Difference between revisions of "Solanaceae"

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{{Redirect|Nightshade}}
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#redirect [[:Category:Solanaceae]]
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{{automatic taxobox
 
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|fossil_range = Early [[Eocene]] to [[Recent]], {{Fossil range|52|0}}
 
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|image = brugmansia_lg.jpg
 
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|image_caption = A flowering ''[[Brugmansia suaveolens]]'' <br>from the [[US Botanic Garden]]
 
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|taxon = Solanaceae
 
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|authority = [[Antoine Laurent de Jussieu|Juss.]]
 
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|subdivision_ranks = [[Family (biology)|Subfamilies]]
 
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|subdivision = [[Cestroideae]]<br>
 
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[[Goetzeoideae]]<br>
 
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[[Nicotianoideae]]<br>
 
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[[Petunioideae]]<br>
 
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[[Schizanthoideae]]<br>
 
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[[Schwenckioideae]]<br>
 
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[[Solanoideae]]<ref name="GRIN">{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/family.pl?1043 |title=''Solanaceae'' Juss., nom. cons. |work=[[Germplasm Resources Information Network]] |publisher=[[United States Department of Agriculture]] |date=2007-04-12 |accessdate=2009-04-16}}</ref>
 
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}}
 
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[[File:Solanums in a box.jpg|thumb|336x336px|Fruits including tomatoes, tomatillos, eggplant, bell peppers and chili peppers.]]
 
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The '''Solanaceae''', or '''nightshades''', are an economically important [[family (biology)|family]] of [[flowering plant]]s. The family ranges from annual and perennial herbs to vines, lianas, epiphytes, shrubs, and trees, and includes a number of important [[agricultural]] crops, medicinal plants, spices, weeds, and ornamentals. Many members of the family contain potent [[alkaloid]]s, and some are highly [[toxin|toxic]], but many, including tomatoes, potatoes, eggplant, bell/chili peppers, and tobacco are widely used by many people. The family belongs to the order [[Solanales]], in the asterid group and class [[Magnoliopsida]] ([[dicotyledon]]s).<ref name=ref_duplicada_2>{{cite book |last1=Olmstead |first1=R. G. |first2=J. A. |last2=Sweere |first3=R. E. |last3=Spangler |first4=L. |last4=Bohs |first5=J. D. |last5=Palmer |year=1999 |chapterurl=http://depts.washington.edu/phylo/OlmsteadPubs/Solanaceae_IV.pdf |chapter=Phylogeny and provisional classification of the Solanaceae based on chloroplast DNA |pages=111–37 |title=Solanaceae IV: advances in biology and utilization |editor1-first=M. |editor1-last=Nee |editor2-first=D. E. |editor2-last=Symon |editor3-first=R. N. |editor3-last=Lester |editor4-first=J. P. |editor4-last=Jessop |publisher=The Royal Botanic Gardens}}</ref> The Solanaceae consists of about 98 genera and some 2,700 species,<ref name="Olmstead 2007">{{cite journal |last1=Olmstead |first1=R.G. |last2=Bohs |first2=L. |year=2007 |title=A Summary of molecular systematic research in Solanaceae: 1982-2006 |url=http://www.actahort.org/books/745/745_11.htm |journal=Acta Horticulturae |volume=745 |pages=255–68}}</ref> with a great diversity of [[habitats]], [[Morphology (biology)|morphology]] and [[ecology]].
 
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The name Solanaceae derives from the genus ''[[Solanum]]'', "the nightshade plant". 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 ''solari'', meaning "to soothe", presumably referring to the soothing [[pharmacology|pharmacological]] properties of some of the [[psychoactive]] [[species]] of the family.
 
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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 [[Physalis infinemundi|fossil tomatillo]] found in the [[Patagonia]]n region of Argentina, dated to 52 million years B.P.  The finding has pushed back the earliest appearance of the Solanaceae plant family.<ref>[http://science.sciencemag.org/content/355/6320/71.full "Eocene lantern fruits from [[Gondwana]]n Patagonia and the early origins of Solanaceae", Wilf et al, ''Science'', 06 Jan 2017, Vol. 355, Issue 6320, pp. 71-75, DOI: 10.1126/science.aag2737]</ref> As tomatillos likely developed later than other nightshades, this may mean that the Solanaceae may have first developed during the [[Mesozoic|Mesozoic Era]].<ref>{{Cite news|url=http://www.npr.org/sections/thesalt/2017/01/10/509171881/52-million-year-old-tomatillo-fossil-rewrites-veggie-history|title=52 Million-Year-Old Tomatillo Fossils Rewrite Veggie History|newspaper=NPR.org|language=en|access-date=2017-01-20}}</ref>
 
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The Solanaceae include a number of commonly collected or cultivated species. The most economically important genus of the family{{Citation needed|date=July 2015}} 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 pepper]]s and [[bell pepper]]s.
 
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The genus ''[[Physalis]]'' produces the so-called groundcherries, as well as the [[tomatillo]] (''Physalis philadelphica''), the [[Physalis peruviana|Cape gooseberry]]  and the [[Physalis alkekengi|Chinese lantern]].  The genus ''[[Lycium]]'' contains the boxthorns and the [[wolfberry]] ''Lycium barbarum''.  ''[[Nicotiana]]'' contains, among other species, [[tobacco]].
 
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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 (genus)|Mandragora]]'' (mandrake), and ''[[Atropa belladonna]]'' (deadly nightshade).  Certain species are widely known for their medicinal uses, their psychotropic effects, or for being poisonous.
 
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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).
 
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Many of the Solanaceae, such as tobacco and petunia, are used as [[model organism]]s in the investigation of fundamental biological questions at the [[cell (biology)|cellular]], [[molecular biology|molecular]], and [[Genetics|genetic]] levels.
 
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==Etymology and pronunciation==
 
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The name "Solanaceae" ({{IPAc-en|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 [[New Latin]], from ''[[Solanum]]'', the [[type genus]], + ''[[wikt:-aceae#Suffix|-aceae]]'',<ref name="MWU">{{Citation |author=Merriam-Webster |authorlink=Merriam-Webster |title=Merriam-Webster's Unabridged Dictionary |publisher=Merriam-Webster |url=http://unabridged.merriam-webster.com/unabridged/ |postscript=.}}</ref> a standardized [[suffix]] for plant family names in modern taxonomy. The genus name comes from the [[Classical Latin]] word ''[[wikt:solanum|solanum]]'', referring to nightshades (especially ''[[Solanum nigrum]]''), "probably from [[wikt:sol#Latin|sol]], 'sun', + ''[[wikt:-anum#Latin|-anum]]'', neuter of ''[[wikt:-anus#Latin|-anus]]''."<ref name="MWU"/>
 
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== Description ==
 
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{{refimprove section|date=April 2015}}
 
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[[File:Illustration Solanum dulcamara0 clean.png|thumb|250px|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]]
 
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Plants in the "Solanaceae" can take the form of herbs, [[shrub]]s, [[tree]]s, [[vine]]s and lianas, and sometimes [[epiphytes]]. They can be [[Annual plant|annuals]], [[Biennial plant|biennials]], or [[Perennial plant|perennials]], upright or decumbent. Some have subterranean [[tuber]]s.  They do not have [[laticifer]]s, nor [[latex]], nor coloured [[sap]]s.
 
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They can have a basal or terminal group of leaves or neither of these types. The [[Leaf|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 [[Thorns, spines and prickles|spines]]. The leaves are generally [[Petiole (botany)|petiolate]] or subsessile, rarely sessile. They are frequently inodorous, but on occasions, they 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 [[stoma]]ta are generally confined to one of a leaf's two sides; they are rarely found on both sides.
 
