Difference between revisions of "Nicotiana" - New World Encyclopedia

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{{Taxobox
 
{{Taxobox
 
| name = Tobacco
 
| name = Tobacco
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Numerous, see text
 
Numerous, see text
 
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{{for|the agricultural product|Tobacco}}
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'''''Nicotiana''''' is a [[genus]] of annual and perennial [[herb]]s, [[shrub]]s, and small [[tree]]s of the [[nightshade]] [[family (biology)|family]] ([[Solanaceae]]), characterized by flowers with five petals, five-lobed calyx, and alternate [[leaf|leaves]], and typically a terminal panicle-like cyme and a fruit in the form of a capsule with a large number of small seeds. The genus includes the well-known species known as tobacco plants, which are cultivated to produce [[tobacco]]. Members of ''Nicotiana'' are indigenous to [[North America|North]] and [[South America]], [[Australia]], south-west [[Africa]], and the [[Oceania|South Pacific]].
'''''Nicotiana''''' is a [[genus]] of annual and perennial [[herbaceous]] [[plant]]s and [[shrub]]s of the [[nightshade]] [[family (biology)|family]] ([[Solanaceae]]),
 
  
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Many plants contain [[nicotine]], a [[neurotoxin]] with particular specificity to [[insect]]s, and thus helpful for protecting the plant. In low concentrations, the substance acts as a stimulant in mammals and is one of the main factors responsible for the dependence-forming properties of tobacco smoking. In many [[industrialized]] countries, nicotine is among the most significant [[Drug addiction|addictive substance]]s and a cause for medical concern. However, in preindustrial societies, tobacco smoking was commonly considered a sacred or ritual activity and tightly regulated. Smoking a [[Native Americans in the United States|Native American]] "[[peace pipe]]" would invariably be preceded by paying due homage to the relevant deities and spirits and sacrificing some of the tobacco. Other cultures such as the [[Aztecs]], while smoking tobacco more casually, were nonetheless aware of the fact that it is a potent and addictive drug. Native peoples also used tobacco in other ways as an entheogen (e.g. as an additive to [[ayahuasca]]), and occasionally in [[ethnoveterinary medicine]], for example, to rid [[livestock]] of [[parasite]]s.
  
indigenous to [[North America|North]] and [[South America]], [[Australia]], south west [[Africa]] and the [[Oceania|South Pacific]]. Various ''Nicotiana'' [[species]], commonly referred to as  '''tobacco plants''', are cultivated and grown to produce [[tobacco]]. Of all ''Nicotiana'' species, [[Cultivated Tobacco]] (''N. tabacum'') is the most widely planted and is grown worldwide for production of tobacco leaf for [[cigarette]]s. The genus is named in honor of [[Jean Nicot]], who in 1561 was the first to present tobacco to the French royal court.
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Many species of ''Nicotiana'' are grown as [[ornamental plant]]s. They are popular [[vespertine (biology)|vespertine]]s, their sweet-smelling flowers opening in the evening to be visited by [[hawkmoth]]s and other [[pollinator]]s. Several tobacco plants have been used as [[model organism]]s in [[genetics]]. [[Tobacco BY-2 cells]], derived from ''N. tabacum'' [[cultivar]] 'Bright Yellow-2', are among the most important research tools in plant [[cytology]]. Tobacco has played a pioneering role in [[Callus (cell biology)|callus]] culture research and the elucidation of the mechanism by which [[kinetin]] works, laying the groundwork for modern agricultural [[biotechnology]].
  
Many plants contain [[nicotine]], a powerful [[neurotoxin]] that is particularly harmful to [[insect]]s. However, tobaccos contain a higher concentration of nicotine than most other plants. In addition, unlike many other Solanaceae they do not contain [[tropane alkaloid]]s, which are often poisonous to humans and other animals. Interesting to note that nicotine and tropane alkaloids, e.g. [[atropine]] found in [[datura stramonium]], have exactly the opposite actions in the brain. While nicotine is an [[agonist]] of [[acetylcholine receptors]], atropine acts as an [[Receptor antagonist|antagonist]] at the same receptors. Tobacco leaves and sometimes stems are commonly used as [[entheogen]]s and for pleasure. The leaves are processed into forms that can be smoked, chewed, or sniffed.
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==Overview and description==
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The genus ''Nicotiana'' is a member of the nightshade or potato family, ''Solanaceae'', along with [[potato]]es ''(Solanum tuberosum)'', [[tomato]]es ''(Solanum lycopersicum)'', [[mandrake]]s ''(Mandragora officinarum)'', and deadly nightshade ''(Atropa belladonna)''. Flowers are typically conical or funnel in shape, with five petals, which are usually fused. The leaves are alternate, often with a hairy or clammy surface. Solanaceous plants produce a fruit that is either a berry, as in the case of the tomato or wolfberry, or is a dehiscent capsule (breaks open upon drying, or dehiscing, releasing the seeds) as in the case of Datura. The seeds are usually round and flat, being two to four millimeters in diameter. The stamens are usually present in multiples of four (most commonly four or eight).  
  
