PRESENTED

TO

The University of Toronto

Xc^ >fcA.^>c.^r-5 ^^^^

 f
ON THE

POISON OF YENOMOUS SMKES

AND

THE METHODS OF PREVENTING

DEATH FROM THEIR BITE

EEPEINTED PAPEES
BY

The late(Sir)JOSEPH FAYEER, Bt., K.C.S.L, M.D., F.R.C.R, F.R.S.,

Sir LAUDER BRUNTON, Bt., LL.D., M.D., F.R.C.R, F.R.S.
AND

Major LEONARD ROGERS, LM.S., M.S., F.R.C.P., F.R.C.S.

MACMILLAN AND CO., LIMITED

ST. MARTIN'S STREET, LONDON
1909
QP
 PEEFACE.

Scientific literature is increasing at such a pace at present

that it is difficult to keep completely acquainted with even one
branch of it. In consequence, some workers publish their own
results without taking the trouble to ascertain what other men
have been doing in the same field. Others, again, belong to the
" Ten Year School," who systematically neglect all work except
that done within the last ten years. But there are many more,
thoroughly conscientious workers, who try to find out all that
has been done in their own field of investigation, so that their
own observations may be rightly fitted in and help to build up
a solid structure of knowledge. But even these are often
hampered by the difficulty of obtaining the original papers to
which they would like to refer, and consequently remain
unacquainted with observations which may be of considerable
importance. This seems to have been the case with the papers
here reprinted by Sir Joseph Fayrer and myself, because in his
admirable work on Venoms, Calmette credits Lacerda with the
discovery of the antidotal power of permanganate of potash and
himself witli that of chloride of gold, although both of these
substances, as will be seen from pages 137 and 149 of these
reprints, were shown by us to be active a good many years ago.
With the concurrence of Lady Fayrer and Major Leonard
Eogers, and by permission of the Eoyal Society, I thought it
would be advantageous to republish these papers, not only for
the purpose of giving wider circulation to the work of the late
Sir Joseph Fayrer, but in the hope that they may be useful to
other workers in the same field.
LAUDEE BEUNTOK
London,
January, 1909.
 CONTENTS.

PAGE

On fclie Nature and Physiological Action of tlie Poison of Naja

tripudians and other Indian Venomous Snakes-
Part I r 1
Part II 23

On the Nature and Physiological Action of the C/*o^«Z«6-poison as

compared with that of Naja tripudians and other Indian Venomous
Snakes ; also Investigations into the Nature of the Influence of
Naja- and C rotalus -^^oiiow on Ciliary and Amoeboid Action and on
Fallisneria, and on the Influence of Inspiration of pure Oxygen on
Poisoned Animals ... ... ... ... ... ... ... ... Ill

Note on Independent Pulsation of the Pulmonary Vein? and Vena Cara 134
Note on the Effect of Various Substances in Destroying the Activity of
Cobra-poison 137

Experiments on a Method of Preventing Death from Snake Bit"', capable

of Common and Easy Practical Application 149
ON THE NATUEE AND PHYSIOLOGICAL
ACTION OF THE POISON OF NAJA
TRIPUDIANS AND OTHER INDIAN
VENOMOUS SNAKES.— Part I.
By T. Lauder Brunton, M.D., Sc.D., M.RC.R, and J. Fayrer,
C.S.I, M.D, FRCP. Lond., F.R.S.E., Surgeon-Major Bengal
Army.

(Reprinted f i*om the Proceedings of the Royal Society ^ No. 145, 1873.)

On the Poison of Naja tripudians.

The destruction of life in India by snake-bites is so great, that,
with the hope of preventing or diminishing the mortality, in
1867 Dr. Fayrer began, and has recently completed, a pro-
tracted and systematic series of investigations on the subject in
all its aspects ; and, in a work entitled the TJianatophidia of
India, has published a description of the venomous snakes
found in British India, with an account of a series of experi-
ments on the lower animals, conducted for the purpose of
studying the nature of the poison, its modus operandi, and the
value of the numerous remedies that have been from time to
time reputed as antidotes — that is, as having the power of
neutralising the lethal effects of the virus, and of saving life.
His object in carrying out these investigations has been :—
1st. To ascertain the nature and relative effects of the bite
of the different forms of Indian venomous snakes, and the
conditions and degrees of intensity under which the activity
of the virus is most marked.
2nd. The physiological action of the virus, and its mode of
causing death.
3rd. The value of remedies, and the extent to which we may,
by preventive or therapeutic measures, hope to save life.
(95) B
 Z ON THE NATURE AND ACTION OF THE

