Solanaceae
Temporal range: Early Eocene to Recent, 52–0 Ma
A flowering Brugmansia suaveolens
from the US Botanic Garden
Scientific classification e
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Solanales
Family: Solanaceae
Juss.
Subfamilies[1]
Fruits including tomatoes, tomatillos, eggplant, bell peppers and chili peppers, all of which are closely related members of the Solanaceae.
Fruits including tomatoes, tomatillos, eggplant, bell peppers and chili peppers, all of which are closely related members of the Solanaceae.

The Solanaceae /sɒləˈnʃ/,[citation needed] or nightshades, are a family of flowering plants that ranges from annual and perennial herbs to vines, lianas, epiphytes, shrubs, and trees, and includes a number of agricultural crops, medicinal plants, spices, weeds, and ornamentals. Many members of the family contain potent alkaloids, and some are highly toxic, but many—including tomatoes, potatoes, eggplant, bell and chili peppers—are used as food. The family belongs to the order Solanales, in the asterid group and class Magnoliopsida (dicotyledons).[2] The Solanaceae consists of about 98 genera and some 2,700 species,[3] with a great diversity of habitats, morphology and ecology.

The name Solanaceae derives from the genus Solanum. The etymology of the Latin word is unclear. The name may come from a perceived resemblance of certain solanaceous flowers to the sun and its rays. At least one species of Solanum is known as the "sunberry". Alternatively, the name could originate from the Latin verb solare, meaning "to soothe", presumably referring to the soothing pharmacological properties of some of the psychoactive species of the family.

The family has a worldwide distribution, being present on all continents except Antarctica. The greatest diversity in species is found in South America and Central America. In 2017, scientists reported on their discovery and analysis of a fossil tomatillo, Physalis infinemundi, found in the Patagonian region of Argentina, dated to 52 million years B.P. The finding has pushed back the earliest appearance of the plant family Solanaceae.[4] As tomatillos likely developed later than other nightshades, this may mean that the Solanaceae may have first developed during the Mesozoic Era.[5]

The Solanaceae include a number of commonly collected or cultivated species. The most economically important genus of the family is Solanum, which contains the potato (S. tuberosum, in fact, another common name of the family is the "potato family"), the tomato (S. lycopersicum), and the eggplant or aubergine (S. melongena). Another important genus, Capsicum, produces both chili peppers and bell peppers.

The genus Physalis produces the so-called groundcherries, as well as the tomatillo (Physalis philadelphica), Physalis peruviana (Cape gooseberry) and Physalis alkekengi (Chinese lantern). The genus Lycium contains the boxthorns and the goji berry, Lycium barbarum. Nicotiana contains, among other species, tobacco. Some other important members of Solanaceae include a number of ornamental plants such as Petunia, Browallia, and Lycianthes, and sources of psychoactive alkaloids, Datura, Mandragora (mandrake), and Atropa belladonna (deadly nightshade). Certain species are widely known for their medicinal uses, their psychotropic effects, or for being poisonous.[6]

Most of the economically important genera are contained in the subfamily Solanoideae, with the exceptions of tobacco (Nicotiana tabacum, Nicotianoideae) and petunia (Petunia × hybrida, Petunioideae).

Many of the Solanaceae, such as tobacco and petunia, are used as model organisms in the investigation of fundamental biological questions at the cellular, molecular, and genetic levels.

Etymology and pronunciation

The name "Solanaceae" (US: /ˌsləˈnsi, -siˌ, -siˌ, -siˌ/) comes to international scientific vocabulary from New Latin, from Solanum, the type genus, + -aceae,[7] a standardized suffix for plant family names in modern taxonomy. The genus name comes from the Classical Latin word solanum, referring to nightshades (especially Solanum nigrum), "probably from sol, 'sun', + -anum, neuter of -anus."[7]

Description

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Illustration of Solanum dulcamara, 1.- Flower, 2.- Flower in longitudinal section, without the petals; 3.- Androecium; 4.- Ovary, in transverse section; 5.- Seed viewed from above; 6.- Seed in transverse section, note the curved embryo surrounding the endosperm; A.- Branch with leaves and flowers; B.- Stem with immature and mature fruit
Illustration of Solanum dulcamara, 1.- Flower, 2.- Flower in longitudinal section, without the petals; 3.- Androecium; 4.- Ovary, in transverse section; 5.- Seed viewed from above; 6.- Seed in transverse section, note the curved embryo surrounding the endosperm; A.- Branch with leaves and flowers; B.- Stem with immature and mature fruit

