_Ergota_; _Ergot of Rye_,[2749] _Spurred Rye_; F. _Seigle ergoté_; G.
_Mutterkorn_.
=Botanical Origin=—_Claviceps purpurea_ Tulasne, a fungus of the order
_Pyrenomycetes_, of which ergot is an immature form, it being the
_sclerotium_ (termed in the British Pharmacopœia _compact mycelium_ or
_spawn_) developed within the paleæ of numerous plants of the order
_Gramineæ_.
Ergot is obtained almost exclusively from rye, _Secale cereale_ L.;
but the same fungus is produced on grasses belonging to many other
genera, as _Agropyrum_, _Alopecurus_, _Ammophila_, _Anthoxanthum_,
_Arrhenatherum_, _Avena_, _Brachypodium_, _Calamagrostis_, _Dactylis_,
_Glyceria_, _Hordeum_, _Lolium_, _Poa_, and _Triticum_. Other organisms
of diverse form, but of doubtful specific distinctness, are developed
in _Molinia_, _Oryza_, _Phragmites_, and other grasses. In the order
_Cyperaceæ_ (e.g., _Scirpus_), peculiar ergots are known.
=History=—Although it is hardly possible that so singular a production
as ergot should be unnoticed in the writings of the classical authors,
we believe no undoubted reference to it has been discovered.[2750]
The earliest date under which we find ergot mentioned on account of
its obstetric virtues is towards the middle of the 16th century, by
Adam Lonicer of Frankfort, who describes its appearance in the ears
of rye, and adds that it is regarded by women to be of remarkable
and certain efficacy.[2751] It is also very clearly described in the
writings of Johannes Thalius, who speaks of it as used “_ad sistendum
sanguinem_.”[2752] In the next century it was noticed by Caspar Bauhin,
who termed it _Secale luxurians_,[2753] and by the English botanist
Ray,[2754] with allusion to its medicinal properties.
Rathlaw, a Dutch accoucheur, employed ergot in 1747. Thirty years later
Desgranges of Lyons prescribed it with success; but its peculiar and
important properties were hardly allowed until the commencement of the
present century, when Dr. Stearns of New York succeeded in gaining for
them fuller recognition.[2755] Ergot of rye was not, however, admitted
into the London Pharmacopœia until 1836.[2756]
[2748] Dingler’s _Polytechnisches Journal_, 197 (1870) 177; also
_Chemisches Centralblatt_, 1870. 607.
[2749] From the French _ergot_, anciently _argot_, a cock’s spur.
[2750] Consult Pliny’s _Nat. Hist._ book 18. ch. 44.
[2751] _Kreuterbuch_, ed. 1582. 285 (not in the edition of 1560).
[2752] _Sylva Hercynia_, Francof. 1588. 47.
[2753] _Pinax Theatri Botanici_, Basil. 1623. 23.
[2754] _Hist. Plant._ ii. (1693) 1241.
[2755] Stillé, _Therapeutics and Mat. Med._ ii. (1868) 609.
[2756] From 1825 to 1828 the wholesale price of ergot of rye in London
was from 36_s._ to 50_s._ per lb., that is to say, from twelve to
fifteen times its present value.
The use of flour containing a considerable proportion of ergot, gives
rise to a very formidable disease, distinguished in modern medicine as
_Ergotism_, but known in early times by a variety of names, as _Morbus
spasmodicus_, _convulsivus_, _malignus_, _epidemicus vel cerealis_,
_Raphania_, _Convulsio raphania_[2757] or _Ignis sancti Antonii_.
