FWD 2 HerbalGram: The Genus Ligusticum in North America: An Ethnobotanical Review with Special Emphasis upon Species Commercially Known as ‘Osha’


Issue: 89 Page: 40-51

The Genus Ligusticum in North America: An Ethnobotanical Review with Special Emphasis upon Species Commercially Known as ‘Osha’

by Christina Turi, Susan J. Murch

HerbalGram. 2010; American Botanical Council



The genus Ligusticum consists of 40 to 60 perennial species found within boreal and mountainous regions of the world.1,2 In Asia, native species of Ligusticum are held in great esteem for their therapeutic actions.3 In North America, endemic species collectively known as osha* are also sought after by consumers within the dietary supplement and natural health products industry, in addition to being revered among indigenous groups.4 Although research investigating the medicinal attributes of Ligusticum species commonly used within Traditional Chinese Medicine has been ongoing since the 1960s,3 research into the medicinal efficacy of North American species is more recent. A literature search of “Ligusticum” in Web of Science™ on September 30, 2010, found a total of 257 articles in the scientific literature. Among these articles, approximately 85% pertained to species endemic to Asia (Figure 1).

In North America, First Nation’s (i.e., Native Americans’) uses of Ligusticum species include food, hunting, spirituality, and medicine5 (Table 2). Ligusticums grow in North American regions ranging from north-eastern parts of Quebec, the Maritimes, and Ontario in Canada, to the Eastern United States as far south as Florida. Ligusticum species are also found in the Mexican Sierra Madre and through regions of New Mexico, Colorado, Wyoming, Montana, Alaska, British Columbia, and Western coastal States and Provinces6 (Figure 2). Ecological zones where Ligusticum species known as osha are found in North America include montane to subalpine meadows,7 spruce and aspen belts, windswept parks, oak and mature stands of conifers,8 pine-oak forests of the Northern Sierra Madre Occidental and central to southern Rocky Mountains of the United States,9 mountain ridges, within dry or moist slopes,10 moist forests from the spruce-fir vegetation zone to foothills,11 sagebrush meadows on rocky soils4 and among subalpine zones and altitudes up to 10,000 feet.12 Ligusticum porteri was also identified as a dominant species among Aspen communities in Gunnison County, Colorado (Figure 3).13

Botany and Taxonomy

Ligusticums belong to the family Apiaceae, otherwise known as the Umbelliferae, or the carrot family. In North America, there are 12 species of Ligusticum.14 Among these, L. porteri, L. canbyi, L. grayi, L. tenuifolium, and L. filicinum are most commonly known as osha.4,8* Like other genera belonging to Apiaceae, Ligusticums are identified by their compound umbels, white or sometimes pinkish flowers, distinctly ribbed and narrowly winged fruits, taproot system, basal leaves, and ternate to pinnately compound or dissected leaves.15,16 Though challenging at times, differentiation between species collectively known as osha is accomplished through examination of plant height, leaf morphology, and habitat (Table 1, Figure 2). Furthermore, species of osha can be mistaken with other members of the Apiaceae family, including the genera Angelica, Conium, and Lomatium.4 This can be problematic, since Ligusticum species resemble toxic species such as water-hemlock (Cicuta douglasii, Apiaceae) and poison-hemlock (Conium maculatum, Apiaceae).12 Differentiation of Ligusticum species from poison-hemlock is achieved through examining root morphology and scent. Typically, but not always, the roots of poison-hemlock are smooth, purplish, without leaf base remnants15 and believed to have a “dead mouse like scent.”8 Roots from water-hemlock are said to be tuberous or fibrous, soft in texture, purplish, emerging from a thickened crown, and exuding a parsnip scent.15 Root crowns of Ligusticums, on the other hand, have basal leaves,16 and possess a distinctive celery-like odor similar to lovage (Levisticum officinale).8,12