In many [[industrialized]] countries, nicotine is among the most significant [[Drug addiction|addictive substance]]s and a cause for medical concern; see [[Health effects of tobacco smoking]] and [[Smokeless tobacco#Health issues]] for details. By contrast, in preindustrial societies, tobacco smoking was almost invariably considered a sacred or ritual activity and tightly regulated. Smoking a [[Native Americans in the United States|Native American]] "[[peace pipe]]" would invariably be preceded by paying due homage to the relevant deities and spirits and sacrificing some of the tobacco. Other cultures such as the [[Aztecs]], while smoking tobacco more casually, were nonetheless aware of the fact that it is a potent and addictive drug. See also [[Religious views on smoking]]. Native peoples also used tobacco in other ways as an entheogen (e.g. as an additive to [[ayahuasca]]), and occasionally in [[ethnoveterinary medicine]], e.g. to rid [[livestock]] of [[parasite]]s.
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[[Image:Nicotiana obtusifolia flower.jpg|thumb|left|240px|''[[Nicotiana obtusifolia]]'']]
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Members of ''Nicotiana'' range from desert ephermerals a few inches high, to annual [[herbaceous plant]]s, to annual and perennial [[shrub]], to perennial trees up to twenty-five feet in height (Goodspeed 1947). They exhibit a wide range in habit, inflorescence, and flower. Typically, they have an inflorescence in the form of a terminal panicle-like cyme (Hyde and Wursten 2008). Goodspeed (1947) notes however, that inflorescence types include a thyrse panicle and a flat spray as extremes, with such intermediates and specializations as mono-, di-, and pleiochasial cymes, solitary flowers, and paniculate-cymose mixtures. Nicotianas have a five-lobed calyx, which is much shorter than the corolla tube; a corolla that is white, yellow, pink, red, or purplish; and five stamens (Hyde and Wursten 2008). The fruit is a capsule with a large number of very small seeds, and the capsule is enclosed by a persistent papery calyx (Hyde and Wursten 2008).
  
==Description==
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There are about 67 species in the genus, located in America (largely in Andean South America and Mexico), Australia, South Pacific Islands, and one in Namibia (Hyde and Wursten 2008). Goodspeed (1947) had identified 58 species.
  
How many species. USDA has 22 (see tobacco article), but how many does ITIS have? The annual dictionary says 70 herbaceous species
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Various ''Nicotiana'' [[species]], commonly referred to as '''tobacco plants''', are cultivated and grown to produce [[tobacco]]. More than 20 species of tobacco plants have been identified. Tobacco leaves and sometimes stems are commonly used as [[entheogen]]s and for pleasure. The leaves are processed into forms that can be smoked, chewed, or sniffed. Of all ''Nicotiana'' species, [[cultivated tobacco]] ''(N. tabacum)'' is the most widely planted and is grown worldwide for production of tobacco leaf for [[cigarette]]s. The genus ''Nicotiana'' is named in honor of [[Jean Nicot]], who in 1561 was the first to present tobacco to the French royal court.
  
may need to correct tobacco article
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Many plants contain [[nicotine]], a powerful [[neurotoxin]] that is particularly harmful to [[insect]]s. However, tobaccos contain a higher concentration of nicotine than most other plants. In addition, unlike many other Solanaceae, they do not contain [[tropane alkaloid]]s, which are often poisonous to humans and other animals. It is interesting to note that nicotine and tropane alkaloids, such as [[atropine]] found in [[datura stramonium]], have exactly the opposite actions in the brain. While nicotine is an [[agonist]] of [[acetylcholine receptors]], atropine acts as an [[Receptor antagonist|antagonist]] at the same receptors.
  
 
== Cultivation ==
 
== Cultivation ==
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[[Image:Tobacco.jpg|thumb|left|Tobacco plantation]]
 
[[Image:Tobacco.jpg|thumb|left|Tobacco plantation]]
  
{{main|Cultivation of tobacco}}
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Tobacco plants were long grown and/or harvested by local peoples. The [[Takelma]], for example, utilized ''[[Nicotiana bigelovii|N. bigelovii]]'', and tobacco was very important to the [[Aztec]]s who considered it one of the [[Aztec entheogenic complex|sacred herbs]] of [[Xochipilli]], the "Flower Prince" (also known as ''Macuilxochitl'', "Five Flowers"), a deity of [[agriculture]] and especially [[psychoactive]] plants.
  
Tobacco plants were long grown and/or harvested by local peoples. The [[Takelma]] for example utilized ''[[Nicotiana bigelovii|N. bigelovii]]'', and tobacco was very important to the [[Aztec]]s who considered it one of the [[Aztec entheogenic complex|sacred herbs]] of [[Xochipilli]], the "Flower Prince" (also known as ''Macuilxochitl'', "Five Flowers"), a deity of [[agriculture]] and especially [[psychoactive]] plants. Indeed, the origins of [[Cultivated Tobacco]] (''N. tabacum'') are obscure; it is not known from the wild and appears to be a [[Hybrid (biology)|hybrid]] between [[Woodland Tobacco]] (''N. sylvestris''), ''[[Nicotiana tomentosiformis|N. tomentosiformis]]'' and another species (perhaps ''[[Nicotiana otophora|N. otophora]]''), deliberately selected by humans a long time ago.<ref>Ren & Timko (2001)</ref>
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The origins of [[cultivated tobacco]] ''(N. tabacum)'' are obscure; it is not known from the wild and appears to be a [[Hybrid (biology)|hybrid]] between [[woodland tobacco]] ''(N. sylvestris)'', ''[[Nicotiana tomentosiformis|N. tomentosiformis]]'' and another species (perhaps ''[[Nicotiana otophora|N. otophora]]''), deliberately selected by humans a long time ago (Ren and Timko 2001).
  