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The [[flower]]s are generally [[hermaphrodite]]s, although some are [[monoecious]], [[andromonoecious]], or [[Dioecy|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 [[Floral symmetry#Actinomorphic|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 (flower)|corolla]] (with five sepals and five petals, respectively) an androecium with five [[stamen]]s and two [[carpel]]s forming a [[gynoecium]] with a superior [[ovary (botany)|ovary]]<ref>{{cite web | url = http://www.efloras.org/florataxon.aspx?flora_id=5&taxon_id=10828 | title = Solanaceae | author = Yasin J. Nasir | work = Flora of Pakistan }}</ref> (they are therefore referred to as pentamers and tetracyclic). The [[stamen]]s are [[Stamen#Descriptive terms|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 together 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 together 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.
 
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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 (botany)|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.
 
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The gynoecium is bicarpelar (rarely three- or five-locular) with a [[Hypogynous#Superior ovary|superior]] [[ovary (plants)|ovary]] and two [[locule]]s, which may be secondarily divided by false [[Septum|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 (botany)|style]] and one [[Stigma (botany)|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 [[Ovule|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 (botany)|berry]] as in the case of the tomato or wolfberry a [[Dehiscence (botany)|dehiscent]] [[capsule (fruit)|capsule]] as in ''Datura'', or a [[drupe]]. The fruit has [[Placentation#Placentation in plants|axial placentation]].  The capsules are normally septicidal or rarely loculicidal or valvate. The [[seed]]s are usually endospermic, oily (rarely starchy), and without obvious hairs. The seeds of most Solanaceae are round and flat, about {{convert|2|-|4|mm|in|abbr=on}} in diameter. The embryo can be straight or curved, and has two cotyledons. Most species in the Solanaceae have 2n=24 [[chromosome]]s,<ref name="Fujii1934">{{cite book|last=Fujii|first=Kenjiro|title=Cytologia|url=https://books.google.com/books?id=If8ZAAAAIAAJ|year=1934|publisher=Botanical Institute|page=281}}</ref> but the number may be a higher multiple of 12 due to [[polyploidy]]. Wild [[potato]]es, 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.
 
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=== Diversity of characteristics ===
 
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Despite the previous description, the Solanaceae exhibit a large morphological variability, even in their reproductive characteristics.  Examples of this diversity include:<ref name="HUNZIKER79">[[Armando Theodoro Hunziker|Hunziker, A.T.]] 1979: South American Solanaceae: a synoptic review. In: [[William Gerald D'Arcy|D'ARCY, W.G.]], 1979: The Biology and Taxonomy of the Solanaceae. Linn. Soc. Symp. Ser. 7: p 48-85. Linnean Soc. & Academic Press; London.</ref><ref name="Balken">Balken, J.A. THE PLANT FAMILY SOLANACEAE:
 
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FRUITS IN SOLANACEAE [http://www.hvanbalken.com/fruit.html]</ref>
 
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* The number of carpels that form the gynoecium
 
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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''.
 
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* The number of locules in the ovary
 
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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'').
 
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* Type of ovules and their number
 
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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''.
 
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* The type of fruit
 
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The fruits of the great majority of the Solanaceae are berries or capsules (including pyxidia) and less often drupes.
 
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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]]'').
 
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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.
 
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Drupes are typical of the Lycieae tribe and in Iochrominae.
 
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== Alkaloids ==
 
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[[Alkaloid]]s are nitrogenous organic substances produced by plants as a [[Plant secondary metabolism|secondary metabolite]] and which have an intense physiological action on animals even at low doses.{{citation needed|date=September 2014}}  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 the henbane (''Hyoscyamus albus''), belladonna (''[[Atropa belladonna]]''), datura or jimson (''[[Datura stramonium]]''), mandrake (''[[Mandragora autumnalis]]''), tobacco, and others. Some of the main types of alkaloids are:
 
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[[File:Solanine.svg|thumb|Chemical structure of solanine]]
 
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* [[Solanine]]: A [[toxic]] [[glycoalkaloid]] with a bitter taste, it has the formula C<sub>45</sub>H<sub>73</sub>NO<sub>15</sub>. It is formed by the alkaloid solanidine with a [[carbohydrate]] side chain. It is found in leaves, fruit, and tubers of various Solanaceae such as the potato and tomato. Its production is thought to be an adaptive defence strategy against herbivores. [[Substance intoxication]] from solanine is characterized by gastrointestinal disorders ([[diarrhoea]], [[vomiting]], abdominal pain) and neurological disorders ([[hallucination]]s and [[headache]]). The [[median lethal dose]] is between 2 and 5&nbsp;mg per kg of body weight. Symptoms become manifest 8 to 12 hr after ingestion. The amount of these glycoalkaloids in potatoes, for example, varies significantly depending of environmental conditions during their cultivation, the length of storage, and the variety. The average glycoalkaloid concentration is 0.075&nbsp;mg/g of potato.<ref name="Zeiger">Zeiger, E. 1998. Solanine and Chaconine. Review of Toxicological Literature. Integrated Laboratory Systems, USA.[http://ntp.niehs.nih.gov/ntp/htdocs/Chem_Background/ExSumPdf/ChaconineSolanine.pdf]</ref> Solanine has occasionally been responsible for poisonings in people who ate berries from species such as ''[[Solanum nigrum]]'' or ''[[Solanum dulcamara]]'', or green potatoes.<ref>{{cite journal | title = Solanine poisoning | journal = Br Med J. | date = 1979-12-08 | volume = 2 | issue = 6203 | pages = 1458–9 | pmc = 1597169 |pmid=526812 | doi=10.1136/bmj.2.6203.1458-a}}</ref><ref>{{cite journal | title = A Fatal Case of Solanine Poisoning |vauthors=Alexander RF, Forbes GB, Hawkins ES | journal = Br Med J. | date = 1948-09-11 | volume = 2 | issue = 4575 | pages = 518 | pmc = 2091497 |pmid=18881287 | doi = 10.1136/bmj.2.4575.518 }}</ref>
 
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[[File:Tropane svg.svg|thumb|Chemical structure of the tropanes.]]
 
