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Ergot
Claviceps purpurea - Köhler–s Medizinal-Pflanzen-185.jpg
Claviceps purpurea
Scientific classification e
Kingdom: Fungi
Division: Ascomycota
Class: Sordariomycetes
Order: Hypocreales
Family: Clavicipitaceae
Genus: Claviceps
Tul., 1853
Species

About 50, including:
Claviceps africana
Claviceps fusiformis
Claviceps paspali
Claviceps purpurea
Claviceps sorghi
Claviceps zizaniae
Claviceps lutea

Ergot (pron. UR-gət) or ergot fungi refers to a group of fungi of the genus Claviceps.

The most prominent member of this group is Claviceps purpurea ("rye ergot fungus"). 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).

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), C. africana (on sorghum), and C. lutea (on paspalum). 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:

  • 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; consequently, 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 (5×3μm). 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 (such as the onset of spring or a rain period) trigger its fruiting phase. It germinates, forming one or several fruiting bodies with heads and stipes, 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.

Evolution

The evolution of plant parasitism in the Clavicipitaceae dates back at least 100 million years, to the early-mid Cretaceous. An amber fossil discovered in 2014 preserves a grass spikelet and an ergot-like parasitic fungus. The fossil shows that the original hosts of the Clavicipitaceae could have been grasses. The discovery also establishes a minimum time for the conceivable presence of psychotropic compounds in fungi. Several evolutionary processes have acted to diversify the array of ergot alkaloids produced by fungi; these differences in enzyme activities are evident at the levels of substrate specificity (LpsA), product specification (EasA, CloA) or both (EasG and possibly CloA). The "old yellow enzyme", EasA, presents an outstanding example. This enzyme catalyzes reduction of the C8=C9 double-bond in chanoclavine I, but EasA isoforms differ in whether they subsequently catalyze reoxidation of C8–C9 after rotation. This difference distinguishes most Clavicipitaceae from Trichocomaceae, but in Clavicipitaceae it is also the key difference dividing the branch of classical ergot alkaloids from dihydroergot alkaloids, the latter often being preferred for pharmaceuticals due to their relatively few side effects.

Effects on humans and other mammals

Ergotism is the name for sometimes severe pathological syndromes affecting humans or other animals that have ingested plant material containing ergot, such as ergot-contaminated grains. The Hospital Brothers of St. Anthony, an order of monks established in 1095, specialized in treating ergotism victims with balms containing tranquilizing and circulation-stimulating plant extracts. The common name for ergotism is "St. Anthony's Fire", in reference to this order of monks and the severe burning sensations in the limbs which was one of the symptoms. There are two types of ergotism, the first is characterized by muscle spasms, fever and hallucinations and the victims may appear dazed, be unable to speak, become manic, or have other forms of paralysis or tremors, and suffer from hallucinations and other distorted perceptions. The second type of ergotism is marked by violent burning, absent pulses and shooting pain of the poorly vascularized distal organs, such as the fingers and toes, and are caused by effects of ergot alkaloids on the vascular system due to vasoconstriction, sometimes leading to gangrene and loss of limbs due to severely restricted blood circulation.

History

Ergot01
Ergot on wheat heads

Ergotism is the earliest recorded example of mycotoxicosis, or poisoning caused by toxic molds. Early references to ergot poisoning (ergotism) date back as far as 600 BC, an Assyrian tablet referred to it as a 'noxious pustule in the ear of grain'. In 350 BC, the Parsees described 'noxious grasses that cause pregnant women to drop the womb and die in childbed'. In ancient Syria, ergot was called 'Daughter of Blood'. Radulf Glaber described an ailment he called 'hidden fire' or ignus ocultus, in which a burning of the limb is followed by its separation from the body, often consuming the victim in one night. In 1588, Johannes Thallius wrote that it is called 'Mother of Rye', or rockenmutter, and is used to halt bleeding.

Human poisoning due to the consumption of rye bread made from ergot-infected grain was common in Europe in the Middle Ages. The first mention of a plague of gangrenous ergotism in Europe comes from Germany in 857, following this France and Scandinavia experienced similar outbreaks; England is noticeably absent from the historical regions affected by ergotism as their main source of food was wheat, which is resistant to ergot fungi. In 944, a massive outbreak of ergotism caused 40,000 deaths in the regions of Aquitaine, Limousin, Perigord, and Angoumois in France. In Hesse in 1596, Wendelin Thelius was one of the first to attribute ergotism poisoning to grain. In 1778, S. Tessier, observing a huge epidemic in Sologne, France in which more than 8,000 people died, recommended drainage of fields, compulsory cleaning of grain, and the substitution of potatoes for affected grain.