Propagation

In China, growers are expected to follow Good Agricultural Practices (GAP). Currently, several GAP Ligusticum chuanxiong centers for cultivation are under development within the Dujiangyan County, Sichuan Province.17 Traditionally, roots are collected in May and processed using a variety of methods, including stir-frying and sun- or oven-heating.18 In North America, knowledge of the propagation and growth of Ligusticum species is limited and suggests osha is difficult to grow.4,19,20 Renowned herbalist and author Michael Moore described cultivating osha as “[a]lmost impossible. Even in northern New Mexico (elevation averaging from 6,000 to 8,000 feet) where it is most widely used, the people are not able to cultivate it for their own consumption…If high-country curanderas and abuela can’t grow Ligusticum porteri, who am I to try it?”8

Ligusticum porteri seeds can be sown in both garden and greenhouse environments during the fall, midwinter, or spring with an expected germination rate of 70%. Period of germination will vary, ranging between 6 to 12 weeks. However, it is not uncommon for seeds to remain dormant until the following growing season.21 Stratifying seeds at 4.4 C for 12 weeks under moist, cold conditions before placing them on a mist propagation bench at 21.1 C can improve germination rates for unattached seeds. 6 weeks of stratification in a cold, moist environment followed by 12 weeks in a germination chamber is beneficial for attached seeds (although only 11% of attached and unattached seeds from umbels reportedly emerged).20 Other research has shown that seed viability is improved by at least 10 weeks of stratification, exposure to GA3,and by lightly covering seeds with soil before exposing them to light in a greenhouse.22

After 8 weeks, seedlings can be transplanted to a garden with an expected survival rate of 80% after 3 years.22 Crown cuttings may be an effective alternate strategy for propagation of osha. It is believed that the branching cluster of root material associated with the taproot system of osha makes transplanting feasible.21 This notion is reinforced, given that 90% of crown cuttings consisting of both root and stem material from L. porteri appeared to root successfully.20

Harvest and Processing

Ligusticum species known as osha are wild-harvested and sold within the herb and dietary supplement industry (known as the Natural Health Products industry in Canada). Market analysis reports that dried roots from L. porteri will fetch a price between 30 to 50 US dollars per pound.22† Given that successful commercial cultivation is yet to be established, questions regarding sustainable harvesting practices have been raised.4 Data collected over the last 10 years by the American Herbal Products Association (AHPA) indicates that wild collection and demand for L. porteri increased in the late 1990s and has remained constant for the last decade23 (Figure 4). Roots are harvested during early spring before budding,20 or during the months of August and September once the Ligusticum’s flowers have all died8,24 and its leaves have turned gold.21 Typically, older plants are favoured for collection, usually those between 5 to 10 years of age20 and possessing numerous flowering stalks with large rosettes. Seeds can be collected and detached once the umbel has reached maturity in the fall.21 A variety of rituals have been associated with the collection of Ligusticum roots. For example, Turner et al. (1980) observed that the Okanagan-Colville people of southern British Columbia and northern Washington had a tradition of expressing gratitude before digging up the roots of L. canbyi.

Once collected, roots from Ligusticum species should be cleared of soil and dried for approximately 2 to 3 weeks and stored away from moisture to ensure a long shelf life of osha products.8,22,24 Alternatively, roots can be dried in a dehydrator and removed when easily broken apart.21 Turner et al. (1980) found that the Okanagan-Colville would store the roots of L. cambyi, Canby’s Lovage, in a medicine bag or tobacco pouch,24 while Jordan (2008) recorded that the Plains Apache would store roots of L. porteri (also known as medicine fat) in “the family medicine bundle” and use its smoke to purify the bundles annually during spring rituals.25

Ethnobotany

(See Table 2)

Zoopharmacognosy

Kodiak bears (Ursus arctos) have been observed masticating the roots of L. porteri, sometimes known as bear root, and rubbing the salivated contents over their bodies, possibly to medicate infected topical wounds.26 The Navajo believe that bear root’s medicinal use was given to them by bears, which has led them to use its roots for treating ailments including skin abrasions.27-29