In modern tobacco farming, ''Nicotiana'' [[seed]]s are scattered onto the surface of the [[soil]], as their [[germination]] is activated by light. In colonial [[Virginia]], seedbeds were fertilized with wood ash or animal [[manure]] (frequently powdered [[horse]] manure). [[Nicotiania attenuata|''Coyote tobacco'']] of the western U.S. requires burned wood to germinate.<ref>{{cite journal|last=Baldwin|first=Ian T.|date=2001-12|title=An Ecologically Motivated Analysis of Plant-Herbivore Interactions in Native Tobacco|journal=Plant Physiol|volume=127|pages=1449-1458|url=http://www.plantphysiol.org/cgi/doi/10.1104/pp.010762|accessdate=2008-08-23}}</ref> Seedbeds were then covered with branches to protect the young plants from frost [[damage]]. These plants were left to grow until around April. Today, in the [[United States]], unlike other countries, ''Nicotiana'' is often fertilized with the mineral [[apatite]] in order to partially starve the plant for [[nitrogen]], which changes the taste of the tobacco.
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In modern tobacco farming, ''Nicotiana'' [[seed]]s are scattered onto the surface of the [[soil]], as their [[germination]] is activated by light. In colonial [[Virginia]], seedbeds were fertilized with wood ash or animal [[manure]] (frequently powdered [[horse]] manure). [[Nicotiania attenuata|''Coyote tobacco'']] of the western U.S. requires burned wood to germinate (Baldwin 2001). Seedbeds were then covered with branches to protect the young plants from frost [[damage]]. These plants were left to grow until around April. Today, in the [[United States]], unlike other countries, ''Nicotiana'' is often fertilized with the mineral [[apatite]] in order to partially starve the plant for [[nitrogen]], which changes the taste of the tobacco.
  
After the plants have reached a certain height, they are transplanted into fields. This was originally done by making a relatively large hole in the tilled earth with a tobacco peg, then placing the small plant in the hole. Various mechanical tobacco planters were invented throughout the late 19th and early 20th century to automate this process, making a hole, fertilizing it, and guiding a plant into the hole with one motion.
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After the plants have reached a certain height, they are transplanted into fields. This was originally done by making a relatively large hole in the tilled earth with a tobacco peg, then placing the small plant in the hole. Various mechanical tobacco planters were invented throughout the late nineteenth and early twentieth century to automate this process, making a hole, fertilizing it, and guiding a plant into the hole with one motion.
 
 
Many species of ''Nicotiana'' are also grown as [[ornamental plant]]s. They are popular [[vespertine (biology)|vespertine]]s, their sweet-smelling flowers opening in the evening to be visited by [[hawkmoth]]s and other [[pollinator]]s. Several tobacco plants have been used as [[model organism]]s  in [[genetics]]. [[Tobacco BY-2 cells]], derived from ''N. tabacum'' [[cultivar]] 'Bright Yellow-2', are among the most important research tools in plant [[cytology]]. Tobacco has played a pioneering role in [[Callus (cell biology)|callus]] culture research and the elucidation of the mechanism by which [[kinetin]] works, laying the groundwork for modern agricultural [[biotechnology]].
 
  
 
== Pathogens ==
 
== Pathogens ==
 
 
[[Image:TMV.jpg|thumb|right|[[Tobacco mosaic virus]] ([[electron micrograph]])]]
 
[[Image:TMV.jpg|thumb|right|[[Tobacco mosaic virus]] ([[electron micrograph]])]]
 
[[Image:Manduca sexta female sjh.JPG|thumb|right|Female ''[[Manduca sexta]]'']]
 
[[Image:Manduca sexta female sjh.JPG|thumb|right|Female ''[[Manduca sexta]]'']]
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Despite containing enough nicotine and/or other compounds such as [[germacrene]] and [[anabasine]] and other [[piperidine]] alkaloids (varying between species) to deter most [[herbivore]]s (Panter et al. 1990), a number of such animals have the ability to feed on ''Nicotiana'' species without being harmed. Nonetheless, tobacco is unpalatable to many species and therefore some tobacco plants (chiefly [[tree tobacco]], ''N. glauca'') have become established as [[invasive weed]]s in some places.
  
{{main|List of tobacco diseases}}
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In the nineteenth century, young tobacco plantings came under increasing attack from flea [[beetle]]s (''[[Epitrix cucumeris]]'' and/or ''[[Epitrix pubescens]]''), causing destruction of half the United States tobacco crop in 1876. In the years afterward, many experiments were attempted and discussed to control the flea beetle. By 1880, it was discovered that replacing the branches with a frame covered by thin fabric would effectively protect plants from the beetle. This practice spread until it became ubiquitous in the 1890s.
 
