English Common Names
(Yellow) evening primrose. Archaic names: tree primrose, scurvish, scabbish (scabish), king’s cure-all, nightwillow herb, sundrops, tree primrose, fever plant.
The name evening-primrose is optionally hyphen-ated. Evening primrose isn’t a “primrose,” a name best applied to the genus Primula. The “evening” in the name relates to the fact that the flowers of many of the 125 species of Oenothera open in the evening and release a scent that attracts moths for pollination.
French Common Names Onagre bisannuelle (commune).
Morphology
Oenothera biennis is a biennial (as the name sug-gests) or short-lived perennial herb producing strong fleshy roots and a basal rosette of lanceolate leaves in the first year. In the second year the stem grows to 1–2 m tall and develops a spicate inflorescence of 4-parted, yellow, tubular flowers. The fruit is a cap-sule containing many seeds which mature in the fall.
The seeds are very small (ca. 0.5 g/1000), but a single plant can easily produce 150,000. The pol-len of many, if not all species of Oenothera, is un-usual in having protruding apertures and viscin threads.
Classification and Geography
Evening primrose, a native of North America, is found in all provinces of Canada, but is more frequent in the east than the west. The species ex-tends south to Florida and Mexico.
Many texts recognize var. canescens T. & G., with dense grayish pubescence, as the predominant plant of western North America, while the eastern plants are referable to var. biennis. The classifica-tion of the transcontinental O. biennis and related
species of both North America and Eurasia is, however, very complex. Cytogenetic races of eve-ning primrose are sometimes segregated as dis-tinct species, although these are usually difficult to distinguish morphologically.
Oenothera biennis is a complete translocation complex-heterozygote, with two sets (the “com-plexes”) of seven chromosomes maintained by a system of balanced lethal genes. This type of in-heritance is known in a few other genera, but was first described in O. biennis, and is the classical ex-ample of the phenomenon discussed in evolution and genetics courses. At meiosis, translocations1 link the chromosomes into a ring of 14, but zig-zag (alternate) separation of the chromosomes gener-ates the original parental sets. Lethal factors kill the pollen carrying one of the sets (so that there is 50%
pollen fertility), and ovule lethal factors limit sur-vival to the set of chromosomes complementary to that in the pollen. Self-pollination generates off-spring with the two chromosome complements found originally in the maternal plant. The perma-nent hybrid vigor resulting from the combination of two quite different genomes is thought to ex-plain the success of evening primrose as a colo-nizing species.
While it is clear that O. biennis is the chief Oenothera species that has been grown as a medici-nal oilseed, related species have also been cultivated, often unknowingly. Other species from which cultivars have been derived include O. glazioviana Micheli (“O. lamarckiana” of many authors) and O. parviflora Micheli.
Ecology
Evening primrose is a frequent weed of road-sides, waste places, and abandoned land, often found in light sandy and gravelly soils. It commonly oc-curs in association with early successional, biennial and perennial weeds.
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Oenot hera biennis
1Translocations are chromosomal abnormalities which occur when chromosomes break and the fragments rejoin to other chro-mosomes. In “reciprocal translocation” two non-homologous chromosomes break and exchange fragments, and during meiosis chromosome pairing and segregation are abnormal, the chromosomes involved often linking together, leading to odd segrega-tion patterns and reducsegrega-tion of viability. Translocasegrega-tion heterozygotes are the product of reciprocal translocasegrega-tions that link all of the chromosomes together. Evolution of new species by chromosome translocations is very common in plants, less so in ani-mals. In humans, translocations can cause severe diseases; for example, Familial Downs Syndrome, which occurs in 1 of 600 births, may result from translocation of part of chromosome 21 to another chromosome [it may also occur as trisomy (three chro-mosomes rather than the normal two) of chromosome 21]. One study (http://www.st-elizabeth.edu/~ikessler/hgen/genlec5.html) found that of live human births, about 6% suffered from some kind of chromosomal abnormality; about 2% of live human births have extra or missing sex chromosomes, about 1.4% have a third non-sex chromosome (i.e. trisomy, mentioned above), and about 2% have translocations.
Medicinal Uses
Evening primrose extracts were used medici-nally by both Indians and early settlers. In Europe during the early 1600s it was called “King’s Cure-all.” An infusion of the whole plant was thought to counter asthmatic cough, gastro-intestinal dis-orders, and whooping cough, and to reduce pain.