Some of the malignant epidemics which visited Europe after seasons of
rain and scarcity during the middle ages have been referred with more
or less of probability to ergot-disease.[2758] The chronicles of the
6th and 8th centuries note the occurrence of maladies which may be
suspected as due to ergotized grain. There is less of doubt regarding
the epidemics that prevailed from the 10th century and were frequent
in France, and in the 12th in Spain. In the year 1596 Hessen (Hessia)
and the adjoining regions were ravaged by a frightful pestilence, which
the Medical Faculty of Marburg attributed to the presence of ergot in
the cereals consumed by the population. The same disease appeared in
France in 1630, in Voigtland (Saxony) in the years 1648, 1649, and
1675; again in various parts of France, as Aquitaine and Sologne, in
1650, 1670, and 1674. Freiburg and the neighbouring region were visited
by the same malady in 1702; other parts of Switzerland in 1715-16;
Saxony and Lusatia in 1716; many other districts of Germany in 1717,
1722, 1736, and 1741-2.[2759] The last epidemic in Europe occasioned by
ergot appears to be that which, after the rainy season of 1816, visited
Lorraine and Burgundy, and proved fatal to many people of the poorer
class. Ergot disease is sometimes observed in Abyssinia at the present
day,[2760] and a few cases of it have even been lately recorded in
Bavaria.[2761]
=Formation=—The true nature of ergot has long been the source of a
great diversity of opinion, not set at rest by the admirable researches
of L. R. Tulasne, from whose _Mémoire sur l’Ergot des Glumacées_,[2762]
the following account is for the most part extracted.
[2757] Pereira, _Elem. of Mat. Med._ ii. (1850) 1007.
[2758] Consult Häser, _Lehrbuch der Geschichte der Medicin und der
Volkskrankheiten_, 1845. i. 256. 830, ii. 94; C. F. Heusinger,
_Recherches de Pathologie comparée_, Cassel, i. (1853) 543-554; Mérat
et De Lens, _Dict. Mat. Med._ iii. 131, vii. 268.
[2759] Tissot of Lausanne, _Phil. Trans._ lv. (1766) 106.—See also
Dodart, _Mém. de l’ Acad. R. des Sciences_, x., années 1666-1699
(Paris, 1730) 561; _Hist. de la Soc. Roy. de Méd._, année 1776. 345;
and _Mém. de Méd. et de Phys. méd._ année 1776. 260-311. 417.
[2760] Th. von Heuglin, _Reise nach Abessinien_ etc. Jena, 1868. 180.
[2761] Wiggers and Husemann, _Jahresbericht_ for 1870. 582.
[2762] _Ann. des Sciences nat._, Bot., xx. (1853) 1-56 and 4
plates.—More recent observations will be found in St. Wilson’s paper,
_Trans. of the Bot. Society of Edinburgh_, xli. (1876) 418-434 with
figures; and especially in Luerssen (quoted at p. 735) 156, _et seqq._
The formation of ergot often affects only a few caryopsides in a single
ear; sometimes, however, more than twenty. In the former case, the
healthy development of the other caryopsides is not prevented, but if
too many are attacked, the entire ear decays. The more isolated ergots
generally grow larger, and attain their greatest size on rye which
springs up here and there among other cereals.
The first symptoms of ergot-formation is the so-called _honey-dew of
rye_, a yellowish mucus, having an intensely sweet taste, and the
peculiar disagreeable odour frequently belonging to fungi. Drops
of this mucus show themselves here and there on the ears in the
neighbourhood of diseased grains, and attract ants and beetles of
various kinds, especially the yellowish-red _Rhagonycha melanura_
Fabr., but not bees. On this account the beetle in question has been
supposed to be instrumental in the development of ergot, and it may
possibly be so, but only by transporting the saccharine mucus from one
plant to another.
The honey-dew of rye contains neither oil-drops nor starch. After
dilution with water, it produces a rapid and abundant separation of
cuprous oxide from an alkaline solution of cupric tartrate. Dried over
sulphuric acid, it solidifies into a crystalline mass. After a few days
the drops of honey-dew dry up and disappear from the ear. The grain at
this period becomes completely disintegrated, and devoid of starch.
