58-664: See text Nicotianoideae is a subfamily within the family Solanaceae . Most genera are found in Australia, but they are also found in America and Africa. The subfamily contains eight genera and about 125 species, 90 of them are included in Nicotiana . Genera: Anthocercis Anthotroche Crenidium Cyphanthera Duboisia Grammosolen Nicotiana Symonanthus Solanaceae The Solanaceae ( / ˌ s ɒ l ə ˈ n eɪ s i . iː , - ˌ aɪ / ), or
116-495: A New World family. Mandragora is suggested to have originated around 20 million years ago, arriving in Eurasia through the agency of birds, with the main split between the species occurring around 10 million years ago. As of March 2015 , major online plant databases (such as Tropicos , The Plant List , and GRIN Taxonomy for Plants ) accept different numbers of species in the genus Mandragora . Three species are accepted in
174-463: A 1998 review of the genus and by GRIN. Other sources keep M. autumnalis and M. chinghaiensis as separate species. Like many members of the Solanaceae, species of Mandragora contain highly biologically active alkaloids, tropane alkaloids in particular. Hanuš et al. reviewed the phytochemistry of Mandragora species. More than 80 substances have been identified; their paper gives
232-694: 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
290-497: A berry ). The genus Mandragora was first used in 1753 by Carl Linnaeus in the first edition of Species Plantarum where the Mediterranean species Mandragora officinarum was described, which is thus the type species of the genus. (Linnaeus later changed his mind and in 1759 placed M. officinarum in the genus Atropa as A. mandragora . ) Jackson and Berry (1979) and Ungricht et al. (1998) have documented
348-656: 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 (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
406-731: 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
464-570: 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
522-449: 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. 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
580-549: 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 its opposite sepals (they alternate with
638-417: 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
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#1732863313942696-428: A rosette at the base of the plant. The flowers are sometimes borne on a short stalk ( scape ), and are solitary, with whorls of five parts. The sepals are joined at the base, as are the petals , both in the shape of a lobed bell. The stamens are shorter than the petals, joined to the floral tube towards the base. The ovary has two chambers ( locules ). After fertilization, a yellow or orange fruit forms ( botanically
754-525: 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 species belonging to the living genus Physalis , Physalis infinemundi , found in the Patagonian region of Argentina, dated to 52 million years ago. The finding has pushed back
812-492: Is a plant genus belonging to the nightshade family ( Solanaceae ). Members of the genus are known as mandrakes . Between three and five species are placed in the genus. The one or two species found around the Mediterranean constitute the mandrake of ancient writers such as Dioscorides . Two or three further species are found eastwards into China. All are perennial herbaceous plants , with large tap roots and leaves in
870-499: 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. Many other species, in particular those that produce alkaloids, are used in pharmacology and medicine ( Nicotiana , Hyoscyamus , and Datura ). 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
928-523: 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. The embryo is curved. The basic chromosome number
986-470: 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
1044-424: 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. The great diversity of flower types within Schizanthus has been
1102-433: Is x=12. It includes four genera and some 30 species distributed throughout South America. The following genera have not yet been placed in any of the recognized subfamilies within the solanaceas ( incertae sedis ) 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
1160-579: The Classical Latin word solanum , referring to nightshades (especially Solanum nigrum ), "probably from sol , 'sun', + -anum , neuter of -anus ." 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
1218-704: The Levant ( Mandragora autumnalis ) from those found in the rest of the Mediterranean area ( Mandragora officinarum ) suggest that there are two clades in the genus - one based in the Mediterranean and beyond to Turkmenistan and Iran, and one in the Sino-Himalayan region. A simplified cladogram based on these studies is shown below. In one of the studies, M. chinghaiensis was embedded within M. caulescens . Mandragora turcomanica Mandragora autumnalis Mandragora officinarum Mandragora caulescens Mandragora chinghaiensis The Solanaceae are primarily
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#17328633139421276-508: The cellular , molecular , and genetic levels. 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
1334-472: The nightshades , is 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
1392-485: The 1820s, Antonio Bertoloni used flowering time to name two species as Mandragora vernalis , the spring-flowering mandrake, and Mandragora autumnalis , the autumn-flowering mandrake. Identifying the former as Linnaeus's M. officinarum , works such as Flora Europaea listed two Mediterranean species of Mandragora : M. officinarum and M. autumnalis . Using statistical analysis of morphological characters, Ungricht et al. found no distinct clusters among
1450-510: 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). This subfamily is characterized by
1508-555: The Solanaceae have 2n=24 chromosomes , 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. Despite
1566-454: The Solanaceae, including subfamilies, tribes and genera, is based on the most recent molecular phylogenetics studies of the family: 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
1624-403: 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
1682-414: 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
1740-573: The case of Atropa . The fruit can be a berry as in the case of the tomato or wolfberry, or 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
1798-449: 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
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1856-493: The detailed chemical structure of 37 of them. The different parts of the plant contain different proportions and concentrations of alkaloids, with the roots having the highest concentrations. Alkaloids present include atropine , apoatropine , belladonnine , cuscohygrine , hyoscyamine , scopolamine (hyoscine), 3α-tigloyloxytropane , and 3α,6β-ditigloyloxytropane . M. caulescens and M. turcomanica are also reported to contain anisodamine . Clinical reports of
