View source for Nicotiana attenuata

{{Short description|Species of flowering plant}}
{{Italic title}}
{{speciesbox
|name = Coyote tobacco
|image = Nicotiana attenuata 4.jpg
|genus = Nicotiana
|species = attenuata
|authority = [[John Torrey|Torr.]] ex [[Sereno Watson|S.Watson]]
}}

'''''Nicotiana attenuata''''' is a species of [[Nicotiana|wild tobacco]] known by the common name '''coyote tobacco'''. It is native to western North America from [[British Columbia]] to [[Texas]] and northern Mexico, where it grows in many types of habitat. It is a glandular and sparsely hairy annual herb exceeding a meter in maximum height. The leaf blades may be {{convert|10|cm|in|0}} long, the lower ones oval and the upper narrower in shape, and are borne on [[Petiole (botany)|petioles]]. The [[inflorescence]] bears several flowers with pinkish or greenish white tubular throats {{convert|2|to|3|cm|in|frac=5}} long, their bases enclosed in pointed [[sepal]]s. The flower face has five mostly white lobes. The fruit is a capsule about {{convert|1|cm|in|frac=2}} long.

== Natural history ==

=== Introduction ===
''Nicotiana attenuata'' has been utilized as an ecological [[Model organism|model species]] since 1994,<ref>{{Cite journal|last1=Baldwin|first1=I. T.|last2=Staszak-Kozinski|first2=L.|last3=Davidson|first3=R.|date=1994-09-01|title=Up in smoke: I. Smoke-derived germination cues for postfire annual, Nicotiana attenuata torr. Ex. Watson|journal=Journal of Chemical Ecology|volume=20|issue=9|pages=2345–2371|doi=10.1007/BF02033207|issn=0098-0331|pmid=24242811|s2cid=31202325}}</ref> thanks in large part to its diverse interactions with a host of different plants, insects, and microorganisms in its native habitat. Work at the [[Max Planck Institute for Chemical Ecology]] in [[Jena]], [[Germany]], has been instrumental in integrating a toolbox of genomic, ecological, and analytical tools alongside field work in the [[Great Basin Desert]] to study the interactions of ''N. attenuata'' in its native environment.

=== Genome ===
''Nicotiana attenuata’s'' [[genome]] is ~2.26 Gb long,<ref>{{Cite web|url=http://nadh.ice.mpg.de/NaDH/others/data|title=Nicotiana attenuata Data Hub - The resource for the coyote tobacco genome|website=nadh.ice.mpg.de|access-date=2017-01-12}}</ref> significantly more than the plant model species ''[[Arabidopsis thaliana]]''. Preferential gene retention after a [[Genome evolution#Gene duplication|genome-wide duplication]] event in the genus ''[[Nicotiana]]'' partially accounts for this large size, which is roughly twice that of ''[[Nicotiana obtusifolia|N. obtusifolia]]'' (~1.23 Gb), a closely related species.<ref>{{Cite journal|last1=Zhou|first1=Wenwu|last2=Brockmöller|first2=Thomas|last3=Ling|first3=Zhihao|last4=Omdahl|first4=Ashton|last5=Baldwin|first5=Ian T.|last6=Xu|first6=Shuqing|date=2016-11-04|title=Evolution of herbivore-induced early defense signaling was shaped by genome-wide duplications in Nicotiana|url=https://elifesciences.org/content/5/e19531v2|journal=eLife|language=en|volume=5|pages=e19531|doi=10.7554/eLife.19531|issn=2050-084X|pmc=5115867|pmid=27813478}}</ref>

=== Predators ===
Two species of hornworm, the [[Manduca sexta|tobacco hornworm]] and the [[Manduca quinquemaculata|tomato hornworm]], use ''N. attenuata'' as a host plant. Each of these species respond negatively to high concentrations of nicotine within plant leaves, with the tobacco hornworm showing a more intense reaction. Nicotine concentrations together with insect predators help to determine where on the plant the hornworms prefer to feed.<ref>{{Cite journal|last1=Kester|first1=Karen M.|last2=Peterson|first2=Steven C.|last3=Hanson|first3=Frank|last4=Jackson|first4=D. Michael|last5=Severson|first5=R. F.|date=2002-03-01|title=The roles of nicotine and natural enemies in determining larval feeding site distributions of Manduca sexta L. and Manduca quinquemaculata (Haworth) on tobacco|journal=Chemoecology|language=en|volume=12|issue=1|pages=1–10|doi=10.1007/s00049-002-8320-6|s2cid=29401231|issn=0937-7409}}</ref>