The ergotized soft ovaries are covered with, and penetrated by a white,
spongy, felted tissue, the _mycelium_ of the young fungus. It is made
up of slender, thread-like cells, the _hyphæ_, the outer layer of
which consists of radially-diverging cells, the _basidia_. The whole
mycelium forms by its crevices and folds a number of cavities opening
externally; from its outer layer, which is also called the _hymenium_
or _spermatophorum_, an immense number of agglutinated, elongated
granules, the _conidia_, are separated. These cells, the products
of the basidia, are not more than four mkm. in length, and give the
floral organs the appearance of being covered with a whitish dust. The
honey-dew likewise contains an abundance of conidia, but it is only
on dilution that they are precipitated and become easily perceptible;
the formation of the honey-dew is intimately connected with that of
the conidia themselves. Ergot in this primary or mycelium stage was
regarded as an independent fungus by Léveillé (1827), who named it
_Sphacelia segetum_. According to Kühn (1863), it may even be directly
by germination of the conidia within the ears of rye.
The mycelium penetrates and envelops the caryopsis, with the exception
of the apex, and thereby prevents its further growth, destroying
especially the epicarp and the embryo. At the base of the caryopsis,
there is formed by tumefaction and gradual transverse separation of
the thread-cells of the mycelium, a more compact kernel-like body (the
future ergot) violet-black without, white within, which gradually
but largely increases in size, and ultimately separates from the
mycelium as the loose tissue of the latter dries and shrinks up after
the completion of its functions. By this growth, the remains of the
caryopsis, still recognizable by their hairs and by the rudiments of
the style, as well as by the surviving portions of the mycelium-tissue,
become visible above the paleæ on the apex of the mature ergot, now
projecting prominently from the ear. Very rarely the ergot is crowned
by a fully developed seed; in the commercial drug, the apex is usually
broken off.
It is evident that in the process of development just described,
the very tissue of the caryopsis of the rye does not undergo a
_transformation_, but that it is _simply destroyed_. Neither in
external form, nor in anatomical structure does ergot exhibit any
resemblance to a caryopsis or a seed, although its development takes
place between the flowering time and that at which the rye begins to
ripen. It has been regarded as a complete fungus, and as such was named
by De Candolle (1816) _Sclerotium Clavus_ and by Fries _Spermædia
Clavus_.
No further change in the ergot occurs while it remains in the ear;
but laid on damp earth, interesting phenomena take place. At certain
points, small orbicular patches of the rind fold themselves back,
and gradually throw out little white heads. These increase in size,
whilst the outer layers of the neighbouring tissue gradually lose their
firmness and become soft and rather granular, at the same time that the
cells, of which they are made up, become empty and extended. In the
interior of the ergot, the cells retain their oil drops unaltered. The
heads assume a greyish-yellow colour, changing to purple, and finally
after some weeks stretch themselves towards the light on slender
shining stalks of a pale violet colour. The stalks often attain an inch
in length, with a thickness of about ½ a line. They consist of thin,
parallel, closely-felted cell-threads, devoid of fat oil. Ergot is
susceptible of this further development only so long as it is fresh,
that is to say, at most until the next flowering time of rye. Within
this period however, even fragments are capable of development. There
are sometimes also produced colourless threads of mould which belong
to other fungi, as _Verticillium cylindrosporum_ Corda, and which
frequently overgrow the _Claviceps_.[2763]
[2763] Ergot of rye collected by myself in August, placed upon earth in
a garden-pot and left in the open air unprotected through the winter,
began to develop the _Claviceps_ on the 20th March, and on another
occasion on the 20th April, at which date some sowed in February also
began to start, Sharp frost appears to retard the vegetation; thus,
after the cold winter of 1869-70, _Claviceps_, even in the greenhouse,
did not make its appearance before the 11th May. The earliest instance
of fully developed _ergots_ which I ever observed, occurred on the
11th of June; more frequently they are seen only in the beginning of
July.—F. A. F.
At the point where the stalk joins the spherical or somewhat flattened
head, the latter is depressed and surrounds the stalk with an annular
border. After a short time there appear on the surface of the head,
which is ⅒ of an inch in diameter, a number of brownish warts, in
which are the openings of minute cavities, the _conceptacula_ or
_perithecia_. On transverse section, they appear arranged radially
round the circumference of the head. In each cavity are a large number
of delicate sacs, only 3-5 mkm. thick, and about 100 mkm. long, the
_thecæ_ or _asci_, each containing, as is usual in fungi, 8 spores.