 
Despite containing enough nicotine and/or other compounds such as [[germacrene]] and [[anabasine]] and other [[piperidine]] alkaloids (varying between species) to deter most [[herbivore]]s,<ref>Panter ''et al.'' (1990)</ref> a number of such animals have [[evolve]]d the ability to feed on ''Nicotiana'' species without being harmed. Nonetheless, tobacco is unpalatable to many species and therefore some tobacco plants (chiefly [[Tree Tobacco]], ''N. glauca'') have become established as [[invasive weed]]s in some places.
 
 
 
In the nineteenth century, young tobacco plantings came under increasing attack from flea [[beetle]]s (''[[Epitrix cucumeris]]'' and/or ''[[Epitrix pubescens]]''), causing destruction of half the United States tobacco crop in 1876. In the years afterward, many experiments were attempted and discussed to control the flea beetle. By 1880 it was discovered that replacing the branches with a frame covered by thin fabric would effectively protect plants from the beetle. This practice spread until it became ubiquitous in the 1890s.
 
  
 
[[Lepidoptera]] whose [[caterpillar]]s feed on ''Nicotiana'' include:
 
[[Lepidoptera]] whose [[caterpillar]]s feed on ''Nicotiana'' include:
  
* [[Dark Sword-grass]] or Black cutworm, ''Agrotis ipsilon''
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* [[Dark sword-grass]] or Black cutworm, ''Agrotis ipsilon''
* [[Turnip Moth]], ''Agrotis segetum''
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* [[Turnip moth]], ''Agrotis segetum''
* [[Mouse Moth]], ''Amphipyra tragopoginis''
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* [[Mouse moth]], ''Amphipyra tragopoginis''
* [[The Nutmeg]], ''Discestra trifolii''
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* [[The nutmeg]], ''Discestra trifolii''
 
* ''[[Endoclita excrescens]]''
 
* ''[[Endoclita excrescens]]''
* [[Blackburn's Sphinx Moth]], ''Manduca blackburni''
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* [[Blackburn's sphinx moth]], ''Manduca blackburni''
* [[Tobacco Hornworm]], ''Manduca sexta''
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* [[Tobacco hornworm]], ''Manduca sexta''
* [[Cabbage Moth]], ''Mamestra brassicae''
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* [[Cabbage moth]], ''Mamestra brassicae''
* [[Angle Shades]], ''Phlogophora meticulosa''
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* [[Angle shades]], ''Phlogophora meticulosa''
* [[Setaceous Hebrew Character]], ''Xestia c-nigrum''
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* [[Setaceous Hebrew character]], ''Xestia c-nigrum''
  
 
These are mainly [[Noctuidae]] and some [[Sphingidae]].
 
These are mainly [[Noctuidae]] and some [[Sphingidae]].
  
 
== Selected species ==
 
== Selected species ==
 
 
[[Image:Nicotiana langsdorfii 20070810-1226-74.jpg|thumb|right|''[[Nicotiana langsdorffii]]'']]
 
[[Image:Nicotiana langsdorfii 20070810-1226-74.jpg|thumb|right|''[[Nicotiana langsdorffii]]'']]
[[Image:Nicotiana obtusifolia flower.jpg|thumb|right|''[[Nicotiana obtusifolia]]'']]
 
 
[[Image:Nicotinana alata0.jpg|thumb|right|''[[Nicotiana × sanderae]]'' ornamental cultivar]]
 
[[Image:Nicotinana alata0.jpg|thumb|right|''[[Nicotiana × sanderae]]'' ornamental cultivar]]
  