Poison and Hunting

According to Lantis (1959), the Eskimo considered mature plants from L. scoticum to be mildly poisonous,30 while Garth (1953) observed that roots of L. grayi were used for hunting and poisoning fish among the Atsugewi of northern California.31 Similarly, Campbell (1958) records that roots from L. porteri were used as a fish poison by the Tarahumar of northern Mexico.9,32

Spirituality, Protection, Smoke or Incense

Ceremonial uses for L. porteri and L. canbyi are documented for North American indigenous groups. Bye and Linares (1981-1986), Curtin (1965), Schulman and Smith (1962), Kennedy (1978), and Ford (1975) observed the Tarahumara using infusions of L. porteri during ritual curing ceremonies or for protecting individuals against witches and rattlesnakes.9 Similarly, Camazine and Bye (1980) report that both patient and medicine man in Zuni culture would chew on the root during healing rituals.33 Jones (1992) describes the Arapahoe and Pawnee using the roots of L. porteri during sweat lodge ceremonies and purification rites.34 While staying with the Tonkawa, Opler (1902) recounted that one must possess L. porteri in order to partake in the peyote ceremony:

“The only way you could get in that ceremonial tent was to have long hair, your face painted to suit them, and an Indian shirt made of any colors you liked. And you had to have buckskin leggings or some kind of leggings such as the Indian used to wear. You had to have a long loin-cloth and you had to wear a sheet wrapped around you. You had to have different kinds of odorous herbs; you had to have Osha.”35

Among the Plains Apache, Jordan (2008) reports that roots of L. porteri are thrown over fire to help console individuals during rituals associated with mourning.25 Turner et al. (1980) found that aromatic smoke produced from roots of L. canbyi were used by the Okanagan-Colville to release individuals from trance, possession by spirits such as the “blujay spirit,” or unconsciousness.24 During the early 1900s, chewing the roots of L. scoticum was considered a substitute for tobacco.36 Similarly, L. canbyi can be added to tobacco37 and smoked in order to produce a menthol taste. It has been suggested that smoking its roots provides a relaxant effect, which most likely explains its common name “Indian marijuana.” Ligusticum canbyi can be used as incense.37 Exposure to excess amounts of smoke is believed to have a strong sedative effect upon children.24 Furthermore, Sturtevant (1958) reports that members of the Cherokee would smoke the roots from L. canadense in order to treat stomach problems.38

Food

Although it is likely that most species have been used for cooking or substituted with other species, L. californium, L. canadense, L. grayi, L. porteri, and L. scoticum are indicated for culinary use within the literature (Table 2). It is believed that Ligusticums produce a chervil-celery-parsley flavor;8 and as a result, the leaves, seeds, and roots are used to season meats, beans, and chilis.8,9,39-43 Ligusticums can be eaten without preparation or by preparing leaves like greens through boiling or adding to salads.31,44-46 Stalks can be used similar to celery,47 while roots can be boiled, thrown into salads and soups, or eaten raw by peeling back the stem.36,48,49

Medicine

The use of osha to treat sickness and promote “well-being” has played an important role among North American indigenous cultures for centuries. In 1988, researchers in Utah discovered a medicine bundle containing L. porteri, thought to be 200 to 400 years old. After analyzing the contents of the package, it was suggested that these items were not simply “stash items,” but rather of great significance.50 Medicinal applications of Ligusticum species include the following: antirheumatic;9 treatment for hair, lice, ticks, wounds, and skin and ear infections;9, 24, 37 as an anti-convulsive, analgesic, and fever treatment,9,31,37 for gastrointestinal problems or dietary aid,9,31,38,51 for colds, coughs, sore throats, and pulmonary or respiratory aid,9,24,31,37,52-58 treatment for anaemia, diabetes, and circulation or heart problems9,37,53 (Table 2). Personal preference towards a specific Ligusticum species varies throughout North America. The dispensatory of the United States indicated in 1918 that L. filicinum “is a highly prized expectorant” within Utah and the surrounding states.56 Smith (1929) states that the Menomini describe 20 kinds of “osha root,” the most powerful being the Mani’k (L. filicinum).57 More recently, L. porteri was identified by Appelt (1985) as a commonly used herb in Hispanic communities of the San Luis Valley, Colorado.59