These are simple thread-shaped cells, filled with a homogeneous solid
mass.
The thicker ends of the spore-sacs (_asci_) open while still within
the perithecium; the spores issue united in a bundle, and are emitted
from the aperture of the perithecium. In consequence of their somewhat
glutinous consistence, they remain united even after their extrusion,
and form white silky flocks; their number in the 20 or 30 heads
sometimes produced from a single ergot, often exceeds a million. The
heads themselves die in two or three weeks after they have begun to
make their appearance. They represent the true fructification of the
fungus. This state of the plant appears to have been first noticed in
1801 by Schumacher, who called it _Sphæria_; it was subsequently known
as _Cordiceps_, _Cordyliceps_, _Kentrosporium_, etc., until Tulasne
proved it to be the final stage of development of ergot.
The three different forms of this structure, namely, the mycelium,
the ergot, and the fruit-bearing heads, are therefore merely
successive states of one and the same biennial fungus, which have
been appropriately united by Tulasne under the name of _Claviceps
purpurea_. The middle stage forms the _sclerotium_, which occurs in a
large number of the most various fungi, and is a special state of rest
of these plants. The direct proof that the mycelium is produced from
spores of the fruit-head sown on ears of rye, was supplied by Kühn in
1863. It has already been mentioned that the same organism is produced
from conidia; whence it appears that a twofold formation of ergot is
possible, as is frequently the case in other fungi.
=Description=—Spurred rye, as found in commerce, consists of fusiform
grains, which it is convenient to term _ergots_. They are from ⅓
to 1½ inch in length, and ½ to 4 lines in diameter; their form is
subcylindrical or obtusely prismatic, tapering towards the ends,
generally arched, with a longitudinal furrow on each side. At the
apex of each ergot, there is often a small whitish easily detached
appendage, while the opposite extremity is somewhat rounded. The
ergots are firm, horny, somewhat elastic, have a close fracture, are
brittle when dry, yet difficult to pulverize. The whitish interior
is frequently laid bare by deep transverse cracks. The tissue is but
imperfectly penetrated by water, even the thinnest sections swelling
but slightly in that fluid.
Ergot of rye has a peculiar offensive odour, and a mawkish, rancid
taste. It is apt to become deteriorated by keeping, especially
when pulverized, partly from oxidation of the oil, and partly from
the attacks of a mite of the genus _Trombidium_. To assist its
preservation, it should be thoroughly dried, and kept in closed bottles.
=Microscopic Structure=—In fully developed ergot, no organs can be
distinguished. It consists of uniform, densely felted tissue of short,
thread-like, somewhat thick-walled cells, which are irregularly packed,
and so intimately matted together that it is only by prolonged boiling
of thin slices with potash, and alternate treatment with acids and
ether, that the individual cells can be made evident. Without such
treatment, the cells even in the thinnest sections, show a somewhat
rounded, nearly isodiametric outline. This pseudo-parenchyme of ergot
exhibits therefore an aspect somewhat different from that of the
loosely felted cells (_hyphæ_) of other fungi. Ergot nevertheless is
not made up of cells differing from those of fungi generally. If thin
longitudinal slices of the innermost tissue are allowed to remain in
a solution of chromic acid containing about 1 per cent., they will
distinctly show the _hyphæ_, which are however considerably shorter
than those of other fungi. They contain numerous drops of fat oil, but
neither starch nor crystals. It is remarkable that this nearly empty
and not much thickened parenchyme should form so compact and solid a
tissue.
The cell-walls of the tissue of ergot are not coloured blue, even after
prolonged treatment with iodine in solution of potassium iodide; or
when the tissue has been previously treated with sulphuric acid, or
kept for days in contact with potash and absolute alcohol at 100° C. In
this respect the cellulose of fungi differs from that of phanerogamic
plants.
