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''Nicotiana attenuata'' is a species of wild tobacco known by the common name coyote tobacco. It is native to western North America from
British Columbia British Columbia (commonly abbreviated as BC) is the westernmost province of Canada, situated between the Pacific Ocean and the Rocky Mountains. It has a diverse geography, with rugged landscapes that include rocky coastlines, sandy beaches, ...
to
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
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 long, the lower ones oval and the upper narrower in shape, and are borne on petioles. The
inflorescence An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...
bears several flowers with pinkish or greenish white tubular throats long, their bases enclosed in pointed
sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 The term ''sepalum'' was coined b ...
s. The flower face has five mostly white lobes. The fruit is a capsule about long.


Natural history


Introduction

''Nicotiana attenuata'' has been utilized as an ecological
model species A model organism (often shortened to model) is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workin ...
since 1994, 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 The Max Planck Institute for Chemical Ecology is located on Beutenberg Campus in Jena, Germany. It was founded in March 1996 and is one of 80 institutes of the Max Planck Society (Max Planck Gesellschaft). Chemical ecology examines the role of che ...
in
Jena Jena () is a German city and the second largest city in Thuringia. Together with the nearby cities of Erfurt and Weimar, it forms the central metropolitan area of Thuringia with approximately 500,000 inhabitants, while the city itself has a popu ...
,
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
, has been instrumental in integrating a toolbox of genomic, ecological, and analytical tools alongside field work in the
Great Basin Desert The Great Basin Desert is part of the Great Basin between the Sierra Nevada and the Wasatch Range. The desert is a geographical region that largely overlaps the Great Basin shrub steppe defined by the World Wildlife Fund, and the Central Basin a ...
to study the interactions of ''N. attenuata'' in its native environment.


Genome

''Nicotiana attenuata’s''
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ge ...
is ~2.26 Gb long, significantly more than the plant model species ''
Arabidopsis thaliana ''Arabidopsis thaliana'', the thale cress, mouse-ear cress or arabidopsis, is a small flowering plant native to Eurasia and Africa. ''A. thaliana'' is considered a weed; it is found along the shoulders of roads and in disturbed land. A winter a ...
''. Preferential gene retention after a genome-wide duplication event in the genus ''
Nicotiana ''Nicotiana'' () is a genus of herbaceous plants and shrubs in the Family (biology), family Solanaceae, that is Native plant, indigenous to the Americas, Australia, Southwestern Africa and the South Pacific. Various ''Nicotiana'' species, common ...
'' partially accounts for this large size, which is roughly twice that of '' N. obtusifolia'' (~1.23 Gb), a closely related species.


Predators

Two species of hornworm, the
tobacco hornworm ''Manduca sexta'' is a moth of the family Sphingidae present through much of the Americas. The species was first described by Carl Linnaeus in his 1763 '' Centuria Insectorum''. Commonly known as the Carolina sphinx moth and the tobacco hawk m ...
and the
tomato hornworm ''Manduca quinquemaculata'', the five-spotted hawkmoth, is a brown and gray hawk moth of the family Sphingidae. The caterpillar, often referred to as the tomato hornworm, can be a major pest in gardens; they get their name from a dark projectio ...
, 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.