* ''[[Nicotiana acuminata]]'' &ndash; Manyflower tobacco<ref name="itis1999">ITIS (1999)</ref>
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* ''[[Nicotiana acuminata]]'' &ndash; Manyflower tobacco (ITIS 1999)
* ''[[Nicotiana africana]]''{{Verify source|date=August 2008}}
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* ''[[Nicotiana africana]]''
* ''[[Nicotiana alata]]'' &ndash; Winged tobacco, Jasmine tobacco, ''tanbaku'' ([[Persian language|Persian]])<ref name="itis1999" />
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* ''[[Nicotiana alata]]'' &ndash; Winged tobacco, Jasmine tobacco, ''tanbaku'' ([[Persian language|Persian]]) (ITIS 1999)
* ''[[Nicotiana attenuata]]'' &ndash; Coyote tobacco<ref name="itis1999" />
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* ''[[Nicotiana attenuata]]'' &ndash; Coyote tobacco (ITIS 1999)
* ''[[Nicotiana benthamiana]]''{{Verify source|date=August 2008}}
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* ''[[Nicotiana benthamiana]]''
* ''[[Nicotiana bigelovii]]''{{Verify source|date=August 2008}}
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* ''[[Nicotiana bigelovii]]''
* ''[[Nicotiana clevelandii]]'' &ndash; Cleveland's tobacco<ref name="itis1999" />
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* ''[[Nicotiana clevelandii]]'' &ndash; Cleveland's tobacco (ITIS 1999)
* ''[[Nicotiana debneyi]]''{{Verify source|date=November 2007}}
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* ''[[Nicotiana debneyi]]''
* ''[[Nicotiana × digluta]]''{{Verify source|date=August 2008}}
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* ''[[Nicotiana × digluta]]''
* ''[[Nicotiana excelsior]]'' &ndash; tobacco<ref name="itis1999" />
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* ''[[Nicotiana excelsior]]'' &ndash; tobacco (ITIS 1999)
* ''[[Nicotiana exigua]]''{{Verify source|date=November 2007}}
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* ''[[Nicotiana exigua]]''  
* ''[[Nicotiana forgetiana]]'' &ndash; tobacco<ref name="itis1999" />
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* ''[[Nicotiana forgetiana]]'' &ndash; tobacco (ITIS 1999)
* ''[[Nicotiana glauca]]'' &ndash; Tree Tobacco, Brazilian Tree Tobacco, Shrub Tobacco, Mustard Tree<ref name="itis1999" />
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* ''[[Nicotiana glauca]]'' &ndash; Tree tobacco, Brazilian tree tobacco, shrub tobacco, mustard tree (ITIS 1999)
* ''[[Nicotiana glutinosa]]'' &ndash; tobacco<ref name="itis1999" />
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* ''[[Nicotiana glutinosa]]'' &ndash; tobacco (ITIS 1999)
* ''[[Nicotiana kawakamii]]''{{Verify source|date=November 2007}}
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* ''[[Nicotiana kawakamii]]''
* ''[[Nicotiana knightiana]]''{{Verify source|date=November 2007}}
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* ''[[Nicotiana knightiana]]''
* ''[[Nicotiana langsdorffii]]'' &ndash; Langsdorff's tobacco<ref name="itis1999" />
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* ''[[Nicotiana langsdorffii]]'' &ndash; Langsdorff's tobacco (ITIS 1999)
* ''[[Nicotiana longiflora]]'' &ndash; Longflower Tobacco<ref name="itis1999" />
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* ''[[Nicotiana longiflora]]'' &ndash; Longflower tobacco (ITIS 1999)
* ''[[Nicotiana obtusifolia]]'' (''N. trigonophylla'') &ndash; Desert Tobacco, ''punche'', ''"[[tabaquillo]]"''<ref name="itis1999" />
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* ''[[Nicotiana obtusifolia]]'' ''(N. trigonophylla)'' &ndash; Desert Tobacco, ''punche'', ''"[[tabaquillo]]"'' (ITIS 1999)
* ''[[Nicotiana otophora]]''{{Verify source|date=August 2008}}
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* ''[[Nicotiana otophora]]''
* ''[[Nicotiana paniculata]]'' &ndash; tobacco<ref name="itis1999" />
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* ''[[Nicotiana paniculata]]'' &ndash; tobacco (ITIS 1999)
* ''[[Nicotiana persica]]''{{Verify source|date=August 2008}}
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* ''[[Nicotiana persica]]''
* ''[[Nicotiana plumbagifolia]]'' &ndash; Tex-Mex tobacco<ref name="itis1999" />
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* ''[[Nicotiana plumbagifolia]]'' &ndash; Tex-Mex tobacco (ITIS 1999)
* ''[[Nicotiana quadrivalvis]]'' &ndash; Indian tobacco<ref name="itis1999" />
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* ''[[Nicotiana quadrivalvis]]'' &ndash; Indian tobacco (ITIS 1999)
* ''[[Nicotiana repanda]]'' &ndash; Fiddleleaf tobacco, Wild tobacco<ref name="itis1999" />
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* ''[[Nicotiana repanda]]'' &ndash; Fiddleleaf tobacco, Wild tobacco (ITIS 1999)
* ''[[Nicotiana rustica]]'' &ndash; Aztec tobacco, Mapacho<ref name="itis1999" />
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* ''[[Nicotiana rustica]]'' &ndash; Aztec tobacco, Mapacho (ITIS 1999)
* ''[[Nicotiana × sanderae]]'' &ndash; Sander's tobacco<ref name="itis1999" />
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* ''[[Nicotiana × sanderae]]'' &ndash; Sander's tobacco (ITIS 1999)
* ''[[Nicotiana stocktonii]]''{{Verify source|date=August 2008}}
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* ''[[Nicotiana stocktonii]]''
* ''[[Nicotiana suaveolens]]'' &ndash; Australian tobacco<ref name="itis1999" />
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* ''[[Nicotiana suaveolens]]'' &ndash; Australian tobacco (ITIS 1999)
* ''[[Nicotiana sylvestris]]'' &ndash; South American tobacco, Woodland Tobacco<ref name="itis1999" />
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* ''[[Nicotiana sylvestris]]'' &ndash; South American tobacco, Woodland Tobacco (ITIS 1999)
* ''[[Nicotiana tabacum]]'' &ndash; Cultivated Tobacco, Common Tobacco (a cultivated hybrid - properly ''Nicotiana × tabacum'')<ref name="itis1999" />
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* ''[[Nicotiana tabacum]]'' &ndash; Cultivated tobacco, common tobacco (a cultivated hybrid - properly ''Nicotiana × tabacum'') (ITIS 1999)
* ''[[Nicotiana tomentosa]]'' &ndash; tobacco<ref name="itis1999" />
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* ''[[Nicotiana tomentosa]]'' &ndash; tobacco (ITIS 1999)
* ''[[Nicotiana tomentosiformis]]''{{Verify source|date=August 2008}}
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* ''[[Nicotiana tomentosiformis]]''
 
