Ergot

This article is about the fungi. For the part of a horse's hoof, see Ergot (horse anatomy).

Ergot
Claviceps purpurea
Scientific classification
Kingdom:	Fungi
Division:	Ascomycota
Class:	Sordariomycetes
Order:	Hypocreales
Family:	Clavicipitaceae
Genus:	Claviceps
Species
About 50, including: Claviceps africana Claviceps fusiformis Claviceps paspali Claviceps purpurea Claviceps sorghi Claviceps zizaniae

Ergot or ergot fungi refers to a group of fungi of the genus Claviceps.^[1] The most prominent member of this group is Claviceps purpurea. This fungus grows on rye and related plants, and produces alkaloids that can cause ergotism in humans and other mammals who consume grains contaminated with its fruiting structure (called ergot sclerotium).^[2][3] Claviceps includes about 50 known species, mostly in the tropical regions. Economically significant species include C. purpurea (parasitic on grasses and cereals), C. fusiformis (on pearl millet, buffel grass), C. paspali (on dallis grass), and C. africana^[4] (on sorghum). C. purpurea most commonly affects outcrossing species such as rye (its most common host), as well as triticale, wheat and barley. It affects oats only rarely.

C. purpurea has at least three races or varieties, which differ in their host specificity:^[5]

• G1 — land grasses of open meadows and fields;
• G2 — grasses from moist, forest, and mountain habitats;
• G3 (C. purpurea var. spartinae) — salt marsh grasses (Spartina, Distichlis).

Life cycle

An ergot kernel called a sclerotium develops when a spore of fungal species of the genus Claviceps infects a floret of flowering grass or cereal. The infection process mimics a pollen grain growing into an ovary during fertilization. Infection requires that the fungal spore have access to the stigma, so plants infected by Claviceps are mainly outcrossing species with open flowers, such as rye (Secale cereale) and ryegrasses (genus Lolium). The proliferating fungal mycelium then destroys the plant ovary and connects with the vascular bundle originally intended for seed nutrition. The first stage of ergot infection manifests itself as a white soft tissue (known as sphacelia) producing sugary honeydew, which often drops out of the infected grass florets. This honeydew contains millions of asexual spores (conidia), which insects disperse to other florets. Later, the sphacelia convert into a hard dry sclerotium inside the husk of the floret. At this stage, alkaloids and lipids accumulate in the sclerotium.

Claviceps species from tropic and subtropic regions produce macro- and microconidia in their honeydew. Macroconidia differ in shape and size between the species, whereas microconidia are rather uniform, oval to globose (5x3μm). Macroconidia are able to produce secondary conidia. A germ tube emerges from a macroconidium through the surface of a honeydew drop and a secondary conidium of an oval to pearlike shape is formed, to which the contents of the original macroconidium migrates. Secondary conidia form a white, frost-like surface on honeydew drops and spread via the wind. No such process occurs in Claviceps purpurea, Claviceps grohii, Claviceps nigricans, and Claviceps zizaniae, all from northern temperate regions.

When a mature sclerotium drops to the ground, the fungus remains dormant until proper conditions trigger its fruiting phase (such as the onset of spring or a rain period). It germinates, forming one or several fruiting bodies with heads and stipe, variously coloured (resembling a tiny mushroom). In the head, threadlike sexual spores form, which are ejected simultaneously when suitable grass hosts are flowering. Ergot infection causes a reduction in the yield and quality of grain and hay, and if livestock eat infected grain or hay it may cause a disease called ergotism. Black and protruding sclerotia of C. purpurea are well known. However, many tropical ergots have brown or greyish sclerotia, mimicking the shape of the host seed. For this reason, the infection is often overlooked.