Of the outermost rows of cells in ergot, a few only are of a violet
colour, but they are not otherwise distinguishable from the colourless
tissue,—or at most by the somewhat greater thickness of their walls.
=Chemical Composition=—The composition of ergot has been elaborately
investigated by Wiggers as early as 1830. The drug contains about 30
per cent. of a non-drying, yellowish oil, chiefly consisting of olein,
palmitin, and small proportions of volatile fatty acids, especially
acetic and butyric, combined with glycerin. The large amount of oil
is remarkable; the fungi, dried at 100°, usually contain not more
than 5 per cent. of fat, mostly much less; they are on the other hand
much richer in albumin than ergot of rye. The oil of the latter, as
extracted by bisulphide of carbon, is accompanied by small quantities
of _resin_ and _cholesterin_ (see p. 420). It is erroneous to attribute
to this oil the poisonous properties of ergot, although it has been
shown by Ganser[2764] to display irritating properties when taken in
doses of about 6 grammes. But the effects observed appear dependent on
the presence in it of resin.
According to Wenzell (1864), ergot of rye contains two peculiar
alkaloids, which he designated _Ecboline_ and _Ergotine_,[2765] and
claimed to be the active principles of the drug. They were, however,
got merely as brownish amorphous substances.
The two bases of ergot are, according to Wenzell, combined with
_Ergotic Acid_, the existence of which has been further admitted by
Ganser. It is said to be a volatile body yielding crystallizable salts.
A crystallized colourless alkaloid, _Ergotinine_, C₃₅H₄₀N₄O₆, has been
isolated (1877-1878) by Tanret, a pharmacien of Troyes. He obtained
it to the amount of about 0·04 per cent., some amorphous ergotinine
moreover being present. Tanret exhausts the powdered drug with boiling
alcohol, which by evaporation affords a fluid resin and an aqueous
solution, besides a fatty layer. Some ergotinine is removed from the
resin by shaking it with ether, and mixed with the main liquid. This
is acidulated and purified by means of ether. Lastly, the ergotinine
is extracted by adding a slight excess of carbonate of potassium and
shaking with ether, and recrystallizing from alcohol. The solutions
of ergotinine turn very soon greenish and red; they are fluorescent.
Sulphuric acid imparts to it a red, violet, and finally blue hue.
Dragendorff and several of his pupils, since 1875, have isolated
the following _amorphous_ principles of the drug under notice:—(1)
_Sclerotic acid_ (doubtful formula C₁₂H₁₉}NO₉), said to be a very
active substance, chiefly in subcutaneous injections. About 4 per
cent. of colourless acid may be obtained from good ergot of rye. (2)
_Scleromucin_, a mucilaginous matter, which may be precipitated by
alcohol from aqueous extracts of the drug. Scleromucin when dried is
no longer soluble in water. (3) _Sclererythrin_, the red colouring
matter, probably allied to anthrachinon and the colouring substances of
madder, chiefly to purpurin. (4) _Sclerojodin_, a bluish-black powder,
soluble in alkalis. (5) _Fuscosclerotinic acid._ (6) _Picrosclerotine_,
apparently a highly poisonous alkaloid. Lastly (7) _Scleroxanthin_,
C₇H₇O₃ + OH₂; and (8) _Sclerocrystallin_, C₇H₇O₃, have been obtained in
crystals; their alcoholic solution is but little coloured, yet assumes
a violet hue on addition of ferric chloride.
Tanret also observed in ergot of rye a volatile _camphoraceous
substance_.
Ergot, in common with other fungi,[2766] contains a sugar termed
_Mycose_, closely allied to cane-sugar, and probably identical with
_Trehalose_ (see p. 417). Mycose crystallizes in rhombic octohedra,
having the composition C₁₂H₂₂O₁₁ + 2H₂O. Mitscherlich obtained of it
about one-tenth per cent. It appears that the sugar exuded in the first
stage of growth of the fungus,—the so-called _rye honey-dew_,—is in
its principal characters different from mycose. Instead of the latter,
Mitscherlich, as well as Fiedler and Ludwig, sometimes obtained from
ergot _Mannite_.