Defenses against herbivory

The main predators of ''N. attenuata'' are the larvae of two hawkmoth species known as the tobacco hornworm (''
Manduca sexta ''Manduca sexta'' is a moth of the family Sphingidae present through much of the Americas. The species was Species description, first described by Carl Linnaeus in his 1763 ''Centuria Insectorum''. Commonly known as the Carolina sphinx moth and ...
'') and tomato hornworm (''
Manduca quinquemaculata ''Manduca quinquemaculata'', the five-spotted hawkmoth, is a brown and gray hawk moth of the family Sphingidae. The caterpillar, often referred to as the tomato hornworm, can be a major pest in gardens; they get their name from a dark projectio ...
''). When these worms eat trichomes on the tobacco leaves the plant produces trypsin
protease inhibitors Protease inhibitors (PIs) are medications that act by interfering with enzymes that cleave proteins. Some of the most well known are antiviral drugs widely used to treat HIV/AIDS and hepatitis C. These protease inhibitors prevent viral replicat ...
as a direct defense, weakening the hornworm's ability to digest plant material. As an indirect defense, when the leaves are eaten by larvae, the plant emits
green leaf volatiles Green leaf volatiles (GLV) are volatile organic compounds that are released when plants suffer tissue damage. Specifically, they include aldehydes, esters, and alcohols of 6-carbon compounds released after wounding. These compounds are very quickly ...
(GLVs) that attract ''
Geocoris ''Geocoris'' is a genus of insects in the family Geocoridae (although in the past the geocorids were subsumed as a subfamily under the family "Lygaeidae"). Commonly known as big-eyed bugs, the species in ''Geocoris'' are beneficial predators, but ...
'' 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 Jasmonic acid (JA) is an organic compound found in several plants including jasmine. The molecule is a member of the jasmonate class of plant hormones. It is biosynthesized from linolenic acid by the octadecanoid pathway. It was first isolat ...
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. It has also been discovered that wild tobacco can undergo defense priming in response to volatile organic compounds (VOCs) emitted from heterospecific neighbors. 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 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. 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. 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
medicinal purposes ''Medicinal Purposes'' is a Big Finish Productions audio drama based on the long-running British science fiction television series ''Doctor Who''. Plot Edinburgh, 1827. Body snatchers William Burke and William Hare are on the loose while the ...
by many Native American groups, and was smoked ceremonially by the
Hopi The Hopi are a Native American ethnic group who primarily live on the Hopi Reservation in northeastern Arizona, United States. As of the 2010 census, there are 19,338 Hopi in the country. The Hopi Tribe is a sovereign nation within the Unite ...
,
Apache The Apache () are a group of culturally related Native American tribes in the Southwestern United States, which include the Chiricahua, Jicarilla, Lipan, Mescalero, Mimbreño, Ndendahe (Bedonkohe or Mogollon and Nednhi or Carrizaleño an ...
,
Navajo The Navajo (; British English: Navaho; nv, Diné or ') are a Native American people of the Southwestern United States. With more than 399,494 enrolled tribal members , the Navajo Nation is the largest federally recognized tribe in the United ...
,
Paiute Paiute (; also Piute) refers to three non-contiguous groups of indigenous peoples of the Great Basin. Although their languages are related within the Numic group of Uto-Aztecan languages, these three groups do not form a single set. The term "Pai ...
, and other groups. Among the
Zuni people The Zuni ( zun, A:shiwi; formerly spelled ''Zuñi'') are Native American Pueblo peoples native to the Zuni River valley. The Zuni are a Federally recognized tribe and most live in the Pueblo of Zuni on the Zuni River, a tributary of the Lit ...
, the smoke is blown over the body to reduce the throbbing from rattlesnake bite. It is also smoked ceremonially among them.Stevenson, p.95


References


External links


Jepson Manual TreatmentCalflora Database: ''Nicotiana attenuata'' (coyote tobacco)Photo gallery
{{-
attenuata ''Attenuata'' is a genus of minute sea snails, marine gastropod mollusks or micromollusks in the family Lironobidae. Species There are 21 species within the genus ''Attenuata'', including: * † '' Attenuata admiranda'' Richardson, 1997 * '' A ...
Tobacco Plants used in traditional Native American medicine Flora of Northwestern Mexico Flora of the Northwestern United States Flora of the Southwestern United States Flora of British Columbia Flora of California Flora of New Mexico Flora of Texas Flora of the California desert regions Flora of the Chihuahuan Desert Flora of the Cascade Range Flora of the Great Basin Flora of the Klamath Mountains Flora of the Sierra Nevada (United States) Flora of the Sonoran Deserts Natural history of the California chaparral and woodlands Natural history of the Peninsular Ranges Natural history of the Santa Monica Mountains Natural history of the Transverse Ranges Tobacco in Mexico Flora without expected TNC conservation status