species). The family contains the cruciferous vegetables, including species such as Brassica oleracea (cultivated as cabbage, kale, cauliflower, broccoli and collards), Brassica rapa (turnip, Chinese cabbage, etc. ), Brassica napus (rapeseed, etc. ), Raphanus sativus (common radish), Armoracia rusticana (horseradish), but also a cut flower Matthiola (stock) and the model organism Arabidopsis thaliana (thale cress). 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 (cabbage looper) 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 (garlic mustard), an extremely invasive species in the United States, can be toxic to their larvae.

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 (in Caulanthus), 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 (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 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, 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.

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 (a so called false septum). 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. 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. 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).

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 October 2023 Plants of the World Online accepts 346 genera.

The name Brassicaceae comes to international scientific vocabulary from Neo Latin, from Brassica, the type genus, + aceae, a standardized suffix for plant family names in modern taxonomy. The genus name comes from the Classical Latin word brassica, referring to cabbage and other cruciferous vegetables. The alternative older name, Cruciferae, meaning "cross bearing", describes the four petals of 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.

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 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. 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 myrosinases inside their cells. When the cell is damaged, the myrosinases hydrolise the glucosinolates, leading to the synthesis of isothiocyanates, 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 Pieridae. 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. 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 butterflies such as P. oleracea, as they preferentially oviposit on it, although it is toxic to their larvae. 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.

This family includes important agricultural crops, among which many vegetables such as cabbage, broccoli, cauliflower, kale, Brussels sprouts, collard greens, Savoy, kohlrabi, and gai lan (Brassica oleracea), turnip, napa cabbage, mizuna, bok choy and rapini (Brassica rapa), rocket salad/arugula (Eruca sativa), garden cress (Lepidium sativum), watercress (Nasturtium officinale) and radish (Raphanus) and a few spices like horseradish (Armoracia rusticana), wasabi (Eutrema japonicum), white, Indian and black mustard (Sinapis alba, Brassica juncea and 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. 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 when pork, beef, or seal meat was used up. The Brassicaceae also includes ornamentals, such as species of Aethionema, Alyssum, Arabis, Aubrieta, Aurinia, Cheiranthus, Erysimum, Hesperis, Iberis, Lobularia, Lunaria, Malcolmia, and Matthiola. 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.