Poultices were used to treat bruises and wounds Evening primrose has attracted great interest for its seed oil, used medicinally as a nutritional supplement. The health value of the seed oil re-sides in an unusual polyunsaturated fatty acid, γ-linolenic acid (gamma-γ-linolenic acid) or simply GLA. The seeds contain 17–25% oil, of which only 7–10% is GLA, although climate and maturity affect oil content and qualitative composition, as well as overall yield. GLA is one of the so-called essential fatty acids needed by humans for maintenance of cell functions. It is a precursor in the biosynthesis of prostaglandins, especially prostaglandin E1, a hor-mone-like substance that has been clinically shown to regulate metabolic functions in mammals; it af-fects cholesterol levels, dilates blood vessels, re-duces inflammation, and has additional effects.
GLA is thought to be important for development of brain tissue and other tissue growth, and nature seems to provide for human infants with high lev-els of GLA in human milk. GLA is a normal con-version product of linoleic acid, a major constituent of most vegetable oils, so that it would appear that humans should not experience a shortage. Never-theless, some people, perhaps 10–20% of the popu-lation, evidently do not have adequate levels, even when receiving large amount of linoleic acid. The
deficiency seems due to lack of an enzyme that metabolizes GLA from linolenic acid, so that there is a deficiency of GLA in the blood. Useful for treating atopic eczema, GLA has therapeutic prom-ise for premenstrual syndrome, diabetes, multiple sclerosis, alcoholism, inflammation, heart disease and stroke. Rubbing GLA into the skin is thought to be an alternative route of assimilation, and so cosmetic preparations sometimes incorporate GLA.
Pharmaceutical and food companies are developing GLA-containing supplements and specialty foods for infants, the elderly, and people with health problems.
Toxicity
Side effects of consumption of GLA-fortified foods and supplements have been documented, so that use should be guided by doctors and pharma-cists.
Chemistry
Gamma-linolenic acid, the constituent of chief medicinal interest, is discussed above.
Non-medicinal Uses
There are ornamental forms of Oenothera biennis with attractive habit and flowers. There are also forms with fleshy edible roots, used as a vegeta-ble, which were more commonly grown in the nineteenth century than today. Evening primrose leaves, shoots, roots, and seed pods were con-sumed by American Indians as food.
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Oenothera biennis L.
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Agricultural and Commercial Aspects The most significant current economic value of the species lies in its use as a diversification crop.
Although GLA has been obtained by fermentation of some yeasts and other fungi, and from currants (Ribes species), the chief commercial sources are evening primrose and borage (Borago officinalis L.). Companies have engaged in a boastful debate about the comparative efficacy of GLA in their preparations made from evening primrose on the one hand, and from borage on the other. Whether borage or evening primrose is more competitive for GLA production depends on climatic and edaphic factors at a particular location. In Canada, both species are grown. Borage has a higher GLA con-tent, but non-shattering cultivars are not grown in Canada, so that harvest is difficult. Borage is much more suitable for the Canadian prairies, where avail-able cultivars of evening primrose do not overwinter reliably. However it isn’t essential to grow evening primrose as a biennial: in Eastern Canada it is of-ten started in greenhouses in mid-winter and transplanted to the field where it is grown as an annual.
As a cultivated plant, evening primrose is toler-ant of a variety of soil types and a range of pH, but soils that are prone to crusting after rains and waterlogged soils should be avoided. If planted at too high a density (150 plants m2) the plants may not bolt.
Evening primrose crops are raised in temperate areas of northern and eastern Europe, North Amer-ica, and Australasia. US production is centered in North and South Carolina, Texas, and Oregon. Ca-nadian production is centered in Nova Scotia and Ontario. Experimental production in Manitoba has been disappointing. Annual world production of seed has increased at least 20 fold in the last 20 years, and is currently about 4,000 tonnes. Com-bined US, and Canadian annual production is less than 200 tonnes. In good market years, several hundred ha of evening primrose may be grown in Canada.
Wild evening primrose plants shed their seeds when a pod matures, and since the pods don’t ma-ture simultaneously, harvest of seeds is difficult.
Nevertheless, seed is gathered from wild plants in northeast China. Most modern evening primrose cultivars have non-shedding pods, which has sim-plified harvest and reduced seed loss. Crop yields of over 2 tonnes/ha have been recorded in Nova Scotia, although much lower yields are frequent. In Ontario, depending on the rather volatile market
and variable production, a hectare may result in a gross financial return of $1,000–2,000.
