ON THE PROPERTIES OF SEVERAL SPECIES OF

THE POLYPORACEAE AND OF A NEW VARIETY

OF CLITOCYBE, CLITOCYBE DEALBATA SUDORI-
FICA, PECK

WILLIAM W. FORD AND JOSEPH L. SHERRICK

From the Laboratory of Hygiene and Bacteriology, Johns Hopkins University

Received for publication, June 16, 1911

The properties of several species of boleti including Boletus

clintonianus PECK, Boletus cavipes KALCHBRENNER, Boletus pal-
uster PECK and Boletus chrysenteron FRIES, variety sphagnorum
PECK, have been described previously by one of us (1) and the
importance of a more extended examination of the forms included
in this group indicated. Several of the tube-bearing fungi are
known to possess great dietetic value, such species as Boletus
edulis, BULLIARD, the Steinpilz of the Germans, Boletus scaber
FRIES, and Boletus granulatus LINNAEUS being regarded by mycol-
ogists as important esculents. Other species such as Boletus
satanus LENZ and Boletus luridus SCHAEFFER have long been
looked upon as poisonous. Little is known of the toxic principles
present in either of the latter forms, except for the fact that
Kober (2) has found muscarine in Boletus luridus. The Poly-
pores in general are large fungi containing an abundance of firm
fleshy material and would be thoroughly suitable for the table
were it not for the fact that certain species are apt to be infested
with worms while others have a bitter taste. At the same time
the tube-bearing fungi grow very commonly in America and in
the majority of instances are recognised without great difficulty
in view of the vast amount of study devoted to them by such
authors as Peck (3), Atkinson (4) and the members of the Boston
Mycological Club. The number of definitely poisonous species
of boleti is not large, and since even the most toxic have such
549
550 WILLIAM W. FORD AND JOSEPH L. SHERRICK

properties as a bad taste, a violent emetic or purgative action

associated with or inherent in the poisonous principles which they
contain, the plants are either not eaten in sufficient quantity to
cause poisoning or the active vomiting and purging which their
ingestion induces prevents serious intoxication from them. In-
deed the number of deaths which can be traced to the consump-
tion of the Polypores is very small, although transient illness
from their use is not so very uncommon.
Recently, through the kindness of Miss Jennie F. Conant, we
have received several specimens of tube-bearing fungi from the
1910 collections of the Boston Mycological Club and take great
pleasure in expressing our indebtedness to her and to the other
members of the Club for the opportunity of studying this group.
We have also received from Dr. C. H. Peck of Albany, N. Y., a
number of plants representing a new variety of Clitocybe dealbata.
This form when eaten by man provoked marked symptoms of
poisoning and when tested upon animals has been found to possess
most interesting and important properties.
The various fungi have all been examined by the methods
already described, being studied for the presence of haemolysins,
agglutinins, and poisons. The plant extracts have been examined
chiefly with reference to their action on rabbits’ blood corpuscles,
while their toxicity has been determined by the subcutaneous
injection of suitable amounts of the extract heated to 65#{176}
C. half an hour into both rabbits and guinea-pigs. The following
species of Polypores have been investigated and their dietetic
properties established as far as possible by reference to the exist-
ing literature of the subject.

BOLETUS FELLEUS Bulliard

This species was free from haemolysins or agglutinins. It

contained no muscarine but produced in both guinea-pigs and
rabbits a chronic intoxication from which the animals died in from
two to three weeks. Thus a guinea-pig weighing 460 grams died
in four days from 5 cc. of an extract made from 5 grams of the
dried plant in 50 cc. H2O. At autopsy a tumor mass was found in
 PROPERTIES OF SEVERAL SPECIES OF POLYPORACEAE 551

the abdominal cavity and another guinea-pig weighing 530

grams was therefore given 5 cc. of a similar extract. This animal
perished in sixteen days of a progressive cachexia. A rabbit
weighing 1315 grams succumbed in twenty days to 5 cc. of the
extract while a second rabbit weighing 1270 grams died from the
same dosage after the lapse of thirty-one days. In both instances
the animals showed a steady emaciation, their weight reaching
but a little over 800 grams at death. There were no especial
lesions at autopsy. Boletus felleus is thus definitely poisonous to
both rabbits and guinea-pigs. Because of its intense and lasting
bitter taste it has long been avoided by mycologists. It should
probably be classed as a poisonous boletus.