Medicinal Chemistry

Much of the research that exists investigating the molecular composition of Ligusticum species includes derivatives of coumarins and phthalides (such as Z-ligustilide and butylidenephthalide).3,60 Over the last 50 years, Asian species of the genus Ligusticum have facilitated the discovery of many novel compounds. Approximately 40 phthalides have been isolated from L. chuanxiong,20 from L. officinale, and 20 from L. wallichi.3 Table 3 presents a summary of compounds identified from species commonly known as osha.3,60-74 Although numerous compounds have been identified, Z-ligustilide and butylidenephthalide are most commonly purported to facilitate their therapeutic effects. Medicinally, Z-ligustilide is used for the following effects: smooth muscle relaxation, vasodilation, and as an insecticidal, antibacterial, antifungal, and anti-inflammatory. Further, antianginal, antihypertensive, antispasmodic, vasodilatory, and serotonergic activity—as well as selectivity upon the central cholinergic neuronal system in rats—have been associated with butylidenephthalide.3

Bioassay

Although a variety of therapeutic properties have been associated with Ligusticums found within Traditional Chinese Medicine, the exact mechanisms which underline these attributes are still unclear.3 With respect to pharmacology, extracts of L. porteri have been shown to inhibit the infective capacity of several strains of bacteria.75 On the other hand, essential oil and dichloromethane extracts did not exhibit significant antimicrobial activity against a norfloxacin-resistant strain of Staphylococcus aureus, but sensitivity to norfloxacin was restored.64 Additionally, ethanol extracts taken from roots of L. porteri (0.05g/ml) have shown inhibitory activity against both Bacilus subtilis and Pseudemonae syringae.76 In various model systems to test pain susceptibility, extracts of L. porteri have shown significant antinociceptive activity.77 Orally administered extracts of L. porteri have shown antihyperglycymic effects in mice, with (Z)-6,6’,7,3’a-Diligustilide, (Z)-ligustilide, and 3-(Z)-butylidenephthalide contributing to the observed effects.78 In contrast, models of cancer metabolism did not find significant inhibition of proliferation of tumor cells or cytotoxic effects.79 It is interesting to note that some toxicity has been determined in roots of L. porteri (LD50 = 1085 mg/kg in mice and CL50 = 777.98 µg/ml with Artemia salina lethality test 80) suggesting further inquiry into safe dosage should be undertaken.

Conclusions

There is still much to learn about North American Ligusticum species with respect to their botanical diversity, growth, cultivation, phytochemistry, and potential therapeutic uses. These studies are especially timely since demand for North American Ligusticum species sold as osha has increased steadily and some collections may have reduced wild populations of certain species. Therefore, monitoring of wild populations and sustainable wild-crafting and cultivation will help ensure that sufficient osha germplasm is available for future generations of the natural health products and herbal dietary supplement industry.

Christina E. Turi is a PhD student in the department of Biology at the University of British Columbia, Okanagan Campus, Kelowna, British Columbia, Canada.

Susan J. Murch, PhD, is an Associate Professor of Chemistry and Canada Research Chair in Natural Products Chemistry at the University of British Columbia, Okanagan Campus, Kelowna, British Columbia, Canada.