The future of evening primrose as a pharmaco-logical crop in Canada is uncertain because of competition from other countries and the unreli-ability of the present market. Hemp (Cannabis sativa L.) is attracting considerable interest as a new crop in Canada, to be grown not only for fi-ber but also for its high-GLA seed oil. Still another potential source of competition is the possibility that genetic engineers will splice the capacity to produce GLA into crops such as canola (Brassica species).
Certainly the demand for GLA will continue to grow, and at least from time to time it may be an-ticipated that evening primrose crops will be grown in Canada on a contracted basis. With respect to climate and native germplasm, Canada is in a good position to develop its share of the evening prim-rose market.
Myths, Legends, Tales, Folklore, and Interesting Facts
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Some evening primrose seeds have been shown to live to 80 years in the soil.Oenothera biennis (evening primrose)
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Hugo de Vries (1848–1935) was a world-famous student of evolution who, at the beginning of the 20th century, theorized that new species arise by spontaneous changes in individuals called mu-tations (Charles Darwin had earlier learned that such changes (called “sports” at the time) oc-cur, but did not appreciate their importance for the mechanism of evolution). Unfortunately for de Vries, he chose evening primrose to demon-strate his theory. Later, scientists learned that the odd genetic system of evening primrose was re-sponsible for the generation of altered individu-als that de Vries was labelling as mutations and that this system did not occur in many other plants. As a result, his theory was discredited, although de Vries did contribute substantially to evolutionary theory.Selected References
Baker, J. 1998. Evening primrose. In Richters second commercial herb growing conference - transcripts.
Edited by R. Berzins, H. Snell and C. Richter.
Richters, Goodwood, ON. pp. 67–85.
Barthell, J.F., and Knops, J.M.H. 1997. Visitation of evening primrose by carpenter bees: evidence of a
“mixed” pollination syndrome. Southwestern Natu-ralist 42(1): 86–93.
Baskin, C.C., and Baskin, J.M. 1993. Germination re-quirements of Oenothera biennis seeds during burial under natural seasonal temperature cycles. Can. J.
Bot. 72: 779–782.
Belisle, D. 1991. Potential of evening primrose in Can-ada. Alternative Crops Notebook 4: 4–5. [Reprinted from BioOptions, Newsletter of the Center for Alter-native Plant & Animal Products 1(5), 1990.]
Brandle, J.E., Court, W.A., and Roy, R.C. 1993.
Heritability of seed yield, oil concentration and oil quality among wild biotypes of Ontario evening prim-rose. Can. J. Plant Sci. 73: 1067–1070.
Briggs, C.J. 1986. Evening primrose: La belle de nuit, the king’s cureall. Can. Pharm. J. 199: 248–252.
Budeiri, D., Po, A.L.W., and Dornan, J.C. 1996. Is evening primrose oil of value in the treatment of premenstrual syndrome? Controlled Clin. Trials 17(1): 60–68.
Court, W.A., Hendel, J.G., and Pocs, R. 1993. Deter-mination of the fatty acids and oil content of eve-ning primrose (Oenothera biennis L). Food Res.
Int.. 26:181–186.
Cisowski, W., Zielinska-Stasiek, M., Luczkiewicz, M., and Stolyhwo, A. 1993. Fatty acids and triacylglycerols of developing evening primrose (Oenothera biennis) seeds. Fitoterapia 64: 155–162.
Dietrich, W.M., and Wagner, W.L. 1988. Systematics of Oenothera section Oenothera subsection Raimannia and subsection Nutantigemma (Onagracceae). Syst.
Monogr. 24: 1–91.
Dietrich, W., Wagner, W.L., and Raven, P.H. 1997.
Systematics of Oenothera section Oenothera sub-section Oenothera (Onagraceae). American Society of Plant Taxonomists, Ann Arbor, MI. Syst. Bot.
Monogr. 50. 234 pp.
Ensminger, P.A., and Ikuma, H. 1987. Photoinduced seed germination of Oenothera biennis L. I. Gen-eral characteristics. Plant Physiol. 85: 879–884.
Ensminger, P.A., and Ikuma, H. 1987. Photoinduced seed germination of Oenothera biennis L. II. Analy-sis of the photoinduction period. Plant Physiol. 85:
885–891.