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* [[Tropane]]s: The term "tropane" comes from a [[genus]] in which they are found, ''[[Atropa]]'' (the [[Atropa belladonna|belladonna]] genus). ''Atropa'' is named after the [[Moirai|Greek Fate]], [[Atropos]], who cut the thread of life. This nomenclature reflects its toxicity and lethality. They are bicyclic organic nitrogen compounds ([[IUPAC]] nomenclature: 8-Methyl-8-azabicyclo[3.2.1]octane), with the chemical formula of C<sub>8</sub>H<sub>15</sub>N. These alkaloids include, among others, atropine, cocaine, scopolamine, and hyoscyamine. They are found in various species, such as mandrake (''Mandragora autumnalis''), black henbane or stinking nightshade (''[[Hyoscyamus niger]]''), belladonna (''[[Atropa belladonna]]'') the stramonium (''[[Datura stramonium]]'') and ''[[Brugmansia]]'' species, as well as many others in the Solanaceae family.<ref>{{cite journal |vauthors=Griffin WJ, Lin GD |title=Chemotaxonomy and geographical distribution of tropane alkaloids |journal=Phytochemistry |volume=53 |issue=6 |pages=623–37 |date=March 2000 |pmid=10746874 |doi=10.1016/S0031-9422(99)00475-6}}</ref> [[Pharmacologically]], they are the most powerful known [[anticholinergic]]s in existence, meaning they inhibit the [[neurological]] signals transmitted by the [[endogenous]] [[neurotransmitter]], [[acetylcholine]]. More commonly, they can halt many types of [[allergic]] reactions. Symptoms of overdose may include [[xerostomia|dry mouth]], [[mydriasis|dilated pupils]], [[ataxia]], [[urinary retention]], [[hallucination]]s, [[seizure|convulsions]], [[coma]], and death. Atropine, a commonly used [[ophthalmology|ophthalmological]] agent,  dilates the pupils and thus facilitates examination of the interior of the eye.  In fact, juice from the berries of ''A. belladonna'' were used by Italian courtesans during the [[Renaissance]] to exaggerate the size of their eyes by causing the dilation of their pupils. Despite the extreme toxicity of the tropanes, they are useful drugs when administered in extremely small dosages. They can reverse [[cholinergic]] poisoning, which can be caused by overexposure to organophosphate [[insecticide]]s and [[chemical warfare]] agents such as [[sarin]] and [[VX (nerve agent)|VX]].  [[Scopolamine]] (found in ''Hyoscyamus muticus'' and ''Scopolia  atropioides''), is used as an [[antiemetic]] against [[motion sickness]] or for people suffering from [[nausea]] as a result of receiving [[chemotherapy]].<ref name="Sneden ">Sneden, A. The tropane alkaloids. Medicinal Chemistry and Drug Design. Virginia Commonwealth University [http://www.people.vcu.edu/~asneden/tropane%20alkaloids.pdf]{{unreliable source?|date=January 2015}}{{dead link|date=January 2015}}</ref><ref name="Evans ">Evans, W.C. 1979. Tropane alkaloids of the Solanaceae. En: HAWKES, LESTER and SHELDING (eds.). The biology and taxonomy of the Solanaceae. Linn. Soc. Symp. Ser. 7:241-254. Linnean Soc. & Academic Press., London.</ref> Scopolamine and hyoscyamine are the most widely used tropane alkaloids in pharmacology and medicine due to their effects on the parasympathetic nervous system. Atropine has a [[stimulant]] effect on the [[central nervous system]] and heart, whereas scopolamine has a [[sedative]] effect. These alkaloids cannot be substituted by any other class of compounds, so they are still in demand. This is one of the reasons for the development of an active field of research into the metabolism of the alkaloids, the enzymes involved, and the genes that produce them. Hyoscyamine 6-β hydroxylase, for example, catalyses the hydroxylation of hyoscyamine that leads to the production of scopolamine at the end of the tropane’s biosynthetic pathway. This enzyme has been isolated and the corresponding gene cloned from three species: ''H. niger'', ''A. belladonna'' and '' B. candida''.<ref name="MATSUDA">{{cite journal |first1=Jun |last1=Matsuda |first2=Souichi |last2=Okabe |first3=Takashi |last3=Hashimoto |first4=Yasuyuki |last4=Yamada |year=1991 |title=Molecular cloning of hyoscyamine 6β-hydroxylase, a 2-oxoglutarate-dependent dioxygenase, from cultured roots of ''Hyoscyamus niger'' |journal=The Journal of Biological Chemistry |volume=266 |issue=15 |pages=9460–4 |pmid=2033047 |url=http://www.jbc.org/cgi/pmidlookup?view=long&pmid=2033047}}</ref><ref name="ROCHA">{{cite journal |first1=Pedro |last1=Rocha |first2=Olaf |last2=Stenzel |first3=Adrian |last3=Parr |first4=Nicholas |last4=Walton |first5=Paul |last5=Christou |first6=Birgit |last6=Dräger |first7=Mark J |last7=Leech |date=June 2002 |title=Functional expression of tropinone reductase I (''trI'') and hyoscyamine-6β-hydroxylase (''h''6''h'') from ''Hyoscyamus niger'' in ''Nicotiana tabacum'' |journal=Plant Science |volume=162 |issue=6 |pages=905–13 |doi=10.1016/S0168-9452(02)00033-X}}</ref><ref name="CARDILLO">{{cite journal |first1=Alejandra B. |last1=Cardillo |first2=Ana M. |last2=Giulietti |first3=Patricia L. |last3=Marconi |date=June 2006 |title=Analysis and sequencing of h6hmRNA, last enzyme in the tropane alkaloids pathway from anthers and hairy root cultures of ''Brugmansia candida'' (''Solanaceae'') |journal=Electronic Journal of Biotechnology |volume=9 |issue=3 |pages= |doi=10.4067/S0717-34582006000300004|doi-broken-date=2017-01-16 }}</ref>
 
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[[File:Nicotine-2D-skeletal.png|thumb|Chemical structure of nicotine.]]
 
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* [[Nicotine]]: Nicotine ([[IUPAC]] nomenclature (S)-3-(1-methylpyrrolidin-2-il) pyridine) is a [[pyrrolidine]] alkaloid produced in large quantities in the [[tobacco]] plant (''Nicotiana tabacum'').  Edible Solanaceae such as eggplants, tomatoes, potatoes, and peppers also contain nicotine, but at concentrations ~100,000–1,000,000 times less than tobacco.<ref>{{cite journal |url=http://pubs.acs.org/cgi-bin/abstract.cgi/jafcau/1999/47/i08/abs/jf990089w.html |title=Determination of the Nicotine Content of Various Edible Nightshades (Solanaceae) and Their Products and Estimation of the Associated Dietary Nicotine Intake |last1=Siegmund |first1=Barbara |last2=Leitner |first2=Erich |last3=Pfannhauser |first3=Werner |journal=J. Agric. Food Chem. |volume=47 |issue=8 |pages=3113–3120 |doi=10.1021/jf990089w |date=1999-07-23 |accessdate=2017-04-25}}</ref><ref>{{cite journal |last=Moldoveanu |first=Serban C. |last2=Scott |first2=Wayne A. |last3=Lawson |first3=Darlene M. |title=Nicotine Analysis in Several Non-Tobacco Plant Materials |url=https://www.degruyter.com/dg/viewarticle/j$002fcttr.2016.27.issue-2$002fcttr-2016-0008$002fcttr-2016-0008.xml |journal=Beiträge zur Tabakforschung International/Contributions to Tobacco Research |language=en |volume=27 |issue=2 |doi=10.1515/cttr-2016-0008 |issn=1612-9237 |date=2016-04-01 |access-date=2017-05-05}}</ref> Nicotine's function in a plant is to act as a defense against [[herbivore]]s, as it is a very effective [[neurotoxin]], in particular against [[insect]]s. In fact, nicotine has been used for many years as an [[insecticide]], though its use is currently being replaced by synthetic molecules derived from its structure. At low concentrations, nicotine acts as a stimulant in mammals, which causes the dependency in smokers. Like the tropanes, it acts on cholinergic neurons, but with the opposite effect (it is an [[agonist]] as opposed to an [[Receptor antagonist|antagonist]]). It has a higher specificity for [[nicotinic acetylcholine receptor]]s than other ACh proteins.
 
−
[[File:Kapsaicyna.svg|thumb|250px|Chemical structure of capsaicin]]
 
−
* [[Capsaicin]]: Capsaicin ([[IUPAC]] nomenclature 8-methyl-N-vanillyl-''trans''-6-nonenamide) is structurally different from nicotine and the tropanes. It is found in species of the genus ''[[Capsicum]]'', which includes [[Capsicum frutescens|chilies]] and [[Capsicum chinense|habaneros]] and it is the active ingredient that determines the [[Scoville scale|Scoville rating]] of these spices. The compound is not noticeably toxic to humans. However, it stimulates specific pain receptors in the majority of mammals, specifically those related to the perception of heat in the oral mucosa and other [[epithelium|epithelial tissues]]. When capsaicin comes into contact with these mucosae, it causes a burning sensation little different from a burn caused by fire. Capsaicin affects only mammals, not birds. Pepper seeds can survive the digestive tracts of birds; their fruit becomes brightly coloured once its seeds are mature enough to germinate, thereby attracting the attention of birds that then distribute the seeds. Capsaicin extract is used to make [[pepper spray]], a useful deterrent against aggressive mammals.
 
−
 
 
−
== Distribution ==
 
−
[[File:Solanaceae map.svg|thumb|Map showing the distribution of the Solanaceae throughout the world (light green areas)]]
 
−
 
 
−
Even though members of the Solanaceae are found on all [[continent]]s 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 [[ecosystem]]s, from [[desert]]s to [[rainforest]]s, and are often found in the secondary vegetation that colonizes disturbed areas. In general, plants in this family are of tropical and temperate distribution.
 
