Oenothera
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''Oenothera'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of about 145 species of herbaceous flowering plants native to the Americas. It is the
type genus In biological taxonomy, the type genus is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearing type of a nominal ...
of the
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
Onagraceae The Onagraceae are a family of flowering plants known as the willowherb family or evening primrose family. They include about 650 species of herbs, shrubs, and treesPrimula ''Primula'' () is a genus of herbaceous flowering plants in the family Primulaceae. They include the primrose ('' P. vulgaris''), a familiar wildflower of banks and verges. Other common species are '' P. auricula'' (auricula), '' P. veris'' (cow ...
'').


Description

The species vary in size from small alpine plants 10 centimeters tall, such as ''O. acaulis'' from
Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the east a ...
, to vigorous lowland species growing to 3 meters, such as ''O. stubbei'' from
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
. The leaves form a basal rosette at ground level and spiral up to the flowering stems. The blades are dentate or deeply lobed (
pinnatifid Pinnation (also called pennation) is the arrangement of feather-like or multi-divided features arising from both sides of a common axis. Pinnation occurs in biological morphology, in crystals, such as some forms of ice or metal crystals, and i ...
). The
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...
s of many species open in the evening, hence the name "evening primrose". They may open in under a minute. Most species have yellow flowers, but some have white, purple, pink, or red. Most native desert species are white. ''
Oenothera caespitosa ''Oenothera caespitosa'', known commonly as tufted evening primrose, desert evening primrose, rock-rose evening primrose, or fragrant evening primrose, is a perennial plant of the genus ''Oenothera'' native to much of western and central North A ...
'', a species of western North America, produces white flowers that turn pink with age. One of the most distinctive features of the flower is the stigma, which has four branches in an X shape.


Ecology

''Oenothera'' flowers are
pollinated by insects Entomophily or insect pollination is a form of pollination whereby pollen of plants, especially but not only of flowering plants, is distributed by insects. Flowers pollinated by insects typically advertise themselves with bright colours, some ...
, such as
moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...
s and bees. Like many other members of the Onagraceae, however, the pollen grains are loosely held together by viscin threads, so only insects that are morphologically specialized to gather this pollen can effectively pollinate the flowers. Bees with typical
scopa ''Scopa'' (; literally "broom") is an Italian card game, and one of the three major national card games in Italy, the others being '' Briscola'' and ''Tresette''. It is also popular in Argentina and Brazil, brought in by Italian immigrants, most ...
cannot hold it. Also, the flowers open at a time when most bee species are inactive, so the bees which visit ''Oenothera'' are generally
vespertine ''Vespertine'' is the fourth studio album by Icelandic recording artist Björk. It was released on 27 August 2001 in the United Kingdom by One Little Independent Records and in the United States by Elektra Entertainment. Production on the album ...
temporal specialists: bees that forage in the evening. The
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiospe ...
s ripen from late summer to fall. ''Oenothera'' are used as food plants by the
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
e of some Lepidoptera species, including the large white-lined sphinx. The flower moths '' Schinia felicitata'' and '' S. florida'' both feed exclusively on the genus, and the former is limited to ''O. deltoides''. In the wild, some species of evening primrose act as primary colonizers, quickly appearing in recently cleared areas. They germinate in disturbed soils, and can be found in habitat types such as dunes, roadsides, railway embankments, and waste areas. They are often casual and are eventually out competed by other species.


Origin

The genus ''Oenothera'' may have originated in
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
and
Central America Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. ...
, and spread farther north in North America and into
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the sout ...
. With the advent of international travel, species are now found in most temperate regions of the world. In Europe alone there are about 70
introduced species An introduced species, alien species, exotic species, adventive species, immigrant species, foreign species, non-indigenous species, or non-native species is a species living outside its native distributional range, but which has arrived ther ...
of ''Oenothera''. During the
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was finally confirmed in ...
era a succession of
ice age An ice age is a long period of reduction in the temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Earth's climate alternates between ice ages and gre ...
s swept down across North America, with intervening warm periods. This occurred four times, and the genus experienced four separate waves of colonization, each hybridizing with the survivors of previous waves. This formed the present-day subsection ''Euoenothera''. The group is
genetically Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar working i ...
and morphologically diverse and the species are largely interfertile, so the species boundaries have been disputed amongst taxonomists.