Plants in the Solanaceae can take the form of herbs, shrubs, trees, vines and lianas, and sometimes epiphytes. They can be annuals, biennials, or perennials, upright or decumbent. Some have subterranean tubers. They do not have laticifers, nor latex, nor coloured saps. They can have a basal or terminal group of leaves or neither of these types. The leaves are generally alternate or alternate to opposed (that is, alternate at the base of the plant and opposed towards the inflorescence). The leaves can be herbaceous, leathery, or transformed into spines. The leaves are generally petiolate or subsessile, rarely sessile. They are frequently inodorous, but some are aromatic or fetid. The foliar lamina can be either simple or compound, and the latter can be either pinnatifid or ternate. The leaves have reticulated venation and lack a basal meristem. The laminae are generally dorsiventral and lack secretory cavities. The stomata are generally confined to one of a leaf's two sides; they are rarely found on both sides.

The flowers are generally hermaphrodites, although some are monoecious, andromonoecious, or dioecious species (such as some Solanum or Symonanthus). Pollination is entomophilous. The flowers can be solitary or grouped into terminal, cymose, or axillary inflorescences. The flowers are medium-sized, fragrant (Nicotiana), fetid (Anthocercis), or inodorous. The flowers are usually actinomorphic, slightly zygomorphic, or markedly zygomorphic (for example, in flowers with a bilabial corolla in Schizanthus species). The irregularities in symmetry can be due to the androecium, to the perianth, or both at the same time. In the great majority of species, the flowers have a differentiated perianth with a calyx and corolla (with five sepals and five petals, respectively) an androecium with five stamens and two carpels forming a gynoecium with a superior ovary[8] (they are therefore referred to as pentamers and tetracyclic). The stamens are epipetalous and are typically present in multiples of four or five, most commonly four or eight. They usually have a hypogynous disk. The calyx is gamosepalous (as the sepals are joined forming a tube), with the (4)5(6) segments equal, it has five lobes, with the lobes shorter than the tube, it is persistent and often accrescent. The corolla usually has five petals that are also joined forming a tube. Flower shapes are typically rotate (wheel-shaped, spreading in one plane, with a short tube) or tubular (elongated cylindrical tube), campanulated or funnel-shaped.

The androecium has (2)(4)5(6) free stamens within it, oppositsepals (that is, they alternate with the petals), they are usually fertile or, in some cases (for example in Salpiglossideae) they have staminodes. In the latter case, there is usually either one staminode (Salpiglossis) or three (Schizanthus). The anthers touch on their upper end forming a ring, or they are completely free, dorsifixed, or basifixed with poricide dehiscence or through small longitudinal cracks. The stamen's filament can be filliform or flat. The stamens can be inserted inside the coralline tube or exserted. The plants demonstrate simultaneous microsporogenesis, the microspores are tetrad, tetrahedral, or isobilateral. The pollen grains are bicellular at the moment of dehiscence, usually open and angular.

The gynoecium is bicarpelar (rarely three- or five-locular) with a superior ovary and two locules, which may be secondarily divided by false septa, as is the case for Nicandreae and Datureae. The gynoecium is located in an oblique position relative to the flower's median plane. They have one style and one stigma; the latter is simple or bilobate. Each locule has one to 50 ovules that are anatropous or hemianatropous with axillar placentation. The development of the embryo sack can be the same as for Polygonum or Allium species. The embryo sack's nuclear poles become fused before fertilization. The three antipodes are usually ephemeral or persistent as in the case of Atropa. The fruit can be a berry as in the case of the tomato or wolfberry a dehiscent capsule as in Datura, or a drupe. The fruit has axial placentation. The capsules are normally septicidal or rarely loculicidal or valvate. The seeds are usually endospermic, oily (rarely starchy), and without obvious hairs. The seeds of most Solanaceae are round and flat, about 2–4 mm (0.079–0.157 in) in diameter. The embryo can be straight or curved, and has two cotyledons. Most species in the Solanaceae have 2n=24 chromosomes,[9] but the number may be a higher multiple of 12 due to polyploidy. Wild potatoes, of which there are about 200, are predominantly diploid (2 × 12 = 24 chromosomes), but triploid (3 × 12 = 36 chromosomes), tetraploid (4 × 12 = 48 chromosomes), pentaploid (5 × 12 = 60) and even hexaploid (6 × 12 = 72 chromosome) species or populations exist. The cultivated species Solanum tuberosum has 4 × 12 = 48 chromosomes. Some Capsicum species have 2 × 12 = 24 chromosomes, while others have 26 chromosomes.