−
 
 
−
== Taxonomy ==
 
−
The following taxonomic synopsis of the solanaceas, including subfamilies, tribes and genera, is based on the most recent [[molecular phylogenetics]] studies of the family:<ref name=ref_duplicada_2 /><ref name="Olmstead 2007" /><ref name="Olmster">{{cite journal |first1=Richard G. |last1=Olmstead |first2=Jeffrey D. |last2=Palmer |year=1992 |title=A Chloroplast DNA Phylogeny of the Solanaceae: Subfamilial Relationships and Character Evolution |journal=Annals of the Missouri Botanical Garden |volume=79 |issue=2 |pages=346–60 |jstor=2399773 |doi=10.2307/2399773}}</ref><ref name="Martins TR">{{cite journal |first1=Talline R. |last1=Martins |first2=Todd J. |last2=Barkman |year=2005 |title=Reconstruction of Solanaceae Phylogeny Using the Nuclear Gene SAMT |journal=Systematic Botany |volume=30 |issue=2 |pages=435–47 |jstor=25064071 |doi=10.1600/0363644054223675}}</ref>
 
−
 
 
−
[[File:Cladograma de las solanáceas.jpg|thumb|Cladogram showing the relationship between the three genera of the Solanaceae family]]
 
−
 
 
−
=== Cestroideae (Browallioideae) ===
 
−
[[File:Cestrum elegans0.jpg|thumb|''Cestrum elegans'', a cestroidea used as an ornamental]] [[File:Browallia americana Flowers Closeup 1350px.jpg|thumb|''Browallia americana'']] [[File:Salpiglossis sinuata flower front view.jpg|thumb| 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 ''Cestrum'' genus 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&nbsp;µm in length), when the rest of the family generally possesses short chromosomes (for example between 1.5 and 3.52&nbsp;µm in the Nicotianoideae)
 
−
* Browallieae <small>[[Hunz.]]</small>
 
−
** ''[[Browallia]]'' <small>[[L.]]</small>, genus with six species distributed throughout the [[neotropic ecozone]] to [[Arizona]] in the [[United States]]
 
−
** ''[[Streptosolen]]'' <small>[[John Miers (botanist)|Miers]]</small>, monotypic genus native to the Andes
 
−
* Cestreae tribe <small>[[George Don|Don]]</small>,  three genera of woody plants, generally shrubs
 
−
** ''[[Cestrum]]'' <small>L.</small>, some 175 species distributed throughout the neotropic ecozone
 
−
** ''[[Sessea]]'' <small>[[Ruiz & Pav.]]</small>, 16 species from the Andes
 
−
** ''[[Vestia (plant)|Vestia]]'' <small>[[Willd.]]</small>, monotypic genus from [[Chile]]
 
−
* Salpiglossideae tribe <small>([[Benth.]]) Hunz.</small>
 
−
** ''Reyesia'' <small>Gay</small>, four species, distributed throughout Argentina and Chile
 
−
** ''[[Salpiglossis]]'' <small>Ruiz & Pav.</small>, two species originating from southern South America
 
−
 
 
−
=== 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'' <small>[[Bosser]] & D'Arcy</small> should be included in this subfamily, endemic to [[Madagascar]], and ''Metternichia'' to the southeast of [[Brazil]].
 
−
Goetzeaceae <small>[[Airy Shaw]]</small> is considered as a synonym of this subfamily.<ref name=ref_duplicada_1>{{cite journal |last1=Olmstead |first1=R.G. |last2=Bohs |first2=L. |year=2007 |title=A Summary of molecular systematic research in Solanaceae: 1982-2006 |journal=Acta Horticulturae |volume=745 |pages=255–68 |url=http://www.actahort.org/books/745/745_11.htm}}</ref>
 
−
* ''Coeloneurum'' <small>[[Radlk.]]</small>, monotypic genus endemic to [[Hispaniola]]
 
−
* ''Espadaea'' <small>[[Rchb.]]</small>, monotypic, from [[Cuba]]
 
−
* ''[[Goetzea]]'' <small>[[Wydler]]</small>, includes two species from the Antilles
 
−
* ''Henoonia'' <small>[[Griseb.]]</small>, monotypic, originating in [[Cuba]]
 
−
 
 
−
=== Petunioideae ===
 
−
 
 
−
[[File:Nierenbergia frutenscens0.jpg|thumb|''Nierenbergia frutescens'', a petunoidea.]] 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.
 
−
* ''Benthamiella'' <small>[[Speg.]]</small>,  12 species native to Patagonia
 
−
* ''[[Bouchetia (plant)|Bouchetia]]'' <small>[[Dunal]]</small>, three neotropical species 
 
−
* ''[[Brunfelsia]]'' <small>[[L.]]</small>, around 45 species from the neotropics
 
−
* ''Combera'' <small>Sandw.</small>, two species from Patagonia
 
−
* ''[[Fabiana (plant)|Fabiana]]'' <small>[[Ruiz & Pav.]]</small>, 15 species native to the Andes 
 
−
* ''Hunzikeria'' <small>D'Arcy</small>, three species from the southwest United States and [[Mexico]] 
 
−
* ''[[Latua pubiflora|Latua]]'' <small>[[Phil.]]</small>, one species from the south of Chile
 
−
* ''Leptoglossis'' <small>[[Benth.]]</small>, seven species from  western South America
 
−
* ''[[Nierembergia]]'' <small>Ruiz & Pav.</small>, 21 species from South America
 
−
* ''Pantacantha'' <small>Speg.</small>,  monospecific genus from Patagonia
 
−
* ''[[Calibrachoa]]'' <small>[[Cerv.]] ex [[La Llave]] & [[Lex.]]</small>, consists of 32 species from the neotropics. The morphological data suggest this genus should be included within the ''Petunia''. However, the molecular and cytogenetic data indicate both should be kept separate. In fact, ''Calibrachoa'' has a basic chromosome number x=9, while that of ''Petunia'' is x=7.<ref name="Ando">{{cite journal |last1=Ando |first1=Toshio |last2=Kokubun |first2=Hisashi |last3=Watanabe |first3=Hitoshi |last4=Tanaka |first4=Norio |last5=Yukawa |first5=Tomohisa |last6=Hashimoto |first6=Goro |last7=Marchesi |first7=Eduardo |last8=Suárez |first8=Enrique |last9=Basualdo |first9=Isabel L.  |year=2005 |title=Phylogenetic Analysis of ''Petunia sensu'' Jussieu (Solanaceae) using Chloroplast DNA RFLP |journal=Annals of Botany |volume=96 |issue=2 |pages=289–97 |doi=10.1093/aob/mci177 |pmid=15944177 |pmc=4246877}}</ref><ref name="Kei-Ichiro ">{{cite journal |last1=Mishiba |first1=Kei-Ichiro |last2=Ando |first2=Toshio |last3=Mii |first3=Masahiro |last4=Watanabe |first4=Hitoshi |last5=Kokubun |first5=Hisashi |last6=Hashimoto |first6=Goro |last7=Marchesi |first7=Eduardo |year=2000 |title=Nuclear DNA Content as an Index Character Discriminating Taxa in the Genus ''Petunia sensu'' Jussieu (Solanaceae) |journal=Annals of Botany |volume=85 |issue=5 |pages=665–73 |doi=10.1006/anbo.2000.1122}}</ref>
 
−
* ''[[Petunia]]'' <small> ([[Juss.]]) Wijsman</small>, 18 species from South America
 
−
* ''Plowmania'' <small>[[Hunz.]] & Subils</small>, monotypic genus from Mexico and Guatemala
 
−
 
 
−
=== Schizanthoideae ===
 
−
[[File:Schizanthus pinnatus.jpg|thumb|Zygomorphic flowers, with bilabiate corolla of ''Schizanthus pinnatus'', a schizanthoidea ornamental]]
 
−
The Schizanthoideae include annual and [[biennial plant|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 stamenodes, anther dehiscence is explosive. 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.<ref name="Olmster" /><ref name="Martins TR" /> 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.<ref name="perz 2006">{{cite journal |last1=Pérez |first1=Fernanda |last2=Arroyo |first2=Mary T. K. |last3=Medel |first3=Rodrigo |last4=Hershkovitz |first4=Mark A. |year=2006 |title=Ancestral reconstruction of flower morphology and pollination systems in ''Schizanthus'' (Solanaceae) |journal=American Journal of Botany |volume=93 |issue=7 |pages=1029–38 |pmid=21642168 |doi=10.3732/ajb.93.7.1029}}</ref>
 
−
* ''[[Schizanthus]]'' <small>Ruiz & Pav.</small>, 12 species originating from Chile.
 