Insects, including flies and moths, carry conidia of Claviceps species, but it is unknown whether insects play a role in spreading the fungus from infected to healthy plants.^[6]

Effects on humans and other mammals

Ergot-derived drug to stop postnatal bleeding

Main article: Ergotism

The ergot sclerotium contains high concentrations (up to 2% of dry mass) of the alkaloid ergotamine, a complex molecule consisting of a tripeptide-derived cyclol-lactam ring connected via amide linkage to a lysergic acid (ergoline) moiety, and other alkaloids of the ergoline group that are biosynthesized by the fungus.^[7] Ergot alkaloids have a wide range of biological activities including effects on circulation and neurotransmission.^[8]

Ergotism is the name for sometimes severe pathological syndromes affecting humans or animals that have ingested plant material containing ergot alkaloid, such as ergot-contaminated grains. The Hospital Brothers of St. Anthony (monks) specialized in treating ergotism victims^[9] with balms containing tranquilizing and circulation-stimulating plant extracts. The common name for ergotism is "St. Anthony's Fire",^[9] in reference to monks who cared for victims as well as symptoms, such as severe burning sensations in the limbs.^[10] These are caused by effects of ergot alkaloids on the vascular system due to vasoconstriction of blood vessels, sometimes leading to gangrene and loss of limbs due to severely restricted blood circulation.

The neurotropic activities of the ergot alkaloids may also cause hallucinations and attendant irrational behaviour, convulsions, and even death.^[7][8] Other symptoms include strong uterine contractions, nausea, seizures, and unconsciousness. Since the Middle Ages, controlled doses of ergot were used to induce abortions and to stop maternal bleeding after childbirth.^[11] Ergot extract has been used in pharmaceutical preparations, including Ergot alkaloids in products such as Cafergot (containing caffeine and ergotamine^[11] or ergoline) to treat migraine headaches, and ergometrine, used to induce uterine contractions and to control bleeding after childbirth.^[12] In addition to ergot alkaloids, Claviceps paspali also produces tremorgens (paspalitrem) causing "paspalum staggers" in cattle.^[13] The fungi of the genera Penicillium and Aspergillus also produce ergot alkaloids, notably some isolates of the human pathogen Aspergillus fumigatus,^[14] and have been isolated from plants in the family Convolvulaceae, of which morning glory is best known.

Ergot contains no lysergic acid diethylamide (LSD) but instead contains ergotamine, which is used to synthesize lysergic acid, an analog of and precursor for synthesis of LSD. Moreover, ergot sclerotia naturally contain some amounts of lysergic acid.^[15]

The January 4, 2007 issue of the New England Journal of Medicine includes a paper that documents a British study of more than 11,000 Parkinson's Disease patients. The study found that two ergot-derived drugs, Pergolide and Cabergoline, commonly used to treat Parkinson's Disease may increase the risk of leaky heart valves by up to 700%.^[16]

History

Ergot on wheat spikes

Human poisoning due to the consumption of rye bread made from ergot-infected grain was common in Europe in the Middle Ages. The epidemic was known as Saint Anthony's fire,^[9] or ignis sacer, and some historical events, such as the Great Fear in France during the Revolution have been linked to ergot poisoning.^[17] Linnda R. Caporael posited in 1976 that the hysterical symptoms of young women that had spurred the Salem witch trials had been the result of consuming ergot-tainted rye.^[18] However, Nicholas P. Spanos and Jack Gottlieb, after a review of the historical and medical evidence, later disputed her conclusions.^[19] Other authors have likewise cast doubt on ergotism as the cause of the Salem witch trials.^[20]

American author John Grigsby contends that the presence of ergot in the stomachs of some of the so called 'bog-bodies' (Iron Age human remains from peat bogs N E Europe such as Tollund Man) is indicative of use of ergot in ritual drinks in a prehistoric fertility cult akin to the Eleusinian Mysteries cult of ancient Greece. In his book Beowulf and Grendel, he argues that the Anglo-Saxon poem Beowulf is based on a memory of the quelling of this fertility cult by followers of Odin. He writes that Beowulf, which he translates as barley-wolf, suggests a connection to ergot which in German was known as the 'tooth of the wolf'.^[21]

Kykeon, the beverage consumed by participants in the ancient Greek cult of Eleusinian Mysteries, might have been based on hallucinogens from ergot,^[22] and lysergic acid diethylamide (LSD) is a potent hallucinogen, which was first synthesized from ergot alkaloids by the Swiss chemist, Albert Hofmann, in 1938.