Ensminger, P.A., and Ikuma, H. 1988. Photoinduced seed germination of Oenothera biennis L. III. Anal-ysis of the postinduction period by means of tem-perature. Plant Physiol. 86: 475–481.
Gates, R.G. 1957. A conspectus of the genus Oenothera in eastern North America. Rhodora 59: 9–17.
Gates, R.G. 1958. Taxonomy and genetics of Oenothera.
Uitgeverij Dr. W. Junk, Den Haag, Cambridge, MA.
115 pp.
Gregory, D.P. 1963. Hawkmoth pollination in the ge-nus Oenothera. Aliso 5: 357–384.
Gross, K.L. 1985. Effects of irradiance and spectral quality on the germination of Verbascum thapsus and Oenothera biennis seeds. New Phytol. 101: 531–542.
Gross, K.L., and Kromer, M.L. 1986. Seed weight ef-fects on growth and reproduction in Oenothera biennis L. Bull. Torrey Bot. Club 113(3): 252–258.
Hall, I.V., Steiner, E., Threadgill, P., and Jones, R.W.
1988. The biology of Canadian weeds. 84. Oenothera biennis L. Can. J. Plant Sci. 68: 163–173.
Hanczakowski, P., Szymczyk, B., and Wolski, T. 1993.
The nutritive value of the residues remaining after oil extraction from seeds of evening primrose (Oenothera biennis L.). J. Sci. Food Agric. 63: 375–376.
Horrobin, D.E. 1990. Gamma linolenic acid. Reviews in Contemporary Pharmacology 1: 1-45.
Hulan, H.W., Hall, I.V., Nash, D.M., and Proudfoot, F.G.
1987. Composition of native evening primrose seeds collected from western Nova Scotia. Crop. Res. Edin-burgh (Scottish Academic Press) 27(1): 1–9.
Kerscher, M.J., and Korting, H.C. 1992. Treatment of atopic eczema with evening primrose oil: rationale and clinical results. Clin. Investig. 70(2): 167–171.
Kromer, M., and Gross, K.L. 1987. Seed mass, geno-type, and density effects on the growth and yield of Oenothera biennis L. Oecologia (Berlin) 73: 207–
212.
Lapinskas, P. 1989. Commercial exploitation of alterna-tive crops, with special reference to evening prim-rose. In New crops for food and industry. Edited by G.E. Wickens, N. Haq and P. Day. Chapman and Hall, London. pp. 216–221.
Levin, D.A., Howland, G.P., and Steiner, E. 1972. Pro-tein polymorphism and genic heterozygosity in a pop-ulation of the permanent translocation heterzygote, Oenothera biennis. Proc. Nat. Acad. Sci. USA 69(6):
1475–1477.
Levy, A., Palevitch, D., Ranen, C. 1993. Increasing gamma linolenic acid in evening primrose grown under hot temperatures by breeding early cultivars.
Acta Hort. 330: 219–225.
Loughton, A., Columbus, M.J., and Roy, R.C. 1991.
The search for industrial uses of crops in the diver-sification of agriculture in Ontario. Altern. Crops Notebook 5: 21–27.
Morrison, K.D., and Reekie, E.G. 1995. Pattern of de-foliation and its effect on photosynthetic capacity in Oenothera biennis. J. Ecol. 83: 759–767.
Mukherjee, K.D., and Kiewitt, I. 1987. Formation of gamma linolenic acid in the higher plant evening primrose (Oenothera biennis L.). J. Agric. Food.
Chem. 35: 1009–1012.
Munz, P.M. 1965. North American Flora, Series II, Part 5 - Onagraceae. The New York Botanical Gar-den, NY. 231 pp.
Paccalin, J., Mendy, F., Bernard, M., Delhaye, N., and Spielmann, D. 1983. Rediscovery of an oleaginous plant: Oenothera biennis. The importance of gamma-linolenic acid in nutrition. Bull. Acad. Natl. Med.
167: 923–931. [In French.]
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Raven, P.E., Dietrich, W., and Stubbe, W. 1979. An outline of the systematics of Oenothera subsect.
Euoenothera (Onagraceae). Syst. Bot. 4: 242–252.
Reekie, E.G., and Reekie, J.Y.C. 1991. The effect of re-production on canopy structure, allocation and growth in Oenothera biennis. J. Ecol. 79: 1061–1071.
Reeleder, R.D. 1994. Factors affecting infection of evening primrose (Oenothera biennis) by Septoria oenotherae. Can. J. Plant Pathol. 16: 13–20.