4th. To ascertain and make known the actual state of our

information in connection with tliese tliree points of inquiry,
and to substitute scientific and rational knowledge for vague,
empirical, and dangerous theories.
He has had the honour of submitting a copy of this work to
the Eoyal Society; and it is therefore unnecessary to occupy
its time by repeating much of what is therein related on the
1st, the ord, and part of the 4th heads.
But on that which is involved in the 2nd, and partly in the
4th, much is still required to be done ; and therefore on the
question of the nature and physiological action of the virus on
life, and the application of that knowledge in the treatment of
those poisoned, the following investigations have been made.
That the subject is one of interest in a purely scientific as
well as sanitary point of view we believe will be admitted ; for
it is as important to humanity as to science that the nature
and properties of a poison which, in India alone, probably
destroys over 20,000 human beings annually should be
determined.
We are aware that these figures may excite astonishment
and even mistrust ; but the sources from which the informa-
tion isderived place it, we think, beyond a doubt, being derived
from official returns for the year 1869, supplied to Dr. Fayrer
by the Government of India.
He has received reports from Bengal, the North-west
Provinces, Punjaub, Oude, Central Provinces, Central India,
Eajpootana, British Burniah, showing the loss of life from
snake-poisoning in those provinces in the year 1869.
These records represent, it is true, only a portion of India, as
the Madras and Bombay Presidencies, as well as other parts of
India, are not included. Had similar information been
obtained from these provinces, the list of mortality would
doubtless have been much larger; as it is, the number of
deaths is perfectly appalling, and the subject merits considera-
tion, with the view of providing, if possible, some remedy.
He has roughly classified the deaths under the headings of
the snakes that inflicted the fatal wound ; but the records are
rather vague on this point, and the information not perhaps
 POISON OF SOME INDIAN VENOMOUS SNAKES. 6

always very reliable. Still they are sufficiently explicit to

make it clear that, in order of clestructiveness, the cobra {Naja
tTipudians) occupies the first place on the list; the krait
{Bungarus cceruleus) the second place ; whilst under the
headings of " other snakes " and " unknown " must be included
many deaths due to cobra, Bungarus cceruleus, Ophiophagus,
Dahoia, Echis carinata, Bungarus fasciatus, Hyclropkidce, and
some perhaps to the Trimeresuri, though, as to the last, there is
reason to believe that deaths from their bites are comparatively
very rare.
The total number of deaths recorded therefore stands thus '.—
Bengal, including Assam and Orissa 6,645
IN'orth- west Provinces 1,995
Punjaub 755
Oude 1,205
Central Provinces 606
Central India 90
British Burmah 120

Total 11,416

of a population (according to Dr. Hunter) of 120,972,263, or,

in round numbers, about one person in every 10,000.
This total, large as it is, we fear cannot be regarded as the
real mortality in these provinces, nor may the numbers be
accepted as an absolutely true indication of the relative
frequency of deaths in each.
The information from which these records were framed was,
though official, probably only partial and imperfect. Dr. Fayrer
believes that if systematic returns could be kept, as he has
suggested that they should be, by the police in every district,
subdivision, and municipality, the number of deaths would be,
excluding all doubtful cases, much larger. He believes also
that were such information collected throughout the whole of
Hindoostan, it would be found that more than 20,000 persons
die annually from snake-bite.
The result of his investigations in India has been, we think,
to show that, so far, no agent or antidote, as that term is
(95) B 2
4 ON THE NATUIJE AND ACTION OF THE

commonly understood, has been found effective in neutralising

the action of snake-poison. We think it is also pretty clearly
demonstrated that death is caused in most cases, at all events
where a full quantity of the virus has been injected, by its
action on the nerve-centres, though whether on them alone, or
also on the peripheral distribution of the nerves, or on the
muscles themselves, or the exact extent to which each is
affected, there may be some difficulty in determining. The
futility of all the methods of treatment hitherto had recourse
to is probably explained by the mode of death ; their inutility
had long since been demonstrated by Fontana, who, ninety years
ago, among other things, showed that the outward and inward
use of ammonia, as well as its injection into the veins, was as
powerless for good as were all other remedies.
There is apparently some analogy between the nature of the
action of the cobra-virus and that of curara, death in both
cases being brought about by arrest of respiration through
paralysis of the respiratory apparatus.
In the case of the curara it has been demonstrated by
experiment that this is due to paralysis of the peripheral
distribution of the motor nerves ; and it has been further
shown that if respiration be continued artificially for a sufficient
length of time, perfect recovery may take place, as we have
ourselves observed, the poison being eliminated from the
system, and not having, during its presence, so far compromised
the integrity of the parts of the nervous system where it took
effect as to interfere with a resumption of their functions after
its removal. Now it is evident that artificial respiration and
the use of any remedies that may expedite elimination, with
the application of artificial warmth to sustain temperature up
to the normal standard, are the measures which may be
regarded as antidotal in a rational sense to this form of
poisoning ; and such they have proved themselves to be ; for if
an animal apparently dead from curara-poisoning be kept warm
and artificial respiration be kept up for some hours, it will
perfectly recover.
It is in the application of similar principles that we may
hope to realise a similar result in cases of snake-poisoning;
 POISON OF SOME INDIAN VENOMOUS SNAKES. 5