−
 
 
−
=== Schwenckioideae ===
 
−
Annual plants with pericyclic fibres, their flowers are zygomorphic, the androecium has four didynamous stamens or three stamenodes; 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.
 
−
* ''Heteranthia'' <small>[[Nees]] & [[Mart.]]</small>, one species from Brazil
 
−
* ''Melananthus'' <small>[[Walp.]]</small>, five species from Brazil, Cuba, and Guatemala 
 
−
* ''Protoschwenckia'' <small>Soler </small>, monotypic genus from Bolivia and Brazil, some molecular phylogenetic studies have suggested this genus has an uncertain taxonomic position within the subfamily 
 
−
* ''Schwenckia'' <small>L.</small>, 22 species distributed throughout the neotropical regions of America
 
−
 
 
−
===Nicotianoideae===
 
−
[[File:Tabak P9290021.JPG|thumb|Tobacco inflorescence, ''Nicotiana tabacum'']]
 
−
 
 
−
* Anthocercideae <small>[[G. Don]]</small>: This tribe, endemic to Australia, contains 31 species in seven genera. Molecular phylogenetic studies of the tribe indicate it is the sister of ''[[Nicotiana]],'' and the genera ''Anthocercis, Anthotroche, Grammosolen'', and ''Symonanthus'' are [[Monophyly|monophyletic]].  Some characteristics are also thought to be derived from within the tribe, such as the unilocular stamens with semicircular opercula, bracteolate flowers, and berries as fruit.<ref name="garcia2003">{{cite journal |first1=Vicente F. |last1=Garcia |first2=Richard G. |last2=Olmstead |date=June 2003 |title=Phylogenetics of Tribe Anthocercideae (Solanaceae) Based on ''ndh''F and ''trn''L/F Sequence Data |journal=Systematic Botany |volume=28 |issue=3 |pages=609–15 |jstor=25063900 |doi=10.1043/02-52.1 |doi-broken-date=2017-01-16 }}</ref>
 
−
** ''[[Anthocercis]]'' <small>[[Labill.]]</small>, 10 species, Australia
 
−
** ''[[Anthotroche]]'' <small>[[Endl.]]</small>, four species, Australia
 
−
** ''Crenidium'' <small>[[Haegi]]</small>, monotypic genus, Australia
 
−
** ''[[Cyphanthera]]'' <small>[[John Miers (botanist)|Miers]]</small>, 9 species, Australia
 
−
** ''[[Duboisia]]'' <small>[[R.Br.]]</small>, four species, Australia
 
−
** ''Gramnosolen'' <small>Haegi</small>, two species, Australia
 
−
** ''Symonanthus'' <small>Haegi</small>, two species, Australia
 
−
* Nicotianeae tribe <small>[[Dum.]]</small>
 
−
** ''[[Nicotiana]]'' <small>L.</small>, genus widely distributed, with 52 American species, 23 Australian, and one African
 
−
 
 
−
=== Solanoideae ===
 
−
[[File:Tabasco peppers.JPG|thumb|''Capsicum frutescens'' cultivar "tabasco", a solanoidea]]
 
−
[[File:Henbane1.JPG|thumb|''Flor de beleño'' (''Hyoscyamus niger'')]] 
 
−
[[File:Nicandra physalodes fax 01.JPG|thumb|''Nicandra physalodes'' flower|alt=Nicandra physalodes flower]] 
 
−
[[File:Solandra maxima 1.jpg|thumb|''Solandra maxima'' flower]]
 
−
[[File:Physalis einzeln.jpg|thumb|In the fruit of ''Physalis peruviana'' (tomatillo),  the persistent calyx surrounds the fruit.]]  [[File:Iochroma australe 2.jpg|thumb|''Iochroma australe'' flower]]
 
−
[[File:Jaltomata procumbens flower.jpg|thumb| ''[[Jaltomata procumbens]]'' flower]] 
 
−
[[File:Buenos-Aires-Nachtschatten.jpg|thumb|''Solanum bonariense'' flower]] 
 
−
[[File:Cyphomandra betacea1.jpg|thumb|Flower of ''Solanum betaceum'' (=''Cyphomandra betacea'')]] 
 
−
[[File:Acnistus arborescens flowers.jpg|thumb|''Acnistus arborescens'' flower]]   
 
−
[[File:Scopolia carniolica2.jpg|thumb|''Scopolia carniolica'' flower]]   
 
−
 
 
−
* Capsiceae <small>Dumort</small> 
 
−
** ''[[Capsicum]]'' <small>L.</small> includes 40 accepted neotropical species<ref name=PlantList/>
 
−
** ''[[Lycianthes]]'' <small>(Dunal) Hassler</small>, some 200 species distributed throughout America and Asia
 
−
* Datureae <small>G. Don</small>, two genera are perfectly differentiated at both the morphological and molecular levels, ''Brugmansia'' includes tree species, while ''Datura'' contains herbs or shrubs, the latter genus can be divided into three sections: ''Stramonium'', ''Dutra'' and ''Ceratocaulis''.<ref name="Mace,">{{cite journal |last1=Mace |first1=E. S. |last2=Gebhardt |first2=C. G. |last3=Lester |first3=R. N. |year=1999 |title=AFLP analysis of genetic relationships in the tribe Datureae (Solanaceae) |journal=Theoretical and Applied Genetics |volume=99 |issue=3–4 |pages=634–41 |doi=10.1007/s001220051278 |pmid=22665199}}</ref>
 
−
** ''[[Brugmansia]]'' <small>Persoon</small>, six species from the Andes
 
−
** ''[[Datura]]'' <small>L.</small>, 12 neotropical species
 
−
* Hyoscyameae <small>Endl.</small> 
 
−
** ''[[Anisodus]]'' <small>Link</small>, four species from [[China]], [[India]] and the [[Himalayas]] 
 
−
**  ''[[Atropa]]'' <small>L.</small>, four Euro-Asiatic species<ref name=PlantList>{{cite web | title=The Plant List, ''Atropa'' | url=http://www.theplantlist.org/tpl1.1/search?q=atropa |publisher=Royal Botanic Garden, Kew }}</ref> 
 
−
**  ''Atropanthe'' <small>Pascher</small>, monotypic genus from China 
 
−
** ''[[Hyoscyamus]]'' <small>L.</small>, 10 accepted species<ref name=PlantList/> distributed from the Mediterranean to China 
 
−
**  ''[[Physochlaina]]'' <small> G. Don</small>, 6 accepted Euro-Asiatic species<ref name=PlantList/>
 
−
** ''Przewalskia'' <small>Maxim.</small>, one species from China
 
−
**  ''[[Scopolia]]'' <small>Jacq.</small>, disjointed distribution with one European species and another from Japan
 
−
* Jaboroseae <small>Miers</small>
 
−
** ''[[Jaborosa]]'' <small>Juss.</small>, genus that includes 23 species from South America. 
 