*One of the major problems with commercial osha is the misidentification of species. Some suppliers, product manufactures, and some herbalists do not differentiate between species of Ligusticum, using osha interchangeably as a common name. A CITES report from 2000 suggests herbal companies often purchase unidentified species of Ligusticum from wildcrafters which they call ‘L. porteri or osha.’ Consequently, L. filicinum, L. canbyi, and L. tenuifolium are species often found within the herbal market as osha or ‘L. porteri.’ Moore’s Medicinal Plants of the Mountain West (2003) refers to L. filicinum, L. grayi, L. porteri, L. apifolium, L. californicum, L. canbyi, L. tenuifolium, and L. verticillatum as osha but emphasizes the importance of L. porteri as ‘True Osha.’ Furthermore, Tilford’s Edible and Medicinal Plants of the West (1997) places greater emphasis upon L. canbyi as osha, while also acknowledging other species of Ligusticum as osha including L. porteri. Finally, Cech’s Growing at-risk Medicinal Herbs (2002) considers all species belonging to the family Ligusticum as osha. A major need for research is the development of an effective way to differentiate and identify dried root material to distinguish species but such methods are currently not available.

† In 1999, a group of scholars from the University of Maryland Program in Sustainable Development and Conservation Biology submitted a draft proposal to the US Fish & Wildlife Service Office of Scientific Authority. This document suggested that L. porteri be listed in Appendix II of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES).4,81 Currently, no Ligusticum species known commercially as osha are CITES listed.82