Reeleder, R.D., Monet, S., Roy, R.C., and Court, W.A.
1996. Dieback of evening primrose: Characteristics of Septoria oenotherae, its interactions with Botry-tis cinerea, and use of fungicides to manage dis-ease. Can. J. Plant Pathol. 18: 261–268.
Rostanski, K. 1982. The species of Oenothera L. in Britain. Watsonia 14: 1–34.
Roy, R.C. 1990. Health food plant may be alternative for tobacco growers. Communication Branch, Agri-culture Canada, Ottawa. Agri-Features 2106: 1–3.
Roy, R.C., White, P.H., More, A.F., Hendel, J.G., Pocs, R., and Court, W.A. 1993. Effect of transplanting date on the fatty acid conposition, oil content and yield of evening primrose (Oenothera biennis L.) seed. Can. J. Plant Sci. 74: 129–131.
Russell, G. 1988. Physiological restraints on the eco-nomic viability of the evening primrose crop in east-ern Scotland. Crop. Res. (Edinburgh) 28(1): 25–33.
Simpson, M.J.A. 1994. A description and code of de-velopment of evening primrose (Oenothera spp.).
Ann. Appl. Biol. 125: 391–397.
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331: 121–128.
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Skvarla, J.J., Raven, P.H., and Praglowski, J. 1976.
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Soc. Symp. Ser. 1: 447–479.
Wagner, W.L., Stockhouse, R.E., and Klein, W.K. 1985.
The systematics and evolution of the Oenothera caespitosa species complex (Onagraceae). Missouri Botanical Gardens, Allen Press, Inc., Lawrence, KS.
103 pp.
Wilson, R. 1989. An alternative crop profile. Crop Devel-opment Division, Agriculture Canada. Altern. Crop Notebook 1: 1–5.
Yaniv, Z., and Perl, M. 1987. The effect of temperature on the fatty acid composition of evening primrose (Oenothera) seeds during their development, stor-age and germination. Acta Hort. 215: 31–38.
Yaniv, Z., Ranen, C., Levy, A., and Palevitch, D. 1989.
Effect of temperature on the fatty acid composition and yield of evening primrose (Oenothera lamarckiana) seeds. J. Exp. Bot. 40: 609–614.
World Wide Web Links
(Warning. The quality of information on the internet varies from excellent to erroneous and highly misleading. The links below were chosen because they were the most informative sites located at the time of our internet search. Since medici-nal plants are the subject, information on medicimedici-nal usage is often given. Such information may be flawed, and in any event should not be substituted for professional medical guidance.)
Evening primrose, Oenothera biennis, Agriculture & Agri-Food Canada, Southern Crop Protection &
Food Research Centre:
http://res.agr.ca/lond/pmrc/study/newcrops/eprim.html Sask Ag & Food: Farmfacts, evening primrose:
http://www.agr.gov.sk.ca/saf/farmfact/sce0190.htm Paintings of Mary Vaux Walcott [color painting]:
http://chili.rt66.com/hbmoore/Images/Walcott/Oenothera_biennis-2.jpg
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Oplopanax horridus (devil’s club)
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Oplopanax horridus (J.E. Smith) Miq. Devil’s Club
Synonyms: Echinopanax horridus (Sm.) Decne. &
Planch. ex H.A.T. Harms; Fatsia horrida (Sm.) Benth. & Hook. f.; Panax horridus Sm.
The genus name is derived from the Greek hoplon, weapon, + Panax, i.e., an “armed ginseng,”
references to the fearsome spines of the plant and to its membership in the same family (Araliaceae) as ginseng. Frequently the genus name has been considered to be a neuter name in Latin, so that the specific epithet ends in “um” (O. horridum). How-ever, the Botanical Code of Nomenclature requires that names ending in panax be treated as mascu-line, so that the name must be spelled E. horridus.
English Common Names
Devil’s club (sometimes ungrammatically hyphen-ated as “devil’s-club”), Alaskan ginseng.
Devil’s club is sometimes confused with devil’s claw (Harpagophytum procumbens DC.); the lat-ter, an African plant, is commonly marketed in North America as an over-the-counter herbal prod-uct, with the information that it has been
Devil’s club is sometimes confused with devil’s claw (Harpagophytum procumbens DC.); the lat-ter, an African plant, is commonly marketed in North America as an over-the-counter herbal prod-uct, with the information that it has been