Saint Anthony's fire and the Antonites

In 1722, the Russian Tsar Peter the Great was thwarted in his campaign against the Ottoman Empire as his army, traveling down the Terek steppe, were struck by ergotism and were forced to retreat in order to find edible grains. A diary entry from the time describes that as soon as people ate the poisoned bread they became dizzy, with such strong nerve contractions that those who did not die on the first day found their hands and feet falling off, akin to frostbite. The epidemic was known as Saint Anthony's fire, or ignis sacer, and some historical events, such as the Great Fear in France during the French Revolution have been linked to ergot poisoning. Saint Anthony was a 3rd Century Egyptian ascetic who lived by the Red Sea and was known for long fasting in which he confronted terrible visions and temptations sent from the Devil. He was credited by two noblemen for assisting them in recovery from the disease; they subsequently founded the Order of St. Anthony in honor of him. Anthony was a popular subject for art in the Middle Ages and his symbol is a large blue "T" sewn onto the shoulder of the order's monks, symbolizing the crutch used by the ill and injured.

The Order of St. Anthony, who were also known as Antonites, grew quickly and hospitals spread through France, Germany, and Scandinavia and gained wealth and power as grateful patrons bestowed money and charitable goods to the hospitals. By the end of the Middle Ages, there were 396 settlements and 372 hospitals owned by the order and pilgrimages to such hospitals became popular as well as the donation of limbs lost to ergotism, which were displayed near shrines to the saint. These hagiotherapeutic centers were the first specialized European medical welfare systems and the friars of the order were knowledgeable about treatment of ergotism and the horrifying effects of the poison. The sufferers would receive ergot-free meals, wines containing vasodilating and analgesic herbs, and applications of Antonites-balsalm, which was the first transdermal therapeutic system (TTS) in medical history. Their medical recipes have been lost to time, though some recorded treatments still remain. After 1130 AD, the monks were no longer permitted to perform operations, and so barber surgeons were employed to remove gangrenous limbs and treat open sores. Three barbers founded a hospital in Memmingen in 1214 and accepted those who were afflicted with the gangrenous form of ergotism. Patients were fed and housed with the more able-bodied individuals acting as orderlies and assistants. Patients with the convulsive form of ergotism, or ergotismus convulsivus, were welcomed for only nine days before they were asked to leave as convulsive ergotism was seen as less detrimental. Though the sufferers often experienced irreversible effects, they most often returned to their families and resumed their livelihoods.

An important aspect to the Order of St. Anthony's treatment practices was the exclusion of rye bread and other ergot-containing edibles, which halted the progression of ergotism. There was no known cure for ergotism itself, however there was treatment of the symptoms, which often included blood constriction, nervous disorder, and/or hallucinations; if the sufferer survived the initial poisoning, his limbs would often fall off and he or she would continue to improve in health if they halted consumption of ergot. The trunk of the body remained relatively untouched by the disease until its final stages and the victims, not understanding the cause of their ailment, would continue to imbibe ergot-laden food for weeks until the condition reached their digestive system. It is believed that the peasantry and children were most susceptible to ergotism, though the wealthy were afflicted as well, as at times entire villages relied on tainted crops for sustenance and during times of famine, ergotism reached into every house. Ergot fungus is impervious to heat and water, thus it was most often baked into bread through rye flour; though other grasses can be infected, it was uncommon in Medieval Europe to consume grasses other than rye. The physiological effects of ergot depended upon the concentration and combinations of the ingested ergot metabolites, as well as the age and nutritional status of the afflicted individual. The Antonites began to decline after physicians discovered the genesis of ergotism and recommended methods for removing the sclerotium from the rye crops. In 1776, the cloisters of the Antonites were incorporated into the Maltese Knights Hospitaller, losing much of their medical histories in the process and losing the ergotism cures and recipes due to lack of use and lack of preservation.

See also

Kids robot.svg In Spanish: Claviceps para niños

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