[2764] _Archiv der Pharm._ cxliv. (1870) 200.
[2765] The name _Ergotine_ has also been given to a medicinal extract
of ergot, prepared after a method devised by Bonjean, a pharmacien of
Chambéry, vide _Journ. de Pharm._ iv. (1843) 107; Pereira, _Elem. of
Mat. Med._ ii. (1850) 1012.
[2766] See Müntz in _Comptes Rendus_, lxxvi. (1873) 649.
Schoonbroodt also found in ergot _Lactic Acid_. Several other chemists
have further proved the presence of acetic and formic acids.
Starch is entirely wanting in ergot at all times. The drug yields about
3 per cent. of nitrogen, corresponding probably to a large amount of
albuminoid matter. Ganser, however, obtained only 3·2 per cent. of
albumin _soluble in water_.
When ergot or its alcoholic extract is treated with an alkali
it yields, as products of the decomposition of the albuminoid
matters, ammonia or ammonia-bases, according to Ludwig and Stahl,
_Methylamine_,—according to others, _Trimethylamine_. Manassewitz, as
well as Wenzell, state that phosphate of trimethylamine is present
in an aqueous extract of ergot, but Ganser ascertained that no such
base _pre-exists_ in ergot. We have found that the crystals which
abound in the extract, after it has been kept for some time, are an
acid phosphate of sodium and ammonium with a small proportion of
sulphate.[2767]
=Production and Commerce=—Ergot of rye is to be met with in all the
countries producing cereals; we have seen it in the high valleys of the
Alps, and Schübeler states that it grows in Norway, as far north as 60°
N. lat.
The drug is chiefly imported into London from Vigo in Spain and from
Tenerife; it is also shipped from Hamburg and France. Dr. de Lanessan,
writing to one of us from Vigo in 1872, remarks that vast quantities of
rye are grown in Galicia, and that owing to the humidity of the climate
the grain is extensively ergotized,—in fact the parasite is present
in one ear out of every three. At the time of harvest the ergots are
picked out, and the rye is thus rendered fit for food.
Southern and Central Russia furnish considerable supplies of the drug.
In the central parts of Europe, ergot does not everywhere occur in
sufficient abundance to be collected, and it greatly diminishes as the
state of agriculture improves. We have noticed that ergot from Odessa
was of a slaty hue and in much smaller grains than that from Spain.
=Uses=—Ergot is principally used on account of its specific action on
the uterus in parturition.
=Other Varieties of Ergot=—_Ergot of Wheat_ (Triticum vulgare), which
is in shorter and thicker ergots than that of rye, is picked out by
hand in some parts of Italy and France, from grain intended to be used
for the manufacture of vermicelli and other pastes; and such ergot is
sold to druggists. Carbonneaux Le Perdriel[2768] has endeavoured to
show that it is less prone to become deteriorated by age than that
of rye, and that it never produces the deleterious effects sometimes
occasioned by the latter.
[2767] The red colour of an alcoholic solution may serve for the
detection of small quantities of ergot in flour. The reaction with
potash, and evolution of the characteristic odour of herring brine may
assist in the same object. Extraction of the fatty oil with carbon
bisulphide may also be recommended as a test, inasmuch as good cereal
grains contain but a very small percentage of fat.
[2768] _De l’Ergot de Froment et de ses propriétés méd._ (thèse)
Montpellier, 1862.
The same writer asserts that _Ergot of Oat_ is sometimes collected and
sold either _per se_, or mixed with that of rye. It differs from the
latter in the ergots being considerably more slender.
Ergot of the North African grass _Arundo Ampelodesmos_ Cirillo, known
as _Diss_, has been collected for use, and according to Lallemant[2769]
is twice as active as that of rye. It is from 1 to 3 inches long by
only about ⅒ of an inch broad, generally arched, or in the large
ergots twisted spirally. We find it to share the structural character
of the ergot of rye; it is in all probability the same formation, yet
remarkably modified.