−
* Solandreae <small>Miers</small>
 
−
** Subtribe Juanulloinae consists 10 genera of trees and epiphytic shrubs with a neotropical distribution .<ref name="Knapp, S., V. Persson & S. Blackmore. 1997">{{cite journal |first1=Sandra |last1=Knapp |first2=Viveca |last2=Persson |first3=Stephen |last3=Blackmore |year=1997 |title=A Phylogenetic Conspectus of the Tribe Juanulloeae (Solanaceae) |journal=Annals of the Missouri Botanical Garden |volume=84 |issue=1 |pages=67–89 |jstor=2399954 |doi=10.2307/2399954}}</ref> Some of these genera (''Dyssochroma'', ''Merinthopodium'' and ''Trianaea'') show a clear dependency on various species of [[bat]]s both for pollination and dispersion of seeds.<ref name="Sazima">{{cite journal |doi=10.1093/aob/mcg190|pmid=14500325|pmc=4244854|title=Dyssochroma viridiflorum (Solanaceae): A Reproductively Bat-dependent Epiphyte from the Atlantic Rainforest in Brazil|journal=Annals of Botany|volume=92|issue=5|pages=725–30|year=2003|last1=Sazima|first1=M.|last2=Buzato|first2=S|last3=Sazima|first3=I}}</ref>
 
−
*** ''Dyssochroma'' <small>Miers</small>, two species from the south of Brazil 
 
−
*** ''Ectozoma'' <small>Miers</small>
 
−
*** ''Hawkesiophyton'' <small>Hunz.</small>
 
−
*** ''Juanulloa'' <small>Ruiz & Pav.</small>, 11 species from South and Central America
 
−
*** ''[[Markea]]'' <small>Rich.</small>, 9 species from South and Central America
 
−
*** ''Merinthopodium'' <small>J. Donn. Sm.</small> three species originating from South America
 
−
*** ''Rahowardiana'' <small>D' Arcy</small>
 
−
*** ''Schultesianthus'' <small>Hunz.</small>, eight neotropical species
 
−
*** ''[[Trianaea]]'' <small>Planch. & Linden</small>, six South American species 
 
−
** Subtribe Solandrinae, a monotypical subtribe, differs from Juanulloinae in that its embryos have incumbent cotyledons and semi-inferior ovaries.<ref name="Knapp, S., V. Persson & S. Blackmore. 1997"/> 
 
−
** ''[[Solandra]]'' <small>Sw.</small>, 10 species from the neotropical regions of America 
 
−
* Lycieae <small>Hunz.</small> has three genera of woody plants, which grow in arid or semiarid climates. The cosmopolitan genus ''Lycium'' is the oldest in the tribe and it has the greatest morphological variability.<ref name="bernadello 1987">{{cite journal |last1=Bernardello |first1=Luis M. |year=1987 |title=Comparative Floral Morphology in Lycieae (Solanaceae) |journal=Brittonia |volume=39 |issue=1 |pages=112–29 |jstor=2806983 |doi=10.2307/2806983}}</ref> Molecular phylogenetic studies suggest both ''Grabowskia'' and ''Phrodus'' should be included in the ''Lycium'',<ref name="Levin 2005">{{cite journal |last1=Levin |first1=achel A. |last2=Mille |first2=Jill S. |year=2005 |title=Relationships within tribe Lycieae (Solanaceae): Paraphyly of Lycium and multiple origins of gender dimorphism |journal=American Journal of Botany |volume=92 |issue=12 |pages=2044–53 |jstor=4125537 |doi=10.3732/ajb.92.12.2044 |pmid=21646122}}</ref> and this genus, along with ''Nolana'' and ''Sclerophylax'', form a clade (Lyciina), which currently lacks a taxonomic category.<ref name=ref_duplicada_1 /> The red fleshy berries dispersed by birds are the main type of fruit in ''Lycium''. The different types of fruit in this genus have evolved from the type of berry just mentioned to a drupe with a reduced number of seeds.<ref name="Bernardello 1998">{{cite book |last1=Bernardello |first1=L. |first2=F. |last2=Chiang-Cabrera |year=1998 |chapter=A cladistic study on the American species of ''Lycium'' (Solanaceae) based on morphological variation |title=Proceedings of the VI Congreso Latinoamericano de Botánica, Mar del Plata, Argentina, 2-8 October, 1994 |series=Monographs in Systematic Botany from the Missouri Botanical Garden |pages=33–46 |publisher=Missouri Botanical Garden Press |isbn=978-0-915279-58-6 |editor1-first=Renée H |editor1-last=Fortunato |editor2-first=Nélida M |editor2-last=Bacigalupo}}</ref> 
 
−
** ''[[Grabowskia]]'' <small>Schltdl.</small>, three species from South America 
 
−
** '' [[Lycium]]'' <small>L.</small>, 83  cosmopolitan species
 
−
** ''Phrodus'' <small>Miers</small>, two species endemic to the north of Chile 
 
−
* [[Mandragoreae]] <small>(Wettst.) Hunz. & Barboza</small> tribe does not have a defined systematic position according to molecular phylogenetic studies.<ref name=ref_duplicada_1 />
 
−
**  ''[[Mandragora (genus)|Mandragora]]'' <small>L.</small>, two species from Eurasia
 
−
* Nicandreae <small>Wettst.</small> is a tribe with two South American genera. Molecular phylogenetic studies indicate the genera are not interrelated nor are they related with other genera of the family, so their taxonomic position is uncertain.<ref name=ref_duplicada_1 />
 
−
** ''Exodeconus'' <small>Raf.</small>, six species from  western South America 
 
−
** '' [[Nicandra]]'' <small>Adans</small>, one species distributed throughout neotropical regions 
 
−
* Nolaneae<small> Rchb.</small> are mostly herbs and small shrubs with succulent leaves, they have very beautiful flowers that range from white to various shades of blue, their fruit is schizocarpal, giving rise to various nuts. 
 
−
** ''[[Nolana]]'' <small>L.</small>, 89 species distributed throughout western South America
 
−
* Physaleae <small>Miers</small>, is a large tribe that is the sister of Capsiceae. 
 
−
** Subtribe Iochrominae <small>(Miers) Hunz.</small>, a clade within the Physaleae tribe. contains 37 species, mainly distributed in the Andes, assigned to six genera. The members of this subtribe are characterized by being woody shrubs or small trees with attractive tubular or rotated flowers. They also possess great floral diversity, containing every type is present in the family. Their flowers can be red, orange, yellow, green, blue, purple, or white. The corolla can be tubular to rotated, with a variation of up to eight times in the length of the tube between the various species.<ref name="de witt">{{cite journal |last1=Smith |first1=Stacey DeWitt |last2=Baum |first2=David A. |year=2006 |title=Phylogenetics of the florally diverse Andean clade Iochrominae (Solanaceae) |journal=American Journal of Botany |volume=93 |issue=8 |pages=1140–53 |jstor=4122802 |doi=10.3732/ajb.93.8.1140 |pmid=21642180}}</ref>
 
−
*** ''[[Acnistus]]'' <small>Schott</small>, one species distributed throughout the neotropics
 
−
*** ''Dunalia'' <small>Kunth.</small>, five species from the Andes 
 
−
*** '' [[Iochroma]]'' <small>Benth.</small>, 24 species from the Andes
 
−
*** ''Saracha'' <small>Ruiz & Pav.</small>, two species from the Andes. 
 
−
*** ''Vassobia'' <small>Rusby</small>, two South American species
 
−
*** ''Eriolarynx'' <small>Hunz.</small>, three species from Argentina and Bolivia
 
−
** Physalinae <small>(Miers) Hunz. </small>, a monophyletic subtribe, contains 10 genera and includes herbs or woody shrubs with yellow, white, or purple solitary axillary flowers pollinated by bees. Once pollination occurs, the corolla falls and the calyx expands until it entirely covers the boll that is developing (the calyx is called accrescent). In many species, the calyx turns yellow or orange on maturity. The berries contain many greenish to yellow-orange seeds, often with red or purple highlights.<ref name="whiton">{{cite journal |last1=Whitson |first1=Maggie |last2=Manos |first2=Paul S. |year=2005 |title=Untangling ''Physalis'' (Solanaceae) from the Physaloids: A Two-Gene Phylogeny of the Physalinae |journal=Systematic Botany |volume=30 |issue=1 |pages=216–30 |jstor=25064051 |doi=10.1600/0363644053661841}}</ref>
 
−
*** ''Brachistus'' <small>Miers</small>, three species from Mexico and Central America 
 
−
*** ''[[Chamaesaracha]]'' <small>(A.Gray) Benth. & Hook.</small>, has 10 species from Mexico and Central America. 
 