Genetics

The pattern of repeated colonizations resulted in a unique genetic conformation in the ''Euoenothera'' whereby the
chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
s at
meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately r ...
can form circles rather than pairs. This is the result of several reciprocal translocations between chromosomes such that the pairing occurs only at the tips. This phenomenon apparently has non-Mendelian genetic consequences; with this mode of chromosome segregation and a system of balanced-lethal genes, genetic recombination is prevented and the plants display the hybrid vigor of
heterosis Heterosis, hybrid vigor, or outbreeding enhancement is the improved or increased function of any biological quality in a hybrid offspring. An offspring is heterotic if its traits are enhanced as a result of mixing the genetic contributions o ...
. This resulted in the
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
of many sympatric races in North America east of the
Rocky Mountains The Rocky Mountains, also known as the Rockies, are a major mountain range and the largest mountain system in North America. The Rocky Mountains stretch in straight-line distance from the northernmost part of western Canada, to New Mexico ...
. Analysis of the
cytology Cell biology (also cellular biology or cytology) is a branch of biology that studies the structure, function, and behavior of cells. All living organisms are made of cells. A cell is the basic unit of life that is responsible for the living an ...
of these races and of artificial hybrids between them increased understanding of the genetic and geographic evolution of the ''Euoenothera''. This subject was a major area of genetic research during the first half of the 20th century. The appearance of sudden changes in ''
Oenothera lamarckiana ''Oenothera glazioviana'' is a species of flowering plant in the evening primrose family known by the common names large-flowered evening-primrose and redsepal evening primrose. ''Oenothera lamarckiana'' was formerly believed to be a different s ...
'' led the pioneering geneticist
Hugo de Vries Hugo Marie de Vries () (16 February 1848 – 21 May 1935) was a Dutch botanist and one of the first geneticists. He is known chiefly for suggesting the concept of genes, rediscovering the laws of heredity in the 1890s while apparently unaware o ...
to propose what he called " mutation theory" in 1901 (''Mutationstheorie'' in the German the original article was written in). This asserted that speciation was driven by sudden large mutations able to produce new varieties in a single step. The understanding that the observed changes in hybrids of the plant were caused by chromosome duplications (
polyploidy Polyploidy is a condition in which the biological cell, cells of an organism have more than one pair of (Homologous chromosome, homologous) chromosomes. Most species whose cells have Cell nucleus, nuclei (eukaryotes) are diploid, meaning they ha ...
) rather than gene mutation did not come until much later.


Taxonomy

Evening primroses were originally assigned to the genus ''Onagra'', which gave the family Onagraceae its name. ''Onagra'' ' ood of
onager The onager (; ''Equus hemionus'' ), A new species called the kiang (''E. kiang''), a Tibetan relative, was previously considered to be a subspecies of the onager as ''E. hemionus kiang'', but recent molecular studies indicate it to be a distinct ...
' was first used in
botany Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek w ...
in 1587, and in English in
Philip Miller Philip Miller FRS (1691 – 18 December 1771) was an English botanist and gardener of Scottish descent. Miller was chief gardener at the Chelsea Physic Garden for nearly 50 years from 1722, and wrote the highly popular ''The Gardeners Dicti ...
's 1754 ''Gardeners Dictionary: Abridged''. The modern name ''Oenothera'' was published by
Carolus Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, th ...
in his '' Systema Naturae''. Its
etymology Etymology ()The New Oxford Dictionary of English (1998) – p. 633 "Etymology /ˌɛtɪˈmɒlədʒi/ the study of the class in words and the way their meanings have changed throughout time". is the study of the history of the Phonological chan ...
is uncertain, but it is believed to be derived from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
words οίνος θήρα (''oinos thera'') 'wine seeker'.