Claviceps purpurea

Main article: Claviceps purpurea

Mankind has known about Claviceps purpurea for a long time, and its appearance has been linked to extremely cold winters that were followed by rainy summers.^{[citation needed]}

The sclerotial stage of C. purpurea conspicuous on the heads of ryes and other such grains is known as ergot. Favorable temperatures for growth are in the range of 18-30 °C. Temperatures above 37 °C cause rapid germination of conidia.^{[citation needed]} Sunlight has a chromogenic effect on the mycelium, with intense coloration.^{[citation needed]} Cereal mashes and sprouted rye are suitable substrates for growth of the fungus in the laboratory.^{[citation needed]}

Claviceps africana

Claviceps africana infects sorghum and was first observed in south Texas in 1997.^{[citation needed]} It only infects unfertilized ovaries, so self-pollination and fertilization can decrease the presence of the disease, but male-sterile lines are extremely vulnerable to infection.^{[citation needed]} Symptoms of infection by C. africana include the secretion of honeydew (a fluid with high concentrates of sugar and conidia), which attracts insects like flies, beetles, and wasps that feed on it. This in turn contributes to the spread of the fungus to uninfected plants.

C. africana caused ergot disease that caused a famine in 1903-1906 in Northern Cameroon, West Africa, and also occurs in eastern and southern Africa, especially Zimbabwe and South Africa. Male sterile sorghums (also referred to as A-lines) are especially susceptible to infection, as first recognized in the 1960s, and massive losses in seed yield have been noted. Infection is associated with cold night temperatures that are below 12 °C occurring two to three weeks before flowering.^{[citation needed]}