Diversity of characteristics

Despite the previous description, the Solanaceae exhibit a large morphological variability, even in their reproductive characteristics. Examples of this diversity include:[10][11]

In general, the Solanaceae have a gynoecium (the female part of the flower) formed of two carpels. However, Melananthus has a monocarpelar gynoecium, there are three or four carpels in Capsicum, three to five in Nicandra, some species of Jaborosa and Trianaea and four carpels in Iochroma umbellatum.

The number of locules in the ovary is usually the same as the number of carpels. However, some species occur in which the numbers are not the same due to the existence of false septa (internal walls that subdivide each locule), such as in Datura and some members of the Lycieae (the genera Grabowskia and Vassobia).

The ovules are generally inverted, folded sharply backwards (anatropous), but some genera have ovules that are rotated at right angles to their stalk (campilotropous) as in Phrodus, Grabowskia or Vassobia), or are partially inverted (hemitropous as in Cestrum, Capsicum, Schizanthus and Lycium). The number of ovules per locule also varies from a few (two pairs in each locule in Grabowskia, one pair in each locule in Lycium) and very occasionally only one ovule is in each locule as for example in Melananthus.

The fruits of the great majority of the Solanaceae are berries or capsules (including pyxidia) and less often drupes. Berries are common in the subfamilies Cestroideae, Solanoideae (with the exception of Datura, Oryctus, Grabowskia and the tribe Hyoscyameae) and the tribe Juanulloideae (with the exception of Markea). Capsules are characteristic of the subfamilies Cestroideae (with the exception of Cestrum) and Schizanthoideae, the tribes Salpiglossoideae and Anthocercidoideae, and the genus Datura. The tribe Hyoscyameae has pyxidia. Drupes are typical of the Lycieae tribe and in Iochrominae.[12]

Alkaloids

Alkaloids are nitrogenous organic substances produced by plants as a secondary metabolite and which have an intense physiological action on animals even at low doses.[citation needed] Solanaceae are known for having a diverse range of alkaloids. To humans, these alkaloids can be desirable, toxic, or both. The tropanes are the most well-known of the alkaloids found in the Solanaceae. The plants that contain these substances have been used for centuries as poisons. However, despite being recognized as poisons, many of these substances have invaluable pharmaceutical properties. Many species contain a variety of alkaloids that can be more or less active or poisonous, such as scopolamine, atropine, hyoscyamine, and nicotine. They are found in plants such as henbane (Hyoscyamus albus), belladonna (Atropa belladonna), jimson weed (Datura stramonium), mandrake (Mandragora autumnalis), tobacco, and others. Some of the main types of alkaloids are:

Chemical structure of solanine
Chemical structure of solanine
Chemical structure of the tropanes.
Chemical structure of the tropanes.
Chemical structure of nicotine.
Chemical structure of nicotine.
Chemical structure of capsaicin
Chemical structure of capsaicin

Distribution

Map showing the distribution of the Solanaceae throughout the world (light green areas)
Map showing the distribution of the Solanaceae throughout the world (light green areas)

Even though members of the Solanaceae are found on all continents except Antarctica, the greatest variety of species are found in Central America and South America. Centers of diversity also occur in Australia and Africa. Solanaceae occupy a great number of different ecosystems, from deserts to rainforests, and are often found in the secondary vegetation that colonizes disturbed areas. In general, plants in this family are of tropical and temperate distribution.

Plant host

The potato tuber moth (Phthorimaea operculella) is an oligophagous insect that prefers to feed on plants of the family Solanaceae, especially the potato plant (Solanum tuberosum). Female P. operculella use the leaves to lay their eggs and the hatched larvae will eat away at the mesophyll of the leaf. After feeding on the foliage, the larvae will then delve down and feed on the tubers and roots of the plant.[24]

Taxonomy

The following taxonomic synopsis of the Solanaceae, including subfamilies, tribes and genera, is based on the most recent molecular phylogenetics studies of the family:[2][3][25][26]

Cestroideae (Browallioideae)

Cestrum elegans, ( subfamily : Cestroideae ), a shrub used as an ornamental.
Cestrum elegans, ( subfamily : Cestroideae ), a shrub used as an ornamental.
Browallia americana
Browallia americana
Streptosolen jamesonii, Cultivated plant, Chelsea Physic Garden London UK.
Flower of Salpiglossis sinuata, Botanischer Garten Jena, Germany
Flower of Salpiglossis sinuata, Botanischer Garten Jena, Germany

This subfamily is characterised by the presence of pericyclic fibres, an androecium with four or five stamens, frequently didynamous. The basic chromosome numbers are highly variable, from x=7 to x=13. The subfamily consists of eight genera (divided into three tribes) and about 195 species distributed throughout the Americas. The genus Cestrum is the most important, as it contains 175 of the 195 species in the subfamily. The Cestreae tribe is unusual because it includes taxa with long chromosomes (from 7.21 to 11.511 µm in length), when the rest of the family generally possesses short chromosomes (for example between 1.5 and 3.52 µm in the Nicotianoideae)

Goetzeoideae

Goetzea elegans ( subfamily Goetzeoideae ) in bud and flower, South Miami, Florida United States.
Goetzea elegans ( subfamily Goetzeoideae ) in bud and flower, South Miami, Florida United States.
Espadaea amoena ( subfamily Goetzeoideae ).
Espadaea amoena ( subfamily Goetzeoideae ).