Dietary uses and side effects

Certain ''Oenothera'' plants have edible parts. The
root In vascular plants, the roots are the organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often below the su ...
s of ''O. biennis'' are reportedly edible in young plants. Primrose is commonly sold as a dietary supplement in capsules containing the
seed oil Vegetable oils, or vegetable fats, are oils extracted from seeds or from other parts of fruits. Like animal fats, vegetable fats are ''mixtures'' of triglycerides. Soybean oil, grape seed oil, and cocoa butter are examples of seed oils, or fa ...
, and is advertised to treat various medical disorders, such as
eczema Dermatitis is inflammation of the skin, typically characterized by itchiness, redness and a rash. In cases of short duration, there may be small blisters, while in long-term cases the skin may become thickened. The area of skin involved can ...
, diabetic neuropathy,
rheumatoid arthritis Rheumatoid arthritis (RA) is a long-term autoimmune disorder that primarily affects joints. It typically results in warm, swollen, and painful joints. Pain and stiffness often worsen following rest. Most commonly, the wrist and hands are invol ...
, or numerous other conditions. The main phytochemical in primrose seed oil is gamma-linolenic acid. There is no high-quality scientific evidence that primrose or primrose oil has any effect on human diseases or promotion of health, and specifically no evidence that it is effective to treat eczema or
cancer Cancer is a group of diseases involving abnormal cell growth with the potential to invade or spread to other parts of the body. These contrast with benign tumors, which do not spread. Possible signs and symptoms include a lump, abnormal b ...
. Most research indicates that orally-administered evening primrose oil does not relieve symptoms of premenstrual syndrome, neither does orally administered primrose oil seem to have any effect in shortening the length of
pregnancy Pregnancy is the time during which one or more offspring develops (gestation, gestates) inside a woman, woman's uterus (womb). A multiple birth, multiple pregnancy involves more than one offspring, such as with twins. Pregnancy usually occur ...
or labour. Consuming primrose oil may cause headache or stomach upset, may increase the risk of complications during pregnancy, and may increase the risk of bleeding in people given prescription drugs as anticoagulants, such as
warfarin Warfarin, sold under the brand name Coumadin among others, is a medication that is used as an anticoagulant (blood thinner). It is commonly used to prevent blood clots such as deep vein thrombosis and pulmonary embolism, and to prevent st ...
.


Cultivation

A number of perennial members of the genus are commonly cultivated and used in landscaping in the southwestern United States. Popular species include tufted evening primrose (''
Oenothera caespitosa ''Oenothera caespitosa'', known commonly as tufted evening primrose, desert evening primrose, rock-rose evening primrose, or fragrant evening primrose, is a perennial plant of the genus ''Oenothera'' native to much of western and central North A ...
''), Mexican evening primrose ('' Oenothera berlanderii''), and Saltillo evening primrose ('' Oenothera stubbei'').Jones W. and C. Sacamano. ''Landscape Plants for Dry Regions''. Fisher Books. 2000. Annual evening primroses are also popular
ornamental plant Ornamental plants or garden plants are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars that ...
s in gardens. Many are fairly
drought A drought is defined as drier than normal conditions.Douville, H., K. Raghavan, J. Renwick, R.P. Allan, P.A. Arias, M. Barlow, R. Cerezo-Mota, A. Cherchi, T.Y. Gan, J. Gergis, D.  Jiang, A.  Khan, W.  Pokam Mba, D.  Rosenfeld, J. Tierney, an ...
-resistant. The first plants to arrive in Europe reached
Padua Padua ( ; it, Padova ; vec, Pàdova) is a city and ''comune'' in Veneto, northern Italy. Padua is on the river Bacchiglione, west of Venice. It is the capital of the province of Padua. It is also the economic and communications hub of the ...
from
Virginia Virginia, officially the Commonwealth of Virginia, is a state in the Mid-Atlantic and Southeastern regions of the United States, between the Atlantic Coast and the Appalachian Mountains. The geography and climate of the Commonwealth ar ...
in 1614 and were described by the English botanist John Goodyer in 1621. Some species are now also naturalized in parts of
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a subcontinent of Eurasia and it is located entirel ...
and
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
, and can be grown as far north as 65°N in
Finland Finland ( fi, Suomi ; sv, Finland ), officially the Republic of Finland (; ), is a Nordic country in Northern Europe. It shares land borders with Sweden to the northwest, Norway to the north, and Russia to the east, with the Gulf of B ...
. The UK National Council for the Conservation of Plants and Gardens, based at Wisley, maintains an ''Oenothera'' collection as part of its National Collections scheme.


Gallery

File:Oenothera macrocarpa a1.JPG, '' Oenothera macrocarpa'' File:Oenothera rosea.JPG, '' Oenothera rosea'' File:Oenothera caespitosa var marginata 4.jpg, ''
Oenothera caespitosa ''Oenothera caespitosa'', known commonly as tufted evening primrose, desert evening primrose, rock-rose evening primrose, or fragrant evening primrose, is a perennial plant of the genus ''Oenothera'' native to much of western and central North A ...
'' var. ''marginata'' File:Starr 051004-4684 Oenothera stricta subsp. stricta.jpg, '' Oenothera stricta'' ssp. ''stricta'' File:Primrose-brighter.jpg, ''Oenothera'' sp. opening File:Oclelandii.jpg, '' Oenothera clelandii'' File:Oenothera drummondii2.jpg, '' Oenothera drummondii'' File:Oenothera odorata 2.jpg, '' Oenothera odorata''


References


External links


Onagraceae - The Evening Primrose Family.
Smithsonian National Museum of Natural History.
Plants For A Future
* {{Authority control Onagraceae genera Night-blooming plants Medicinal plants