−
*** ''Leucophysalis'' <small>Rydberg</small>, includes 3 species from the south west of the United States and Mexico.
 
−
*** ''Margaranthus'' <small>Schlecht.</small>, with 1 species from Mexico. 
 
−
*** '' [[Oryctes (plant)|Oryctes]]'' <small>S. Watson</small>, monotypic genus from the south west of the United States. 
 
−
*** ''[[Quincula]]'' <small>Raf.</small> with just 1 species from the south west of the United States and from Mexico.
 
−
*** ''[[Physalis]]'' <small>L.</small>, the largest genus of the subtribe, with 85 species distributed through the tropical regions of the Americas and with 1 species in [[China]]. 
 
−
*** ''Witheringia'' <small>L' Heritier</small>, genus with 15 species from neotropical regions. 
 
−
*** ''Tzeltalia'', genus segregated from ''Physalis'', with 2 species distributed throughout Mexico and [[Guatemala]].
 
−
*** ''Darcyanthus'', genus with just 1 species originating in Bolivia and Peru.
 
−
** Subtribe Salpichroinae, this is a subtribe of Physaleae that includes 16 American species distributed in 1 genera:
 
−
*** ''Nectouxia'' <small>Kunth.</small>, monotypic genus that is endemic to Mexico.
 
−
*** ''Salpichroa'' <small>Miers</small>, genus with 15 species from the Andes and other regions of South America. 
 
−
** Subtribe Withaninae, is a subtribe of Physaleae with a broad distribution, including 9 genera:
 
−
*** ''Archiphysalis'' <small>Kuang</small>, with 3 species from China and Japan. 
 
−
*** '' [[Athenaea (plant)|Athenaea]]'' <small>Sendtn.</small>, which includes 7 species from Brazil. 
 
−
*** ''[[Aureliana]]''<small> Sendtn.</small>, with 5 species from South America.
 
−
*** ''[[Mellissia]]'' <small>Hook. f.</small>, monotypic genus from [[Saint Helena]] with the common name Saint Helena Boxwood.
 
−
*** ''Physalisastrum'' <small>Makino</small>, with 9 Asiatic species. 
 
−
*** ''Tubocapsicum'' <small>(Wettst.) Makino</small>, with just one species endemic to China.
 
−
*** '' [[Withania]]'' <small>Pauq.</small>, with 10 species native to the [[Canary Islands]], Africa and [[Nepal]].
 
−
*** '' Cuatresia'' <small>Hunz.</small>, with 11 neotropical species. Molecular studies indicate that this genus, along with ''Deprea'' and ''Larnax'' has an uncertain taxonomic position.<ref name=ref_duplicada_1 /> 
 
−
*** ''Deprea'' <small>Raf.</small>, with 6 neotropical species.
 
−
*** '' [[Larnax (plant)|Larnax]]'' <small>Miers</small>, many taxonomists consider it to be a synonym for ''Deprea'', contains 22 species native to the Andes.
 
−
* Tribe Solaneae. The genera ''Cyphomandra'' <small>Sendtn.</small>, ''Discopodium'' <small>Hochst. </small>, ''Normania'' <small>Lowe</small>, ''Triguera'' <small>Cav.</small>  and ''Lycopersicum'' <small>Mill</small> have been transferred to ''Solanum''. The subtribe is therefore composed of two genera:<ref name=ref_duplicada_1 />   
 
−
*** ''[[Jaltomata]]'' <small>Schltdl.</small>, which contains 50 neotropical species. 
 
−
*** ''[[Solanum]]'' <small>L.</small>, the largest genus in the family and one of the broadest of the angiosperms, with 1,328 species distributed across the whole world. 
 
−
* Genera with doubtful taxonomic positions (''Incertae sedis'')
 
−
The following genera have still not been placed in any of the recognized subfamilies within the solanaceas.   
 
−
*      ''Duckeodendron'' <small>Kuhlmannb</small>, monotypic genus from the [[Amazon rainforest]].
 
−
*      ''[[Parabouchetia]]'' <small>Baillon</small>, poorly-known,monotypic genus from [[Brazil]].
 
−
*      ''Pauia'' <small>Deb. & Dutta</small>, monotypic genus from [[Assam]] and [[Arunachal Pradesh]] in N.E.[[India]]
 
−
 
 
−
=== Genera and distribution of species ===
 
−
[[File:Cestrum parqui.JPG|thumb|Flowers and foliage of ''[[Cestrum parqui]]''.]]
 
−
 
 
−
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.
 
−
 
 
−
{| class="wikitable" border="1"
 
−
! Genera
 
−
! Approximate number of species
 
−
|-
 
−
| ''[[Solanum]]''
 
−
| align="right" |1,330
 
−
|-
 
−
| ''[[Lycianthes]]''
 
−
| align="right" |200
 
−
|-
 
−
| ''[[Cestrum]]''
 
−
| align="right" |150
 
−
|-
 
−
|'' [[Nolana]]''
 
−
| align="right" |89
 
−
|-
 
−
| ''[[Physalis]]''
 
−
| align="right" |85
 
−
|-
 
−
| '' [[Lycium]]''
 
−
| align="right" |85
 
−
|-
 
−
| ''[[Nicotiana]]''
 
−
| align="right" |76
 
−
|-
 
−
| ''[[Brunfelsia]]''
 
−
| align="right" |45
 
−
|-
 
−
| Estimated number of species in the family
 
−
| align="right" |2,700
 
−
|-
 
−
|}
 
−
 
 
−
== Economic importance  ==
 
−
[[File:Brugmansia bianca rose.jpg|thumb|Pink flower of the ''Brugmansia'']]
 
−
[[File:PetuniaHybrida.jpg|thumb|''Petunia hybrida'', a herbaceous annual that is commonly used in gardens]]
 
−
 
 
−
The solanaceas include such important food species as the potato (''[[Solanum tuberosum]]''), the tomato (''[[Solanum lycopersicum]]''), the pepper (''[[Capsicum annuum]]'') and the aubergine or egg plant (''[[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,<ref name="Masis ">Masis, C.  & Madrigal, R. 1994. Lista preliminar de malezas hospedantes de Thrips
 
−
(Thysanoptera) que dañan al ''Chrysanthemum morifolium'' en el valle central de Costa Rica. Agronomía Costarricense 18(1): 99-101. 1994</ref> and certain species of ''Datura'' that play host to various types of virus that are later transmitted to cultivated solanaceas.<ref name="Ormeño ">Ormeño, J., Sepúlveda R., Rojas, R. Malezas del género ''Datura'' como factor epidemiológico del virus del mosaico de la alfalfa (amv), virus del mosaico del pepino (cmv) y virus y de la papa (pvy) en Solanáceas cultivadas. Agricultura técnica Vol. 66, Nº. 4, 2006, 333-341 [http://www.invenia.es/oai:dialnet.unirioja.es:ART0000116933 Summary in Spanish]
 
−
</ref> 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.<ref name="Pedrosa-Macedo">Pedrosa-Macedo, J.,  Olckers, T. & Vitorino, M. 2003. Phytophagous arthropods associated with ''Solanum mauritianum'' Scopoli (Solanaceae) in the first Plateau of Paraná, Brazil: a cooperative project on biological control of weeds between Brazil and South Africa. Neotrop. Entomol. 32: 519-522. [http://www.scielo.br/scielo.php?script=sci_arttext&pid=S1519-566X2003000300026&lng=en&nrm=iso Article in English, with a summary in Portuguese]
 