References

  1. Ben shu G, Fading P, Watson M. Ligusticum linnaeus, sp. pl. 1:250. 1753. Flora of China. 2005;14:140-150. Mabberly D. The Plant Book. New York, NY. USA: Cambridge University Press; 1997.
  2. Beck JJ, Chou S. The structural diversity of phthalides from the Apiaceae. J Nat Prod. 2007;70:891-900.
  3. The Scientific Authority of the United States of America. Species proposal for the 12th Meeting of the Conference of the Parties: Ligusticum porteri. Convention on International Trade in Endangered Species of Wild Fauna and Flora. Convention on International Trade in Endangered Species of Wild Fauna and Flora. 2000.
  4. Moerman D. Native American Ethnobotany. Portland, OR: Timber Press, Inc; 2009. Canadian Museum of Nature Herbarium, Vascular Plant Collection-University of Washington Herbarium, European Environment Agency, et al. Biodiversity occurrence data for Ligusticum species. Available at: data.gbif.org/species/13196690/. Accessed July 6, 2010.
  5. Scotter G, Flygare H. Wildflowers of the Rocky Mountains. North Vancouver, British Columbia: Whitecap Books; 2007. Moore M. Medicinal Plants of the Mountain West. Sante Fe, NM: Museum of New Mexico Press; 2003.
  6. Linares E, Bye RA. A study of 4 medicinal plant-complexes of Mexico and adjacent United States. J Ethnopharmacol. 1987;19:153-183.
  7. Kershaw L. Edible and Medicinal Plants of the Rockies. Edmonton, Alberta: Lone Pine Publishing; 2000. Dodson C, Dunmire W. Mountain Wildflowers of the Southern Rockies. Santa Fe, NM: University of New Mexico Press; 2007.
  8. Tilford G. Edible and Medicinal Plants of the West. Missoula, MT: Mountain Press Publishing Company; 1997. Crawford JL, McNulty SP, Sowell JB, Morgan MD. Changes in Aspen communities over 30 years in Gunnison County, CO. Am Midl Nat. 1998;140:197-205.
  9. United States Department of Agriculture. Ligusticum Species. Available at: plants.usda.gov/java/profile?symbol=LIGUS. Accessed 1/09, 2010.
  10. Applequist W. Root anatomy of Ligusticum species (Apiaceae) sold as osha compared to that of potential contaminants. Journal of Herbs and Medicinal Plants. 2005;11:1-11.
  11. Hitchcock C, Cronquist A. Flora of the Pacific Northwest. Seattle, WA: University of Washington Press; 1996. Li SL, Lin G, Tam YK. Time-course accumulation of main bioactive components in the rhizome of Ligusticum chuanxiong. Planta Med. 2006;72:278-280.
  12. Li S, Yan R, Tam Y, Lim G. Post-harvest alteration of the main chemical ingredients in Ligusticum chuanxiong HoRT. (rhizoma chuanxiong). Chem Pharm Bull. 2007;55:140-144.
  13. West K, Jackson S. Research to determin osha’s economic potential as a sustainable agricultural crop. HerbalGram. 2004;62:15.
  14. Panter KL, Ashley RE, Guernsey KM, Johnson CM. Preliminary studies on propagation of osha. Horttechnology. 2004;14:141-143.
  15. Cech R. Growing at-risk medicinal herbs: cultivation, conservation and ecology. Williams, OR: Horizon Herbs Publication; 2002.
  16. Terrell B, Fennell A. Osha (bear root): Ligusticum porteri J.M. coult. & rose var. porteri. Native Plants Journal. 2009;10:111-118.
  17. The American Herbal Products Association. Tonnage survey of selected North American wild-harvested plants, 2004-2005. Silver Spring, MD: American Herbal Products Association; 2007.
  18. Turner N, Bouchard R, Kennedy D. Ethnobotany of the Okanagan-Colville Indians of British Columbia and Washington. Victoria, BC: British Columbia Provincial Museum; 1980.
  19. Jordan J. Plains Apache Ethnobotany. Norman, OK: University of Oklahoma Press.
  20. Newton P, Wolfe N. Can animals teach us medicine? British Medical Journal. 1992;305:1517-1518.
  21. Huffman M. Animal self-medication and ethnomedicine: exploration and exploitation. Proceedings of the Nutrition Society. 2003;62:371-381.
  22. Cowen R. Medicine on the wild side. Science News. 1990;138:280-282.
  23. Grisanzio L. Animal. 1992:26-30.
  24. Lantis M. Folk medicine and hygiene. Anthropological Papers of the University of Alaska. 1959:1-75.