BOLETUS MINIATO-OLIVACEUS Frost

#{149}An
extract of this plant agglutinated rabbits’ corpuscles in a
dilution of one-tenth of the juice coming from 5 grams of the fun-
gus in 50 cc. 1120. This agglutinin was not destroyed by boiling
half an hour. The extract was not haemolytic. It was poison-
ous to guinea-pigs one animal weighing 95 grams dying in ten
days from 5 cc. and another weighing 420 grams succumbing in
sixteen days to a dose of the same character. No evidence of
the presence of muscarine in the plant was presented by these
animals. Rabbits were not affected by the plant juice. The
guinea-pigs developed a progressive emaciation but nothing
characteristic could be determined at autopsy. According to
Mcllvaine (5) this fungus is an edible species but Collins (6) has
reported the variety “sensibilis” as possessing poisonous prop-
erties. Other authors fail to mention the character of the species.
It should probably be classed with the poisonous forms, at least
until we have more knowledge of its effect when eaten.

BOLETUS CHROMAPES Frost

This species was free from haemolysin and agglutinin. No

muscarine was present. The species was poisonous to guinea-
pigs one animal weighing 370 grams dying in twenty-four hours
552 WILLIAM W. FORD AND JOSEPH L. SHERRICK

from 5 cc. of an extract of 3.2 grams of the plant in 32 cc. H2O

and a larger animal weighing 515 grams dying in fifteen days from
the same amount. The latter animal showed a progressive loss
in weight of over 100 grams. The plant had no poisonous action
on rabbits. According to Mcllvaine (7) Boletus chromapes is
an edible species.
BOLETUS AFFINIS Peck

Contained an agglutinin active,in a dilution of one-half of an

extract of 5 grams in 50 cc. H2O. This agglutinin was destroyed
at 65#{176}C. in half an hour. No haemolysin. No muscarine.
No definite poisonous action upon either guinea-pigs or rabbits.
Said to be edible by Mcllvaine (8).

BOLETUS ORNATIPES Peck

Contained an agglutiin in a one-fortieth dilution of an extrat

of 5 grams in 50 cc. 1120. This agglutiin was destroyed at 65#{176}
C. in half an hour. No haemolysin. No muscarine. No poison-
ous action upon rabbits or guinea pigs. McIlvaine reports it as
an edible species.

BOLETUS BICOLOR Peck

Contained an agglutinin in a one-twentieth dilution of an

extract of 2 grams of the plant in 25 cc. 1120. No haemolysin.
No muscarine. No definite toxic action upon animals. Both
Mcllvaine (10) and Hard (11) regard the species as edible,
Mcllvaine stating that it is one of the very best esculents.

BOLETUS SEPARANS Peck

Contained an agglutinin in a dilution of one-quarter of an

extract made from 5 grams in 50 cc. 1120. This agglutinin re-
sisted a temperature of 65#{176}
C. half an hour, but was destroyed
by heating to 100#{176}C. for the same length of time. No
haemolysin. No muscarine. Non-toxic to rabbits and guinea-
pigs. According to Mcllvaine (12) and Hard (13) this is an edible
plant.
 PROPERTIES OF SEVERAL SPECIES OF POLYPORACEAE 553

BOLETUS RAVANELII Berkeley and Curtis

No haeinolysin. No agglutinin. No muscarine. Non-toxic

to rabbits and guinea-pigs. Dietetic qualities not reported.

BOLETUS ROXANAE Frost.

Contained an agglutiin in a dilution of one-fourth of an extract

made from 4 grams in 40 cc. 1120. This agglutiin resisted
a temperature of 65#{176}
C. half an hour but was destroyed by
boiling the same length of time. No haemolysin. No musca-
rine. Non-toxic to guinea-pigs. A rabbit weighing 1205 grams
treated with 4 cc. of this extract developed a gradual loss in weight
and was found dead thirty-six days later. At autopsy this ani-
mal showed no lesions like coccidiosis or pneumonia which could
account for its death. It probably died of a chronic intoxication
due to some peculiar substances present in the plant. No note
concerning the dietetic properties of this species could be found
in the literature.