−
</ref>
 
−
 
 
−
Various solanaceas species are grown as ornamental trees or shrubs.<ref name="citaweb1 ">[http://www.arbolesornamentales.com/Solanaceae.htm Arboles ornamentales cultivados en España. Solanáceas]</ref> 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 solanacea 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 [[Nothotaxon|nothogenus]] called ''× Petchoa'' G. Boker & J. Shaw) that is being sold as an ornamental.<ref name="Shaw">Shaw, J. 2007. A new hybrid genus for ''Calibrachoa'' × ''Petunia'' (Solanaceae). HANBURYANA 2: 50–51 [https://www.rhs.org.uk/NR/rdonlyres/5A4EBB0E-C5F6-4204-9A0C-61CE4554E77D/0/12Hanburyana50to51.pdf]</ref><ref>[http://hsornamentals.blogspot.ca/2012/10/the-value-of-growing-petchoa-supercal.html The Value of Growing Petchoa SuperCal®. Ornamental News Oct 25 2012]</ref> Many other species, in particular those that produce alkaloids, are used in pharmacology and medicine ''([[Nicotiana]], [[Hyoscyamus]]'', and ''[[Datura]]'').
 
−
 
 
−
=== Solanaceas 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 [[chromosome]]s 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 [[mitochondria]]l genome was carried out in Argentina and the [[chloroplast]] genome was sequenced in the [[European Union]].<ref name="TSP">[http://www.sgn.cornell.edu/about/tomato_sequencing.pl International Tomato Sequencing Project Home]</ref><ref name="SOL">International Solanaceae Genomics Project (SOL), Systems Approach to Diversity and Adaptation.[http://www.sgn.cornell.edu/solanaceae-project/index.pl]</ref>
 
−
 
 
−
==See also==
 
−
* [[List of plants poisonous to equines]]
 
−
 
 
−
==References==
 
−
{{Reflist|30em}}
 
−
* {{cite book| last=D'Arcy |first=William G. | title=Solanaceae |authorlink=William Gerald D'Arcy | year=1986 | publisher=Columbia University Press | isbn=0-231-05780-6 }}
 
−
* {{cite web
 
−
|url = http://www.mobot.org/MOBOT/Research/APweb/orders/solanalesweb.htm#Solanaceae
 
−
|title = Solanaceae
 
−
|accessdate = 2007-11-04
 
−
|last = Stevens |first = P. F.
 
−
|year = 2001{{ndash}}2007
 
−
|id = [http://www.mobot.org/MOBOT/Research/APweb/welcome.html Angiosperm Phylogeny Website], version 8, June 2007.
 
−
}}
 
−
* {{cite web
 
−
|url = http://delta-intkey.com/angio/www/solanace.htm
 
−
|title = Solanaceae
 
−
|accessdate = 2007-11-04
 
−
|last1 = Watson |first1 = L.
 
−
|last2 = Dallwitz |first2=, M. J.
 
−
|id = [http://delta-intkey.com/angio/www/index.htm The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 1 June 2007.]
 
−
}}
 
−
* Dimitri, M. 1987. Enciclopedia Argentina de Agricultura y Jardinería. Tomo I. Descripción de plantas cultivadas. Editorial ACME S.A.C.I., Buenos Aires.
 
−
* {{cite web
 
−
|url = http://www.nhm.ac.uk/research-curation/projects/solanaceaesource//
 
−
|title = Solanaceae Source
 
−
|accessdate = 2007-11-17
 
−
}}
 
−
* Hunziker, Armando T. 2001. The Genera of Solanaceae. A.R.G. Gantner Verlag K.G., Ruggell, Liechtenstein. ISBN 3-904144-77-4.
 
−
 
 
−
==Further reading==
 
−
*{{cite book | last1=Hawkes |first1=J. G. |last2=Lester |first2=R. N. |last3=Skelding |first3=A. D. | title=The biology and taxonomy of the Solanaceae | year=1979 | publisher=Academic Press, London | isbn=0-12-333150-1 }}
 
−
*{{cite book | last=D'Arcy |first=William G. | title=Solanacea | year=1986 | publisher=Columbia University Press | isbn=0-231-05780-6 }}
 
−
*{{cite book | last=Radford |first=Albert E. | title=Fundamentals of Plant Systematics | year=1986 | publisher=Harper & Row, Publishers, Inc. | isbn=0-06-045305-2 }}
 
−
*{{cite journal |first1=Ken-ichi |last1=Kubo |first2=Timothy |last2=Paape |first3=Masaomi |last3=Hatakeyama |first4=Tetsuyuki |last4=Entani |first5=Akie |last5=Takara |first6=Kie |last6=Kajihara |first7=Mai |last7=Tsukahara |first8=Rie |last8=Shimizu-Inatsugi |first9=Kentaro K. |last9=Shimizu |first10=Seiji |last10=Takayama |year=2015 |title=Gene duplication and genetic exchange drive the evolution of S-RNase-based self-incompatibility in ''Petunia'' |journal=Nature Plants |volume= 1|issue= |pages= 14005|doi=10.1038/nplants.2014.5 |pmid=27246052 |laysummary=http://www.eurekalert.org/pub_releases/2015-01/uoz-ssp010815.php |laysource=EurekAlert! |laydate=January 8, 2015}}
 
−
 
 
−
==External links==
 
−
{{wikispecies|Solanaceae}}
 
−
{{commons category|Solanaceae}}
 
−
* [http://solgenomics.net Sol Genomics Network]
 
−
* [http://www.solanaceae.net/index.php Solanaceae Network - pictures of plants]
 
−
* [http://www.solanaceaesource.org/ Solanaceae Source] - A worldwide taxonomic monograph of all species in the genus Solanum.
 
−
* [http://www.chileflora.com/Florachilena/FloraEnglish/PIC_FAMILIES_SIMPLE_138.php Solanaceae of Chile, by Chileflora]
 
−
* [http://delta-intkey.com/angio/www/solanace.htm Solanaceae] in [http://delta-intkey.com/angio/ L. Watson and M.J. Dallwitz (1992 onwards). The families of flowering plants: descriptions, illustrations, identification, information retrieval.] http://delta-intkey.com
 
−
* [http://plants.usda.gov/java/ClassificationServlet?source=display&classid=Solanaceae Solanaceae] in USDA Plants Database.
 
−
* [http://flowersinisrael.com/FamSolanaceae.html Family Solanaceae] Flowers in Israel
 
−
* [http://www.sgn.cornell.edu ''SOL Genomics Network'', Universidad de Cornell]
 
−
* [http://www.solanaceae.net/index.php Imagines de various species of Solanaceae]
 
−
* [http://www.chileflora.com/Florachilena/FloraSpanish/PIC_FAMILIES_SIMPLE_138.php Solanaceae de Chile, by Chileflora]
 
−
* [http://www.nhm.ac.uk/solanaceaesource/ ''Solanaceae Source'', site with abundant information regarding Solanaceas]
 
−
* [http://www.ejournal.unam.mx/cns/no69/CNS06908.pdf Chilli: La especia del Nuevo Mundo] (Article in Spanish by Germán Octavio López Riquelme regarding the biology, nutrition, culture and medical aspects of Chile. 
 
−
* [http://solgenomics.net/community/links/related_sites.pl Solanaceae Resources on the Web]
 
−
* Jäpelt RB, Jakobsen J (2013) Vitamin D in plants: a review of occurrence, analysis, and biosynthesis. Front Plant Sci 4, No. 136 [http://www.frontiersin.org/Journal/Abstract.aspx?s=907&name=plant_physiology&ART_DOI=10.3389/fpls.2013.00136 -- Note the reference to higher cholesterol levels (and consequent Vitamin D3 levels) in family ''Solanaceae''] 
 
−
 
 
−
[[Category:Solanaceae| ]]
 
−
[[Category:Asterid families]]
 

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