 
== See also ==
 
 
 
{{commons}}
 
{{wikispecies}}
 
 
 
* [[Agroinfiltration]]
 
* [[Edgar Anderson]]
 
* [[Jamestown, Virginia]]
 
* [[Jean Nicot]]
 
* [[John Rolfe]]
 
* [[List of plants poisonous to equines]]
 
* [[Nancy Tyson Burbidge]]
 
* [[Pectinesterase]]
 
* [[Turkish tobacco]]
 
* [[Indian tobacco]]
 
  
 
== References ==
 
== References ==
  
=== Notes ===
+
* Baldwin, I. T. 2001. [http://www.plantphysiol.org/cgi/doi/10.1104/pp.010762 An ecologically motivated analysis of plant-herbivore interactions in native tobacco] ''Plant Physiol'' 127: 1449-1458. Retrieved September 18, 2008.
{{reflist|colwidth=40em}}
+
* Goodspeed, T. H. 1947. [http://www.pnas.org/content/33/6/158.full.pdf+html On the evolution of the genus ''Nicotiana''] ''PNAS'' 33(6): 158-171.
 
+
* Hyde, M. A., and B. Wursten. 2008. [http://www.zimbabweflora.co.zw/speciesdata/genus.php?genus_id=1254 ''Nicotiana''] ''Flora of Zimbabwe''. Retrieved September 18, 2008.
=== Bibliography ===
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* Integrated Taxonomic Information System (ITIS). 1999. [http://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=30562 ''Nicotiana''] ''ITIS Taxonomic Serial No.: 30562''. Retrieved September 18, 2008.
 
+
* Panter, K. E., R.F. Keeler, T. D. Bunch, and R. J. Callan. 1990. [http://www.ncbi.nlm.nih.gov/pubmed/2089736 Congenital skeletal malformations and cleft palate induced in goats by ingestion of ''Lupinus, Conium and Nicotiana'' species] ''Toxicon'' 28(12): 1377-1385. PMID 2089736. Retrieved September 18, 2008.
{{refbegin}}
+
* Ren, N., and M. P. Timko. 2001. [http://article.pubs.nrc-cnrc.gc.ca/ppv/RPViewDoc?issn=0831-2796&volume=44&issue=4&startPage=559 AFLP analysis of genetic polymorphism and evolutionary relationships among cultivated and wild ''Nicotiana'' species] ''Genome'' 44(4): 559-571. Retrieved September 18, 2008.
* {{aut|[[Integrated Taxonomic Information System]] (ITIS)}} (1999): [http://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=30562 ''Nicotiana'']. Retrieved 2007-NOV-20.
+
* United States Department of Agriculture (USDA). 2006. [http://plants.usda.gov/java/ClassificationServlet?source=display&classid=NICOT Classification ''Nicotiana]'' ''USDA''. Retrieved September 18, 2008.
 
 
* {{aut|Panter, K.E.; Keeler, R.F.; Bunch, T.D. & Callan, R.J.}} (1990): Congenital skeletal malformations and cleft palate induced in goats by ingestion of ''Lupinus, Conium and Nicotiana'' species. ''Toxicon'' '''28'''(12): 1377-1385. <small>PMID 2089736</small> (HTML abstract)
 
 
 
* {{aut|Ren, Nan & Timko, Michael P.}} (2001): AFLP analysis of genetic polymorphism and evolutionary relationships among cultivated and wild ''Nicotiana'' species. ''[[Genome]]'' '''44'''(4): 559-571. <small>{{doi|10.1139/gen-44-4-559}}</small> [http://article.pubs.nrc-cnrc.gc.ca/ppv/RPViewDoc?issn=0831-2796&volume=44&issue=4&startPage=559 PDF fulltext]
 
{{refend}}
 
 
 
* United States Department of Agriculture (USDA). 2006. [http://plants.usda.gov/java/ClassificationServlet?source=display&classid=NICOT ''Nicotiana Plants Classification Report]''
 
 
 
 
 
http://www.zimbabweflora.co.zw/speciesdata/genus.php?genus_id=1254 Nicotiana
 
Mark Hyde and Bart Wursten, 2002-8
 
Hyde, M.A. & Wursten, B. (2008). Flora of Zimbabwe: Genus page: Nicotiana.
 
http://www.zimbabweflora.co.zw/speciesdata/genus.php?genus_id=1254, retrieved 17 September 2008
 
 
 
http://www.pnas.org/content/33/6/158.full.pdf+html
 
PNAS  1947-06  vol. 33  no. 6  158-171
 
Botany:                      T. H. Goodspeed
 
On the Evolution of the Genus NicotianaPNAS 1947 33:158-171
 
 
 
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Latest revision as of 18:56, 22 September 2008

Tobacco
Flowering Nicotiana tabacum
Flowering Nicotiana tabacum
Scientific classification
Kingdom: Plantae
Division: Magnoliophyta
Class: Magnoliopsida
Subclass: Asteridae
(unranked) Euasterids I
Order: Solanales
Family: Solanaceae
Genus: Nicotiana
L.
Species

Numerous, see text

Nicotiana is a genus of annual and perennial herbs, shrubs, and small trees of the nightshade family (Solanaceae), characterized by flowers with five petals, five-lobed calyx, and alternate leaves, and typically a terminal panicle-like cyme and a fruit in the form of a capsule with a large number of small seeds. The genus includes the well-known species known as tobacco plants, which are cultivated to produce tobacco. Members of Nicotiana are indigenous to North and South America, Australia, south-west Africa, and the South Pacific.