STROBILOMYCES STROBILACEUS Berkeley

No haemolysin. No agglutinin. No muscarine. Non-toxic

to guinea-pigs or rabbits. Edible according to Atkinson (14)
Mcllvaine (15) Hard (16) and other authors. Mcllvaine states
that it has a strong woody flavor and a faint taste of anisette
or of musk.
GENERAL CONSIDERATIONS

Owing to the limited number of these observations on the

various species of Polypores no very comprehensive statements
can be made as to the characters of the group. One or two points
of considerable interest and importance may be mentioned how-
ever. Haemolytic substances were not found in any of the spe-
cies examine#{231}l. Agglutinins were present in six species of boleti
out of a total of nine, being found in Boletus affinis, Boletus
ornatipes, Boletus bicolor, Boletus miniato-olivaceus, Boletus rox-
anae and Boletus separans. In two instances Boletus affinis
554 WILLIAM W. FORD AND JOSEPH L. SHERRICK

and Boletus ornatipes the agglutinating substances were destroyed

at 65#{176}
C. in half an hour, in twO species, Boletus roxanae and
Boletus separans they were destroyed only by boiling, and
in two cases Boletus bicolor and Boletus miniato-olivaceus they
resisted even this temperature. Agglutinins were not present
in Boletus chromapes, in Boletus felleus nor in Boletus ray-
enelii. Strobilomyces strobilaceus was likewise free from them.
In but one instance Boletus felleus was there a definite toxi-
city established for both guinea-pigs and rabbits and this
species is notorious for its extremely bitter taste. Several
species which are known to be edible, as Strobilomyces stro-
bilaceus, Boletus separans, Boletus bicolor, Boletus affinis, and
Boletus ornatipes exhibited no toxicity for either guinea-pigs or
rabbits. In these forms the lack of poisonous principles for ani-
mals can be correlated with their inability to produce harm when
ingested by man. In one species, Boletus ravenelii no bad effect
followed the injection of animals with the plant extract. No
note has been found in the literature as to the properties of this
species but in all probability it belongs to the edible fungi. In
two forms Boletus chromapes and Boletus miniato-olivaceus the
extracts were clearly poisonous to guinea-pigs but not to rabbits.
Boletus miniato-olivaceus is regarded by Mcllvaine as an edible
species. Its action on animals and the production of definite
symptoms in man by the ingestion of the closely related variety
“sensibilis” should make us cautious about expressing a decided
opinion about it, and for the present it may be looked upon as
suspicious. Thus far no statements have been found of the char-
acters of Boletus chromapes and a decision as to its classification
may be deferred till more knowledge is available concerning its
edibility. The species Boletus roxanae has a peculiar effect upon
animals, poisoning rabbits but not guinea-pigs. Its qualities
are not described in the literature and no opinion can be given
in regard to its properties.
 PROPERTIES OF SEVERAL SPECIES OF POLYPORACEAE 555