This subfamily is characterized by the presence of drupes as fruit and seeds with curved embryos and large fleshy cotyledons. The basic chromosome number is x=13. It includes four genera and five species distributed throughout the Greater Antilles. Some authors suggest their molecular data indicate the monotypic genera Tsoala Bosser & D'Arcy should be included in this subfamily, endemic to Madagascar, and Metternichia to the southeast of Brazil. Goetzeaceae Airy Shaw is considered as a synonym of this subfamily.[27]

Nicotianoideae

Tobacco inflorescence, Nicotiana tabacum
Tobacco inflorescence, Nicotiana tabacum

Petunioideae

Brunfelsia pauciflora subfamily Petunioideae
Nierembergia frutescens subfamily Petunioideae
Nierembergia frutescens subfamily Petunioideae
Petunia exserta
Petunia exserta

Molecular phylogenetics indicates that Petunioideae is the sister clade of the subfamilies with chromosome number x=12 (Solanoideae and Nicotianoideae). They contain calistegins, alkaloids similar to the tropanes. The androecium is formed of four stamens (rarely five), usually with two different lengths. The basic chromosome number of this subfamily can be x=7, 8, 9 or 11. It consists of 13 genera and some 160 species distributed throughout Central and South America. Molecular data suggest the genera originated in Patagonia. Benthamiella, Combera, and Pantacantha form a clade that can be categorized as a tribe (Benthamielleae) that should be in the subfamily Goetzeoideae.

Schizanthoideae

Zygomorphic flowers, with bilabiate corolla of Schizanthus pinnatus, a schizanthoidea ornamental
Zygomorphic flowers, with bilabiate corolla of Schizanthus pinnatus, a schizanthoidea ornamental

The Schizanthoideae include annual and biennial plants with tropane alkaloids, without pericyclic fibres, with characteristic hair and pollen grains. The flowers are zygomorphic. The androecium has two stamens and three staminodes, anther dehiscence is explosive. In terms of fruit type, the Schizanthoidae retain the plesiomorphic fruit form of the family Solanaceae, capsules, which rely on an anemochorous, abiotic form of dispersal. This is present in Schizanthoidae due both to the genetic constraints of early divergence (see below) as well as Schizanthus evolution and presence in open habitats.[31] The embryo is curved. The basic chromosome number is x=10. Schizanthus is a somewhat atypical genus among the Solanaceae due to its strongly zygomorphic flowers and basic chromosome number. Morphological and molecular data suggest Schizanthus is a sister genus to the other Solanaceae and diverged early from the rest, probably in the late Cretaceous or in the early Cenozoic, 50 million years ago.[25][26] The great diversity of flower types within Schizanthus has been the product of the species’ adaptation to the different types of pollinators that existed in the Mediterranean, high alpine, and desert ecosystems then present in Chile and adjacent areas of Argentina.[32]

Schwenckioideae

Annual plants with pericyclic fibres, their flowers are zygomorphic, the androecium has four didynamous stamens or three staminodes; the embryo is straight and short. The basic chromosome number is x=12. It includes four genera and some 30 species distributed throughout South America.

Solanoideae

Capsicum frutescens cultivar "tabasco", a solanoidea
Capsicum frutescens cultivar "tabasco", a solanoidea
Atropa belladonna ( Deadly Nightshade ) flower
Atropa belladonna ( Deadly Nightshade ) flower
Black Henbane (Hyoscyamus niger)
Black Henbane (Hyoscyamus niger)
Latua pubiflora subfamily Solanoideae
Nicandra physalodes flower
Solandra maxima flower
Solandra maxima flower
In the fruit of Physalis peruviana (Cape gooseberry), the persistent calyx surrounds the fruit.
In the fruit of Physalis peruviana (Cape gooseberry), the persistent calyx surrounds the fruit.
Eriolarynx australis ( known formerly as Iochroma australe ) flower, cultivated plant, UBC Botanical Garden, British Columbia.
Eriolarynx australis ( known formerly as Iochroma australe ) flower, cultivated plant, UBC Botanical Garden, British Columbia.
Jaltomata procumbens flower
Solanum bonariense flower
Solanum bonariense flower
Flower of Solanum betaceum (=Cyphomandra betacea)
Flower of Solanum betaceum (=Cyphomandra betacea)
Acnistus arborescens flower
Scopolia carniolica flower

Incertae sedis

Sclerophylax kurtzii.
Sclerophylax kurtzii.