Many plants contain nicotine, a neurotoxin with particular specificity to insects, and thus helpful for protecting the plant. In low concentrations, the substance acts as a stimulant in mammals and is one of the main factors responsible for the dependence-forming properties of tobacco smoking. In many industrialized countries, nicotine is among the most significant addictive substances and a cause for medical concern. However, in preindustrial societies, tobacco smoking was commonly considered a sacred or ritual activity and tightly regulated. Smoking a Native American "peace pipe" would invariably be preceded by paying due homage to the relevant deities and spirits and sacrificing some of the tobacco. Other cultures such as the Aztecs, while smoking tobacco more casually, were nonetheless aware of the fact that it is a potent and addictive drug. Native peoples also used tobacco in other ways as an entheogen (e.g. as an additive to ayahuasca), and occasionally in ethnoveterinary medicine, for example, to rid livestock of parasites.

Many species of Nicotiana are grown as ornamental plants. They are popular vespertines, their sweet-smelling flowers opening in the evening to be visited by hawkmoths and other pollinators. Several tobacco plants have been used as model organisms in genetics. Tobacco BY-2 cells, derived from N. tabacum cultivar 'Bright Yellow-2', are among the most important research tools in plant cytology. Tobacco has played a pioneering role in callus culture research and the elucidation of the mechanism by which kinetin works, laying the groundwork for modern agricultural biotechnology.

Overview and description

The genus Nicotiana is a member of the nightshade or potato family, Solanaceae, along with potatoes (Solanum tuberosum), tomatoes (Solanum lycopersicum), mandrakes (Mandragora officinarum), and deadly nightshade (Atropa belladonna). Flowers are typically conical or funnel in shape, with five petals, which are usually fused. The leaves are alternate, often with a hairy or clammy surface. Solanaceous plants produce a fruit that is either a berry, as in the case of the tomato or wolfberry, or is a dehiscent capsule (breaks open upon drying, or dehiscing, releasing the seeds) as in the case of Datura. The seeds are usually round and flat, being two to four millimeters in diameter. The stamens are usually present in multiples of four (most commonly four or eight).

Nicotiana obtusifolia

Members of Nicotiana range from desert ephermerals a few inches high, to annual herbaceous plants, to annual and perennial shrub, to perennial trees up to twenty-five feet in height (Goodspeed 1947). They exhibit a wide range in habit, inflorescence, and flower. Typically, they have an inflorescence in the form of a terminal panicle-like cyme (Hyde and Wursten 2008). Goodspeed (1947) notes however, that inflorescence types include a thyrse panicle and a flat spray as extremes, with such intermediates and specializations as mono-, di-, and pleiochasial cymes, solitary flowers, and paniculate-cymose mixtures. Nicotianas have a five-lobed calyx, which is much shorter than the corolla tube; a corolla that is white, yellow, pink, red, or purplish; and five stamens (Hyde and Wursten 2008). The fruit is a capsule with a large number of very small seeds, and the capsule is enclosed by a persistent papery calyx (Hyde and Wursten 2008).

There are about 67 species in the genus, located in America (largely in Andean South America and Mexico), Australia, South Pacific Islands, and one in Namibia (Hyde and Wursten 2008). Goodspeed (1947) had identified 58 species.

Various Nicotiana species, commonly referred to as tobacco plants, are cultivated and grown to produce tobacco. More than 20 species of tobacco plants have been identified. Tobacco leaves and sometimes stems are commonly used as entheogens and for pleasure. The leaves are processed into forms that can be smoked, chewed, or sniffed. Of all Nicotiana species, cultivated tobacco (N. tabacum) is the most widely planted and is grown worldwide for production of tobacco leaf for cigarettes. The genus Nicotiana is named in honor of Jean Nicot, who in 1561 was the first to present tobacco to the French royal court.

Many plants contain nicotine, a powerful neurotoxin that is particularly harmful to insects. However, tobaccos contain a higher concentration of nicotine than most other plants. In addition, unlike many other Solanaceae, they do not contain tropane alkaloids, which are often poisonous to humans and other animals. It is interesting to note that nicotine and tropane alkaloids, such as atropine found in datura stramonium, have exactly the opposite actions in the brain. While nicotine is an agonist of acetylcholine receptors, atropine acts as an antagonist at the same receptors.

Cultivation

Tobacco plantation

Tobacco plants were long grown and/or harvested by local peoples. The Takelma, for example, utilized N. bigelovii, and tobacco was very important to the Aztecs who considered it one of the sacred herbs of Xochipilli, the "Flower Prince" (also known as Macuilxochitl, "Five Flowers"), a deity of agriculture and especially psychoactive plants.