CLITOCYBE DEALBATA SUDORIFICA Peck

The species Clitocybe dealbata SOWERBY is considered an edible

fungus, both in this country and abroad, such authors as Mcli-
vaine (17), Hard (18) and Stevenson (19) putting a high value
upon it. Specimens of a closely related plant collected by Mr.
F. G. Howland at Saratoga Springs, N. Y., produced a profuse
perspiration in a number of individuals. The specimens were
submitted to Dr. C. H. Peck and in his case also their ingestion
was followed by a sweating which began on the forehead, gradu-
ally spread over the entire body, and lasted about five hours. No
other ill effects were experienced. Dr. Peck identified the fun-
gus as a variety of Clitocybe deal bata giving it the varietal name
sudorifica. His account of its botanical characters and the effect
it produced when ingested appears in the New York State Museum
Bulletin, No. 150 (20).
A few plants of the same lot collected by Mr. Howland were
sent to us for an examination into their effect upon animals.
Since but a small quantity of the fungus was available the dried
plant was maccrated in about twice the usual quantity of water,
0.8 grams being extracted with 16 cc. H2O. No tests with blood
corpuscles were carried out. A dose of 7 cc. of the extract was
administered subcutaneously to a rabbit weighing 1520 grams.
In a surprisingly short period of time, from three to five minutes,
a profuse salivation appeared, the animal becoming weak and
sick. The excessive salivation continued, accompanied by a
discharge of urine and faeces, for a period of three hours. Grad-
ually the condition of the animal improved, the salivation became
less marked and eventually wore off completely. By the next
morning the animal appeared as well as ever. It was kept under
observation for a number of weeks and finally died of an infec-
tion. About 3 cc. of the same extract given a guinea-pig
weighing 563 grams produced symptoms of intoxication in a few
moments. The animal began to make peculiar chewing move-
ments, lay motionless on its side and died within fifteen minutes.
Another guinea-pig, 665 grams in weight injected with 4 cc. of
the extract previously boiled half an hour developed a paralysis
556 WILLIAM W. FORD AND JOSEPH L. SHERRICK

of respiration in seven minutes, the heart continuing to beat

two to three minutes after the cessation of respiratory movements.
This animal showed the characteristIc gasping movements of
air hunger. In neither of these animals was there any unusual
appearance at autopsy. In another rabbit weighing 1335 grams,
5 cc. of a similar extract produced a salivation within one minute.
The saliva continued to flow from its mouth in a continuous
stream for twenty-five minutes and then gradually diminished
in amount. At the end of an hour the pupils of the eyes were
contracted as compared with those of an untreated animal. The
rabbit was likewise found to have discharged both urine and
faeces. The symptoms continued till the death of the animal at
the end of about two hours. The saliva from this animal was
collected and 10 cc. injected subcutaneously into another rabbit.
No salivation resulted in the second animal. Finally a third
guinea-pig weighing 370 grams was given 1 cc. of the extract. It
developed a marked conjunctival secretion within one and one-
half minutes, the respiratory movements being increased in force
and frequency at the same time. Within four minutes saliva-
tion appeared, the secretion from nose, mouth and eyes remaining
excessive for a period of twenty minutes. These secretions

now gradually diminished in intensity, and a diarrhoea developed.

The animal, after a period of rapid forced respiration, died of
respiratory paralysis in one and one-half hours after treatment,
the heart continuing to beat for a couple of minutes, after all
respiratory movements had ceased.
The pupillary constriction observable in these rabbits was
brought out in a more characteristic form by the local applica-
tion of the plant extract to a rabbit’s eye. About four drops
were instilled upon the conjunctiva of a normal animal. The
pupil of this eye contracted within twenty minutes, the constric-
tion lasting for a period of four hours.
The action of the fungus upon animals is characteristic of poi-
sons belonging to the muscarine--pilocarpine series, the exces-
sive salivation, conjunctival secretion, urination, diarrhoea,
and stimulated then paralysed respiration being described for
both bodies. Pilocarpine however does not appear to have been
 PROPERTIES OF SEVERAL SPECIES OF POLYPORACEAE 557