=== Defenses against herbivory ===
The main predators of ''N. attenuata'' are the larvae of two hawkmoth species known as the tobacco hornworm (''[[Manduca sexta]]'') and tomato hornworm (''[[Manduca quinquemaculata]]'').<ref>André Kessler, Ian T. Baldwin, Defensive Function of Herbivore-Induced Plant Volatile Emissions in Nature, Science  16 Mar 2001: Vol. 291, Issue 5511, pp. 2141-2144 DOI: 10.1126/science.291.5511.2141</ref> When these worms eat trichomes on the tobacco leaves the plant produces trypsin [[protease inhibitors]] as a direct defense, weakening the hornworm's ability to digest plant material.<ref>Jorge A. Zavala, Aparna G. Patankar, Klaus Gase, Dequan Hui, Ian T. Baldwin Plant Physiology Mar 2004, 134 (3) 1181-1190; DOI: 10.1104/pp.103.035634</ref> As an indirect defense, when the leaves are eaten by larvae, the plant emits [[green leaf volatiles]] (GLVs) that attract ''[[Geocoris]]'' bugs, which are predators of the worm. These GLVs are one of many herbivory-induced plant volatiles (HIPVs) that ''N. attenuata'' emits via [[jasmonic acid]] signaling. When GLVs come into contact with saliva from the hornworm there is a conformational change in the GLVs that attracts ''Geocoris'' bugs and increases predation on the hornworm eggs and larvae.<ref>Schuman et al. eLife 2012;1:e00007. DOI: 10.7554/eLife.00007</ref> It has also been discovered that wild tobacco can undergo defense priming in response to volatile organic compounds (VOCs) emitted from heterospecific neighbors.<ref>Danny Kessler. et al., Current Biology 20, 237–242, February 9, 2010 ª2010 Elsevier Ltd All rights reserved DOI 10.1016/j.cub.2009.11.071</ref>

Another indirect defense that has recently been studied is a change in flowering time and phenology, prompting a change in pollinator from the night-active hawkmoth to day-active hummingbirds. The flowers of ''N. attenuata'' normally open at dusk and are exposed during the night where Hawkmoth pollination occurs coupled with oviposition and thus future herbivory by hawkmoth larvae. Saliva from the hornworm causes a jasmonic acid transduction cascade leading to changes in flower phenology. Flowers reduce [[benzylacetone|benzyl acetone]] (BA) concentrations, a hawkmoth-attracting volatile, and shift corolla opening to dawn, where day-active hummingbird pollination prevails and herbivory by the Hawkmoth larvae is lessened.<ref>Danny Kessler. et al., Current Biology 20, 237–242, February 9, 2010 ª2010 Elsevier Ltd All rights reserved DOI 10.1016/j.cub.2009.11.071</ref> This shift from night opening to morning opening flowers was discovered using a native population of ''N. attenuata'' in Utah. Mesh coverings were placed over selected plants in different test groups with hornworms present or absent, and through a series of trials the ratio of morning opening to night opening flowers after just 8 days was significantly increased in the plants with hornworms present.<ref>Danny Kessler. et al., Current Biology 20, 237–242, February 9, 2010 ª2010 Elsevier Ltd All rights reserved DOI 10.1016/j.cub.2009.11.071</ref>

Collectively, these direct and indirect defenses show the impressive plasticity in behavior of ''N. attenuata'' in responding to herbivore attack.

==Uses==
This plant was used for a great variety of [[Herbalism|medicinal purposes]] by many [[Indigenous peoples of the Americas|Native American groups]], and was [[Tobacco smoking|smoked]] ceremonially by the [[Hopi]], [[Apache]], [[Navajo people|Navajo]], [[Southern Paiute|Paiute]], and other groups.<ref>[http://herb.umd.umich.edu/herb/search.pl?searchstring=Nicotiana+attenuata Ethnobotany]</ref>

Among the [[Zuni people]], the smoke is blown over the body to reduce the throbbing from rattlesnake bite.<ref>Stevenson, Matilda Coxe 1915 Ethnobotany of the Zuni Indians. SI-BAE Annual Report #30 (p. 54)</ref> It is also smoked ceremonially among them.<ref>Stevenson, p.95</ref>

==References==
{{Reflist}}

==External links==
{{Commons}}
*[http://ucjeps.berkeley.edu/cgi-bin/get_JM_treatment.pl?7625,7654,7656 Jepson Manual Treatment]
*[http://www.calflora.org/cgi-bin/species_query.cgi?where-taxon=Nicotiana+attenuata Calflora Database: ''Nicotiana attenuata'' (coyote tobacco)]
*[http://calphotos.berkeley.edu/cgi/img_query?query_src=photos_index&where-taxon=Nicotiana+attenuata Photo gallery]

{{Taxonbar|from=Q311142}}
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[[Category:Nicotiana|attenuata]]
[[Category:Tobacco]]
[[Category:Plants used in traditional Native American medicine]]
[[Category:Flora of Northwestern Mexico]]
[[Category:Flora of the Northwestern United States]]
[[Category:Flora of the Southwestern United States]]
[[Category:Flora of British Columbia]]
[[Category:Flora of California]]
[[Category:Flora of New Mexico]]
[[Category:Flora of Texas]]
[[Category:Flora of the California desert regions]]
[[Category:Flora of the Chihuahuan Desert]]
[[Category:Flora of the Cascade Range]]
[[Category:Flora of the Great Basin]]
[[Category:Flora of the Klamath Mountains]]
[[Category:Flora of the Sierra Nevada (United States)]]
[[Category:Flora of the Sonoran Deserts]]
[[Category:Natural history of the California chaparral and woodlands]]
[[Category:Natural history of the Peninsular Ranges]]
[[Category:Natural history of the Santa Monica Mountains]]
[[Category:Natural history of the Transverse Ranges]]
[[Category:Tobacco in Mexico]]
[[Category:Flora without expected TNC conservation status]]