The origins of cultivated tobacco (N. tabacum) are obscure; it is not known from the wild and appears to be a hybrid between woodland tobacco (N. sylvestris), N. tomentosiformis and another species (perhaps N. otophora), deliberately selected by humans a long time ago (Ren and Timko 2001).

In modern tobacco farming, Nicotiana seeds are scattered onto the surface of the soil, as their germination is activated by light. In colonial Virginia, seedbeds were fertilized with wood ash or animal manure (frequently powdered horse manure). Coyote tobacco of the western U.S. requires burned wood to germinate (Baldwin 2001). Seedbeds were then covered with branches to protect the young plants from frost damage. These plants were left to grow until around April. Today, in the United States, unlike other countries, Nicotiana is often fertilized with the mineral apatite in order to partially starve the plant for nitrogen, which changes the taste of the tobacco.

After the plants have reached a certain height, they are transplanted into fields. This was originally done by making a relatively large hole in the tilled earth with a tobacco peg, then placing the small plant in the hole. Various mechanical tobacco planters were invented throughout the late nineteenth and early twentieth century to automate this process, making a hole, fertilizing it, and guiding a plant into the hole with one motion.

Pathogens

Tobacco mosaic virus (electron micrograph)
Female Manduca sexta

Despite containing enough nicotine and/or other compounds such as germacrene and anabasine and other piperidine alkaloids (varying between species) to deter most herbivores (Panter et al. 1990), a number of such animals have the ability to feed on Nicotiana species without being harmed. Nonetheless, tobacco is unpalatable to many species and therefore some tobacco plants (chiefly tree tobacco, N. glauca) have become established as invasive weeds in some places.

In the nineteenth century, young tobacco plantings came under increasing attack from flea beetles (Epitrix cucumeris and/or Epitrix pubescens), causing destruction of half the United States tobacco crop in 1876. In the years afterward, many experiments were attempted and discussed to control the flea beetle. By 1880, it was discovered that replacing the branches with a frame covered by thin fabric would effectively protect plants from the beetle. This practice spread until it became ubiquitous in the 1890s.

Lepidoptera whose caterpillars feed on Nicotiana include:

  • Dark sword-grass or Black cutworm, Agrotis ipsilon
  • Turnip moth, Agrotis segetum
  • Mouse moth, Amphipyra tragopoginis
  • The nutmeg, Discestra trifolii
  • Endoclita excrescens
  • Blackburn's sphinx moth, Manduca blackburni
  • Tobacco hornworm, Manduca sexta
  • Cabbage moth, Mamestra brassicae
  • Angle shades, Phlogophora meticulosa
  • Setaceous Hebrew character, Xestia c-nigrum

These are mainly Noctuidae and some Sphingidae.

Selected species

Nicotiana langsdorffii
Nicotiana × sanderae ornamental cultivar
  • Nicotiana acuminata – Manyflower tobacco (ITIS 1999)
  • Nicotiana africana
  • Nicotiana alata – Winged tobacco, Jasmine tobacco, tanbaku (Persian) (ITIS 1999)
  • Nicotiana attenuata – Coyote tobacco (ITIS 1999)
  • Nicotiana benthamiana
  • Nicotiana bigelovii
  • Nicotiana clevelandii – Cleveland's tobacco (ITIS 1999)
  • Nicotiana debneyi
  • Nicotiana × digluta
  • Nicotiana excelsior – tobacco (ITIS 1999)
  • Nicotiana exigua
  • Nicotiana forgetiana – tobacco (ITIS 1999)
  • Nicotiana glauca – Tree tobacco, Brazilian tree tobacco, shrub tobacco, mustard tree (ITIS 1999)
  • Nicotiana glutinosa – tobacco (ITIS 1999)
  • Nicotiana kawakamii
  • Nicotiana knightiana
  • Nicotiana langsdorffii – Langsdorff's tobacco (ITIS 1999)
  • Nicotiana longiflora – Longflower tobacco (ITIS 1999)
  • Nicotiana obtusifolia (N. trigonophylla) – Desert Tobacco, punche, "tabaquillo" (ITIS 1999)
  • Nicotiana otophora
  • Nicotiana paniculata – tobacco (ITIS 1999)
  • Nicotiana persica
  • Nicotiana plumbagifolia – Tex-Mex tobacco (ITIS 1999)
  • Nicotiana quadrivalvis – Indian tobacco (ITIS 1999)
  • Nicotiana repanda – Fiddleleaf tobacco, Wild tobacco (ITIS 1999)
  • Nicotiana rustica – Aztec tobacco, Mapacho (ITIS 1999)
  • Nicotiana × sanderae – Sander's tobacco (ITIS 1999)
  • Nicotiana stocktonii
  • Nicotiana suaveolens – Australian tobacco (ITIS 1999)
  • Nicotiana sylvestris – South American tobacco, Woodland Tobacco (ITIS 1999)
  • Nicotiana tabacum – Cultivated tobacco, common tobacco (a cultivated hybrid - properly Nicotiana × tabacum) (ITIS 1999)
  • Nicotiana tomentosa – tobacco (ITIS 1999)
  • Nicotiana tomentosiformis

References
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