found in the fungi while muscarine is a constituent of A manila

muscaria. Amanita pantherina, Russula ernetica, and Boletus
luridus. (See Kobert [21]). The action of this Clitocybe can
thus be referred with greater confidence to muscarine than to
pilocarpine. The probability of muscarine being present was
rendered greater by comparing the effect of this plant extract
with that of Amanita muscaria, various animals being treated
with the two fungi side by side and a careful comparison being
made of the symptoms which followed. Guinea-pigs dosed with
Amanita inuscaria exhibited the same conjunctival secretion, the
same salivation, and the same diarrhoea, dying with symptoms
identical with those previously observed for the Clitocybe ani-
mals. With rabbits the symptoms shown by an animal given
4 cc. of the Clitocybe extract did not materially differ from those
shown by several animals with various doses of Amanita mus-
caria, the salivation and diarrhoea being the same. The Clito-
cybe however seemed to be much more poisonous, a rabbit weigh-
ing 1670 grams dying in less than two hours from a dose of 4 cc.
of an extract made in the proportion of 1 gram to 20 cc. 1120.
Large doses, 6 cc. and 8 cc. of the Amanita muscaria extract made
in the proportion of 1 gram to 10 cc. H2O produced both saliva-
tion and diarrhoea in rabbits but in no instance proved fatal.
This difference in the action of the two plants seemed purely
quantitative and could easily be explained by assuming that the
Clitocybe contained a greater amount of the poison in propor-
tion to the size of the plant than the Amanita. Finally the Cli-
tocybe extract when tested upon the exposed heart of a pithed
frog showed a reaction characteristic of the muscarine-pilocarpine
series. A drop of the extract diminished the frequency of the
heart beat from 50-60 per minute to 5-6 per minute. A drop
of a weak solution of atropine restored the activity of the heart
to normal, after which several drops of the extract had but little
effect, failing to reduce the frequency of the heart’s action to less
than 40 per minute. In another frog four drops of the extract,
introduced into the ventral lymph sac, produced within ten min-
utes a marked diminution in the heart rate which was reduced
to 5 or 6 per minute. Within twenty minutes a complete stop-
558 WILLIAM W. FORD AND JOSEPH L. SHERRICK

page of the heart had developed and more than one hour later
the application of a drop of the atropine solution brought back
the heart’s action to normal within fifteen minutes. This long-
continued pause is characteristic of muscarine, piocarpine pro-
ducing but a temporary cessation of the heart’s activity as Har-
nack and Meyer have shown (22).
It is thus evident that Clitocybe dealbata sudorifica PECK con-
tains a poison similar in its action to muscarine or piocarpine,
with the probability vastly in favor of the poison being muscarine.
The absolute identification of this body can only be accomplished
by a chemical analysis of the plant. We expect to undertake
this analysis in the near future. In view of the poisonous action
of this fungus upon man when ingested nd upon animals as
a result of subcutaneous injection it should probably be given
specific rank and not be regarded as a variety of Clitocybe dealbata
SOWERBY which apparently contains no poisonous principles
acting upOn man.

BIBLIOGRAPHY

(1) Ford: Jour. Pharmacol. and Exp. Therap., vol. ii, no. 4, March, 1911.
p. 285.
(2) Kobert: Lehrbuch der Intoxikationen, 1906, Bd. ii, 2, s. 1224.
(3) Peck: Boleti of the United States. Bulletin of the New York State
Museum, vol. 2, no. 8, Sept., 1889.
(4) Atkinson: Mushrooms, edible, poisonous, etc., 1903.
(5) Mcllvaine and Macadam: One Thousand American Fungi, 1900, p. 424.
(6) Collins: Rhodora, vol. i, Feb., 1899, no. 2, p. 21.
(7) Mcllvaine and Macadam: One Thousand American Fungi, 1900, p. 464.
(8) Mcllvaine and Macadam: One Thousand American Fungi, 1900, p. 448.
(9) Mcllvaine and Macadam: One Thousand American Fungi, 1900, p. 440.
(10) Mcllvaine and Macadam: One Thousand American Fungi, 1900, p. 425.
(11) Hard: The Mushroom, edible and otherwise, 1906, p. 352.
(12) Mcllvaine and Macadam: One Thousand American Fungi, 1900, p. 445.
(13) Hard: The Mushroom, edible and otherwise, 1908, p. 369.
(14) Atkinson: Mushrooms, 1903, p. 184.
(15) Mcllvaine and Macadam: One Thousand American Fungi, 1900, p. 475.
(16) Hard: The Mushroom, edible and otherwise, 1908, p. 380.
(17) Mcllvaine and Macadam: One Thousand American Fungi, 1900, p. 93.
(18) Hard: The Mushroom, edible and otherwise, 1908, p. 104.
(19) Stevenson: British Fungi, 1886, vol. i, p. 78.
(20) Peck: Bulletin of the New York State Museum, no. 150, p. 43.
(21) Kobert: Lehrbuch der Intoxikationen, 1906, Bd. ii, 2, s. 1223, 1224.
(22) Harnack and Meyer: Arch. f. exp. Path. u. Pharmakol., 1880, Bd. 12, s. 366.