  25. Garth T. Atsugewi ethnography. Anthropological Records. 1953;14:140-141.
  26. Campbell P. Tarahumar fish stupefaction plants. Economic Botany. 1958;12:95-102.
  27. Camazine S, Bye R. A study of the medical ethnobotany of the Zuni Indians of New Mexico. Journal of Ethnopharmacology. 1980;2:365-388.
  28. Feather Jones: Fragments of Earth Wisdom. In: Tierra M, American Herbalist Guild, eds. American Herbalism: Essays on Herbs & Herbalism: Crossing Press; 1992: 41-45.
  29. Opler M. A description of Tonkawa peyote meeting held in 1902. American anthropologist. 1939;41:433-439. Hedrick UP. 1919 Sturtevant’s Edible Plants of the World. Available at: www.henriettesherbal.com/eclectic/sturtevant/ligusticum.html, Accessed 07/10, 2010.
  30. Hart J. Montana Native Plants and Early Peoples. Helena, MT: Montana Historical Society Press; 1992.
  31. Sturtevant W. The Mikasuki Seminole:Medical Beliefs and Practices. [PhD]. Yale University; 1954.
  32. Castetter E, Opler E. Ethnobiological studies in the American Southwest III: the ethnobiology of the Chirichahua and Mescalero Apache. University of New Mexico Bulletin. 1936;4:1-63.
  33. Jones A. Nauriat Niginaqtuat = Plants that we Eat. Kotzebue, AK: Maniilaq Association Traditional Nutrition Program; 1983.
  34. Rousseau J. Notes sur l’Ethnobotanique d’Anticosti. Archives de Folklore. 1946;1:60-71.
  35. Anderson J. Plants used by the Eskimo of the northern Bering Sea and arctic regions of Alaska. American Journal of Botany. 1939;26:714-716.
  36. Ager TA, Ager L. Ethnobotany of the Eskimos of Nelson Island, Alaska. Artic Anthropology. 1980;27:26-48.
  37. Perry M. Food Use of “Wild” Plants by Cherokee Indians. [M.S.]. The University of Tennessee; 1975.
  38. Withoft J. Cherokee Indian use of potherbs. Journal of Cherokee Studies. 1977;2:250-255.
  39. Hamel P, Chiltoskey M. Cherokee Plants and their Uses: A 400 Year History. Sylva, NC: Hearld Publishing Co.;1975.
  40. Heller C. Edible and Poisonous Plants of Alaska. Anchorage, AK: University of Alaska; 1953.
  41. Hatfield G. Hatfield’s Herball. London: Penguin Books Ltd.; 2007.
  42. Baker M. The Ethnobotany of the Yurok, Tolowa, and Karok Indians of Northwest Californica. [M.A.]. Arcata, CA: Humbolt State University; 1981.
  43. Harrison M. The botanical parts of the Patterson bundle. The Utah Native Plant Society. UT:2000:7.
  44. Schenck S, Gifford E. Karok ethnobotany. Anthropological Records. 1952;13:377-372.
  45. Gifford E. Ethnographic notes on the Southwestern Pomo. Anthropological Records. 1967;25:10-15.
  46. Goodrich J, Lawson C. Kashaya Pomo Plants. American Indian Studies Center, Los Angeles, CA: University of California Press; 1980.
  47. Sayre L. A Manual of Organic Materia Medica and Pharmacognosy. Available at: www.henriettesherbal.com/eclectic/sayre/ligusticum.html. Accessed July 20, 2010.
  48. Felter H, Lloyd J. 1898 King’s American Dispensatory. Available at: www.henriettesherbal.com/eclectic/kings/angelica-arch.html. Accessed July 20, 2010.
  49. Remington J, Wood H. 1918 The Dispensatory of the United States of America. Available at: www.henriettesherbal.com/eclectic/usdisp/ligusticumfili.html. Accessed July 20, 2010.
  50. Smith H. Materia medica of the Bella Coola and neighbouring tribes of British Columbia. National Museum of Canada Bulletin. 1929;56:47-68.
  51. Train P, Henrichs J, Archer W. Medicinal Uses of Plant by Indian Tribes of Nevada. Washington, DC: Department of Agriculture; 1941.
  52. Appelt GD. Pharmacological aspects of selected herbs employed in Hispanic folk medicine in the San Luis Valley of Colorado, USA .1. Ligusticum porteri (osha) and Matricaria chamomilla (manzanilla). J Ethnopharmacol. 1985;13:51-55.
  53. Zschocke S, Liu JH, Stuppner H, Bauer R. Comparative study of roots of Angelica sinensis and related umbelliferous drugs by thin layer chromatography, high-performance liquid chromatography, and liquid chromatography mass spectrometry. Phytochem Anal. 1998;9:283-290.
  54. Delgado G, Rezagarduno RG, Rios MY, Delrio F. Phthalides and monoterpenes of the hexane extract of the roots of Ligusticum porteri. Planta Med. 1992;58:570-570.