See also

Medicinal mushrooms

References

1. ^ Schardl CL, Panaccione DG, Tudzynski P (2006). "Ergot alkaloids – biology and molecular biology". The Alkaloids: Chemistry and Biology. The Alkaloids: Chemistry and Biology 63: 45–86. doi:10.1016/S1099-4831(06)63002-2. ISBN 9780124695634. PMID 17133714. 
2. ^ ergot, online medical dictionary
3. ^ ergot, Dorland's Medical Dictionary
4. ^ Bandyopadhyay, Ranajit; Frederickson, Debra E.; McLaren, Neal W.; Odvody, Gary N.; Ryley, Malcolm J. (1998). "Ergot: A New Disease Threat to Sorghum in the Americas and Australia". Plant Disease 82 (4): 356. doi:10.1094/PDIS.1998.82.4.356. http://www.apsnet.org/online/feature/ergot/0218-01F.pdf 
5. ^ Sylvie Pazoutova. "Intraspecific variability of C. purpurea". http://www2.biomed.cas.cz/~pazouto/purpurea.htm. 
6. ^ Butler, M.D., Alderman, S. C., Hammond, P.C., Berry, R. E. (2001). "Association of Insects and Ergot (Claviceps purpurea) in Kentucky Bluegrass Seed Production Fields". J. Econ. Entomol. 94 (6): 1471–1476. doi:10.1603/0022-0493-94.6.1471. PMID 11777051. http://www.bioone.org/doi/full/10.1603/0022-0493(2001)094%5B1471%3AAOIAEC%5D2.0.CO%3B2. 
7. ^ ^{a b} Tudzynski P, Correia T, Keller U (2001). "Biotechnology and genetics of ergot alkaloids". Appl Microbiol Biotechnol. 57 (5–6): 4593–4605. doi:10.1007/s002530100801. PMID 11778866. 
8. ^ ^{a b} Eadie MJ (2003). "Convulsive ergotism: epidemics of the serotonin syndrome?". Lancet Neurol. 2 (7): 429–434. doi:10.1016/S1474-4422(03)00439-3. PMID 12849122. 
9. ^ ^{a b c} Microbiology in Action. P115. By J. Heritage, Emlyn Glyn Vaughn Evans, R. A. Killington. Cambridge University Press, 1999.
10. ^ http://www.medicinenet.com/script/main/art.asp?articlekey=14891
11. ^ ^{a b} Untersuchungen über das Verhalten der Secalealkaloide bei der Herstellung von Mutterkornextrakten. Labib Farid Nuar. Universität Wien - 1946 - (University of Vienna)
12. ^ Black, Michael H.; Halmer, Peter (2006). The encyclopedia of seeds: science, technology and uses. Wallingford, UK: CABI. pp. 226. ISBN 978-0-85199-723-0. 
13. ^ Cole RJ, Dorner JW, Lansden JA, Cox RH, Pape C, Cunfer B, Nicholson SS, Bedell DM (1977). "Paspalum staggers: isolation and identification of tremorgenic metabolites from sclerotia of Claviceps paspali". Journal of Agricultural and Food Chemistry 25: 1197-1201. doi:10.1021/jf60213a061. 
14. ^ Rao KK, Rao S (1975). "Effect of tweens on the production of ergot alkaloids by Aspergillus fumigatus". Folia Microbiol. 20 (5): 418–422. doi:10.1007/BF02877045. PMID 1104424. 
15. ^ Correia T, Grammel N, Ortel I, Keller U, Tudzynski P. (2001). "Molecular cloning and analysis of the ergopeptine assembly system in the ergot fungus Claviceps purpurea". Chem Biol. 10 (12): 1281–1292. doi:10.1016/j.chembiol.2003.11.013. PMID 14700635. 
16. ^ NEJM - Dopamine Agonists and the Risk of Cardiac-Valve Regurgitation
17. ^ Matossian, Mary Kilbourne, Poisons of the Past: Molds, Epidemics, and History. New Haven: Yale, 1989 (reedited in 1991) ISBN 0-300-05121-2
18. ^ Caporael LR (April 1976). "Ergotism: the satan loosed in Salem?". Science 192 (4234): 21–6. doi:10.1126/science.769159. PMID 769159. http://web.utk.edu/~kstclair/221/ergotism.html. 
19. ^ Spanos NP, Gottlieb J (December 1976). "Ergotism and the Salem Village witch trials". Science 194 (4272): 1390–4. doi:10.1126/science.795029. PMID 795029. 
20. ^ Woolf A (2000). "Witchcraft or mycotoxin? The Salem witch trials". J Toxicol Clin Toxicol 38 (4): 457–460. doi:10.1081/CLT-100100958. PMID 10930065. 
21. ^ Grigsby, John (2005). Beowulf & Grendel: The Truth Behind England's Oldest Legend. Watkins Publishing. ISBN 1842931539. 
22. ^ "Mixing the Kykeon", ELEUSIS: Journal of Psychoactive Plants and Compounds, New Series 4, 2000

External links

• Claviceps purpurea - Ergot Alkaloid
• Ergot article from North Dakota State University, 2002
• Abundant Respirable Ergot Alkaloids from the Common Airborne Fungus Aspergillus fumigatus
• PBS Secrets of the Dead: "The Witches Curse" (concerning the Salem trials and ergot)
• New England Journal of Medicine - Dopamine Agonists and the Risk of Cardiac-Valve Regurgitation
• Parkinson's Drugs Can Damage Heart Valves. HealthDay. January 3, 2007.
• The Reactions of Claviceps purpurea to Variations of Environment
• A Preliminary Study of Claviceps purpurea in Culture
• Observations on the Poisonous Plants of Michigan
• Viability of Claviceps africana Spores Ingested by Adult Corn Earworm Moths, Helicoverpa zea
• Genetic diversity of Claviceps africana on sorghum and Hyparrhenia
• Windborne spread of ergot disease (Claviceps africana) in sorghum A-lines in Zimbabwe