1914-411: The earliest appearance of the plant family Solanaceae. 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
1972-403: The effects of consumption of plants described as M. autumnalis ( M. offinarum sensu lato ) include severe symptoms similar to those of atropine poisoning, including blurred vision, dilation of the pupils ( mydriasis ), dryness of the mouth, difficulty in urinating, dizziness, headache, vomiting, blushing, and a rapid heart rate ( tachycardia ). Hyperactivity and hallucinations also occurred in
2030-519: 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 ) and Physalis peruviana (Cape gooseberry). Alkekengi officinarum (Chinese Lantern)
2088-573: The form of a rosette . Individual flowers are bell-shaped, whitish through to violet, and followed by yellow or orange berries . Like many members of the Solanaceae, species of Mandragora contain highly biologically active alkaloids that make the plants poisonous. Their roots in particular have a long use in traditional medicine . Mandrakes are involved in many myths and superstitions. Species of Mandragora are perennial herbaceous plants. They have large vertical tap roots, sometimes forked. Their stems are short or virtually absent. The leaves form
2146-498: 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. 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
2204-412: 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. 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. Solanaceae are known for having
2262-428: The main types of alkaloids are: 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
2320-535: The majority of patients. Mandragora species have a long use in traditional medicine, extracts being used for their real or supposed aphrodisiac, hypnotic, emetic, purgative, sedative, and pain-killing effects. Tropane alkaloids are known to be effective as analgesics and anaesthetics, and can be used to increase circulation and dilate pupils, among other effects. Hyoscine and anisodamine are used medicinally in China. Continued use of M. autumnalis in folk medicine
2378-454: The objective of developing a biological control through the use of insects. A wide variety of plant species and their cultivars belonging to the Solanaceae are grown as ornamental trees, shrubs, annuals and herbaceous perennials Examples include Brugmansia × 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
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2436-456: The order Solanales , in the asterid group and class Magnoliopsida ( dicotyledons ). The Solanaceae consists of about 98 genera and some 2,700 species, 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
2494-813: 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
2552-526: 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 filiform or flat. The stamens can be inserted inside
2610-470: 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
2668-664: The previous description, the Solanaceae exhibit a large morphological variability, even in their reproductive characteristics. Examples of this diversity include: 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
2726-447: 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. 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
2784-468: The same tribe or subtribe as at least the first of these genera. Molecular phylogenetic studies suggest that the genus belongs in the large subfamily Solanoideae, but that within this subfamily, it is one of a number of isolated genera with no immediate relatives. It has thus been placed in its own tribe, Mandragoreae. Within the genus, studies have used different circumscriptions of the Mediterranean mandrakes. Two studies that separate plants found in
2842-606: The southeast of Brazil . Goetzeaceae Airy Shaw is considered as a synonym of this subfamily. 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
2900-536: The species, as shown in the table below. Solanum – the genus that typifies the family – includes nearly 50% of the total species of the Solanacea. The name "Solanaceae" ( US : / ˌ s oʊ l ə ˈ n eɪ s i , - s i ˌ aɪ , - s i ˌ eɪ , - s i ˌ iː / ) comes to international scientific vocabulary from Neo-Latin , from Solanum , the type genus , + -aceae , a standardized suffix for plant family names in modern taxonomy. The genus name comes from
2958-594: The specimens they examined and concluded that Linnaeus's M. officinarum is a single, variable species. Other sources divide M. officinarum sensu lato differently. Plants from the western Mediterranean, from Turkey westwards to the Iberian peninsula and Morocco, are placed in M. officinarum ; plants from the eastern Mediterranean, from Syria to Israel, are placed in M. autumnalis . Traditionally, Mandragora has been considered to be closely related to Atropa and Lycium , being grouped together in
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#17328633139423016-436: The subsequent confusion over the number of Mediterranean species of Mandragora and their scientific names. Dioscorides was among those who distinguished between "male" and "female" mandrakes, a distinction used in 1764 when Garsault published the names Mandragora mas and Mandragora foemina . The size and shape of the fruit and the colour and time of appearance of the flowers have been used to distinguish possible species. In
3074-469: 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 Solanaceae family includes a number of commonly collected or cultivated species. The most economically important genus of
3132-465: 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. The following taxonomic synopsis of
3190-711: 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, and certain species of Datura that play host to various types of virus that are later transmitted to cultivated solanaceas. 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
3248-943: 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 . Mandragora (genus) See text Mandragora
3306-694: Was previously included in the genus Physalis (as Physalis alkekengi ), until molecular and genetic evidence placed it as the type species of a new genus. 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. This family has
3364-541: Was reported in Sicily in 2014. M. caulescens (as M. chinghaiensis ) has been listed as a Chinese medicinal plant needing conservation. The presence of deliriant and hallucinogenic alkaloids and the sometimes vaguely humanoid shape of their roots have led to mandrakes being associated with a variety of myths and superstitious practices throughout history. However, the plants used in this way are not always species of Mandragora ; for example, bryony ( Bryonia )
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