The following genera have not yet been placed in any of the recognized subfamilies within the solanaceas (incertae sedis).

Genera and distribution of species

Flowers and foliage of Cestrum parqui.
Flowers and foliage of Cestrum parqui.
Metternichia principis

The Solanaceae contain 98 genera and some 2,700 species. Despite this immense richness of species, they are not uniformly distributed between the genera. The eight most important genera contain more than 60% of the species, as shown in the table below. Solanum – the genus that typifies the family - includes nearly 50% of the total species of the solanaceas.

Genera Approximate number of species
Solanum 1,330
Lycianthes 200
Cestrum 150
Nolana 89
Physalis 85
Lycium 85
Nicotiana 76
Brunfelsia 45
Estimated number of species in the family 2,700

Economic importance

Pink, double-flowered Brugmansia cultivar
Pink, double-flowered Brugmansia cultivar
Triple-flowered Datura metel 'Fastuosa': ancient cultivar created from Datura innoxia by Pre-Columbian horticulturalists in the Greater Antilles
Triple-flowered Datura metel 'Fastuosa': ancient cultivar created from Datura innoxia by Pre-Columbian horticulturalists in the Greater Antilles
Petunia × atkinsiana, a herbaceous annual  commonly cultivated as a summer bedding plant
Petunia × atkinsiana, a herbaceous annual commonly cultivated as a summer bedding plant

The family Solanaceae contains such important food species as the potato (Solanum tuberosum), the tomato (Solanum lycopersicum), the pepper (Capsicum annuum) and the aubergine or eggplant (Solanum melongena). Nicotiana tabacum, originally from South America, is now cultivated throughout the world to produce tobacco. Many solanaceas are important weeds in various parts of the world. Their importance lies in the fact that they can host pathogens or diseases of the cultivated plants, therefore their presence increases the loss of yield or the quality of the harvested product. An example of this can be seen with Acnistus arborescens and Browalia americana that host thrips, which cause damage to associated cultivated plants,[43] and certain species of Datura that play host to various types of virus that are later transmitted to cultivated solanaceas.[44] Some species of weeds such as, Solanum mauritianum in South Africa represent such serious ecological and economic problems that studies are being carried out with the objective of developing a biological control through the use of insects.[45]

A wide variety of plant species and their cultivars belonging to the Solanaceae are grown as ornamental trees, shrubs, annuals and herbaceous perennials[46] Examples include Brugmansia x candida ("Angel’s Trumpet") grown for its large pendulous trumpet-shaped flowers, or Brunfelsia latifolia, whose flowers are very fragrant and change colour from violet to white over a period of 3 days. Other shrub species that are grown for their attractive flowers are Lycianthes rantonnetii (Blue Potato Bush or Paraguay Nightshade) with violet-blue flowers and Nicotiana glauca ("Tree Tobacco") Other solanaceous species and genera that are grown as ornamentals are the petunia (Petunia × hybrida), Lycium, Solanum, Cestrum, Calibrachoa × hybrida and Solandra. There is even a hybrid between Petunia and Calibrachoa (which constitutes a new nothogenus called × Petchoa G. Boker & J. Shaw) that is being sold as an ornamental.[47][48] Many other species, in particular those that produce alkaloids, are used in pharmacology and medicine (Nicotiana, Hyoscyamus, and Datura).[6]

Solanaceae and the genome

Many of the species belonging to this family, among them tobacco and the tomato, are model organisms that are used for research into fundamental biological questions. One of the aspects of the solanaceas’ genomics is an international project that is trying to understand how the same collection of genes and proteins can give rise to a group of organisms that are so morphologically and ecologically different. The first objective of this project was to sequence the genome of the tomato. In order to achieve this each of the 12 chromosomes of the tomato's haploid genome was assigned to different sequencing centres in different countries. So chromosomes 1 and 10 were sequenced in the United States, 3 and 11 in China, 2 in Korea, 4 in Britain, 5 in India, 7 in France, 8 in Japan, 9 in Spain and 12 in Italy. The sequencing of the mitochondrial genome was carried out in Argentina and the chloroplast genome was sequenced in the European Union.[49][50]

See also

References

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Further reading

Wikisource has the text of the 1911 Encyclopædia Britannica article "Nightshade".