  55. Delgado G, Rezagarduno RG, Toscano RA, Bye R, Linares E. Secondary metabolites from the roots of Ligusticum porteri (Umbelliferae) - X-ray structure of Z-6.6’,7.3a’-diligustilide. Heterocycles. 1988;27:1305-1312.
  56. Gillespie SG, Duszynski JN. Phthalides and monoterpenes of the hexane extracts of the roots of Ligusticum porteri, L filicinum, and L tenuifolium. Planta Med. 1998;64:392-392.
  57. Cegiela-Carlioz P, Bessiere JM, David B, Mariotte AM, Gibbons S, Dijoux-Franca MG. Modulation of multi-drug resistance (MDR) in Staphylococcus aureus by osha (Ligusticum porteri L., Apiaceae) essential oil compounds. Flavour Fragrance J. 2005;20:671-675.
  58. Cool LG, Vermillion KE, Takeoka GR, Wong RY. Irregular sesquiterpenoids from Ligusticum grayi roots. Phytochemistry. 2010;71:1545-1557.
  59. Sarker SD, Naharl L. Natural medicine: The genus Angelica. Curr Med Chem. 2004;11:1479-1500
  60. Xie J, Lu J, Qian Z, Yu Y, Duan J, Li S. Optimization and comparison of five methods for extraction of coniferyl ferulate from Angelica sinensis. Molecules. 2009;14:555-565.
  61. Panoutsopoulos GI, Beedham C. Enzymatic oxidation of vanillin, isovanillin and protocatechuic aldehyde with freshly prepared guinea pig liver slices. Cellular Physiology and Biochemistry. 2005;15:89-98.
  62. Ou SY, Kwok KC. Ferulic acid: pharmaceutical functions, preparation and applications in foods. J Sci Food Agric. 2004;84:1261-1269.
  63. Bakkali F, Averbeck S, Averbeck D, Waomar M. Biological effects of essential oils - a review. Food and Chemical Toxicology. 2008;46:446-475.
  64. Edris AE. Pharmaceutical and therapeutic potentials of essential oils and their individual volatile constituents: a review. Phytotherapy Research. 2007;21:308-323.
  65. Stella Nerio L, Olivero-Verbel J, Stashenko E. Repellent activity of essential oils: a review. Bioresour Technol. 2010;101:372-378.
  66. Naser B, Bodinet C, Tegtmeier M, Lindequist U. Thuja occidentalis (arbor vitae): a review of its pharmaceutical, pharmacological and clinical properties. Evidence-Based Complementary and Alternative Medicine. 2005;2:69-78.
  67. Rossi P, Bao L, Luciani A, et al. (E)-methylisoeugenol and elemicin: antibacterial components of Daucus carota L. essential oil against Campylobacter jejuni. J Agric Food Chem. 2007;55:7332-7336.
  68. Dentali S, Hoffman J. Potential antiinfective agents from Eriodictyon angustifolium and Salvia apiana. Int J Pharmacogn. 1992;30:223-231.
  69. Turi C. Herbal Medicine and Pulmonary Tuberculosis in England and North America: An Examination of Herbal Materia Medica from the 16th Century Onwards. [MSC]. Canterbury, UK: Department of Anthropology, University of Kent; 2009.
  70. Deciga-Campos M, Gonzalez-Trujano E, Navarrete A, Mata R. Antinociceptive effect of selected mexican traditional medicinal species. Proceedings of the Western Pharmacology Society, Vol 48. 2005;48:70-72.
  71. Brindis F, Rodrı´guez R, Bye R, Gonza´lez-Andrade M, Mata R. (Z)-3-butylidenephthalide from Ligusticum porteri, an alpha-glucosidase inhibitor. Journal of Natural Products. 2010.
  72. Daniels AL, Van Slambrouck S, Lee RK, et al. Effects of extracts from two native american plants on proliferation of human breast and colon cancer cell lines in vitro. Oncol Rep. 2006;15:1327-1331.
  73. Deciga-Campos M, Rivero-Cruz I, Arriaga-Alba M, et al. Acute toxicity and mutagenic activity of Mexican plants used in traditional medicine. J Ethnopharmacol. 2007;110:334-342.
  74. Medicinal Plants Working Group. 1999 Graduate Program in Sustainable Development and Conservation Biology:Review of Four Species for Potential Listing on the Convention on International Trade. Available at: www.nps.gov/plants/medicinal/pubs/cites-a.htm#lig_top,. Accessed July 20, 2010.
  75. United Nations Environment Programme. Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES), Appendices I,II,and III. Available at: www.cites.org/eng/app/appendices.shtml. July 26, 2010.