and it is with this object that the investigations by Dr. Lauder

Brunton and Dr. Fayrer, since his return to England, of which
the present paper is an instalment, have been pursued.
Our investigations so far confirm the opinion by Dr. Fayrer
already recorded, that death is due to the action of the poison
on the nerve-centres, to which it is conveyed by the blood
with terrible rapidity when the injection of the poison takes
place into a large vein like the crural or jugular. But we
have not yet arrived at absolute conclusions as to the extent to
which this neurotic action is carried, whether it be localised in
the nerve-centres only, or whether there be, and to what
extent, any action on other portions of the nerve-apparatus.
Our experiments so far, though pointing distinctly to the
centres as the seat of its action, in some cases seem to imply
that the nerve-periphery and perhaps even the muscles them-
selves are involved ; but on this head, for the present, we
reserve the expression of a positive opinion.
With reference to remedial measures in cobra-poisoning, we
would remark that, so far as our experiments have as yet gone,
artiticial respiration has certainly had the effect of prolonging
life; and without committing ourselves to any opinion, we
would say that we would not yet abandon hope that it may, as
in the case of the curara, even save it altogether. This must,
of course, depend on, first, the nature of the action of the
poison on the nerve-apparatus — that is, whether it be of
a transient or permanent character. Is it, for example, like
curara, which though it destroys the power of the peripheral
extremity of the motor nerves during its presence, yet leaves
them uninjured and capable of resuming their functions
after the poison is removed (as it may be) by elimination,
life being supported by artificial respiration during that
process.
If so, and the cobra-poison, even though antagonistic and
annihilative of the action of the nerve-centres and peripheral
distribution, or of the muscular irritability itself, be only so
whilst it is present, and would, if removed within reasonable
time, leave the nervous apparatus or muscles in a condition to
resume their operations, then, if elimination could be carried
6 ON THE NATURE AND ACTION OF THE

on whilst respiration is artificially sustained, we might hope to

succeed eventually in cobra as in curara poisoning.
Or could we, indeed, conceive of and find any agent so
subtle as to overtake and neutralise the virus whilst it is in
the system, and before it should have compromised the nerve-
centres or other parts, then we should have the antidote which
has been so long sought for, but yet, we fear, not found.* We
do not now wish to speak of the action of the cobra-virus as it
operates secondarily on the blood, either in those cases where
great vigour of the animal or smallness of the dose have
enabled the creature to resist the immediate and deadly
neurotic effects of the poison. Such cases are to be classed
among other septicaemiaB, and are apart from that we are now
discussing.
The question resolves itself into three points of inquiry :—
1st. Is the nature of the virus such that we may hope
to find any agent that may overtake, neutralise, and so render
it (the virus) harmless or inert ?
2nd. Does the virus exert only a temporarily pernicious
action on the ultimate structure of the nerve-centres or other
parts of the nerve-apparatus ? i.e., is it only inhibitory or
hurtful during its presence in the blood, but if removed would
leave the nerve-apparatus in a condition to resume its
functions (such is curara), or does it enter into some perma-
nent composition or union with the nerve-elements ? or, 3rd,
does it so modify their arrangements as to render them
permanently incapable of resuming their functions, even after
the poison has been eliminated, if it may be so removed, as we
know other poisons may ? Such, we fear, may be snake-
poison !
If the first proposition be correct, then in some subtle
chemical agent, or. if the second, in artificial respiration and
eliminant action we may have hope of success.
* Fontana thought he had discovered such an agent in the " pierre a cautere "
(caustic potash). He says of it :— "Mais on peut point douter cependant de
I'efficacite de ce remede, et on peut affirmer que la pierre a cautere est le vrai
specifique de ce terrible renin." — Sur les Poisons, p. 324 (Florence, 1781).
This agent has been tried in India, but has not proved of any service in cobra-
poisoning.
 POISON OF SOME INDIAN VENOMOUS SNAKES. 7

If the third, what chance have we beyond that of sustaining

life as long as artificial respiration be maintained ? for if the
nerve-apparatus be permanently injured, no resumption of its
functions can take place. Whichever of these propositions be
nearest the truth, there must still be a condition in which
from the smallness of the quantity of virus inoculated, recovery
is possible— one in which the full lethal effect of the virus is
not produced. In such cases, no doubt, remedial measures
may be of avail.
The results of investigations in India have led to the con-
clusion, then, that death is brought about by the action of the
poison on the cerebro-spinal nerve-centres, paralysing them,
and in some cases, where the quantity of virus VN^as large and
introduced into the circulation through the medium of a large
vein, acting directly on the ganglia of the heart, causing arrest
of its action. In those cases where the quantity of virus
inoculated is smaller and of less intensity, according to th-e
condition of the snake or its species (the poison of some genera
being less active than that of others), secondary changes,
though of what precise kind we are not yet prepared to say,
occur in the blood itself, but allied in character to that of other
blood-poisons and probably of a zymotic nature. We would*
merely for the present remark that, in the first class of cases,
we believe that remedies or means of treatment other than
those which may be of a preventive character are as yet of no
avail, whilst in the second it is probable that they may be of
some efficacy. So far we believe little more has been done
than to go over ground that has already been traversed by
previous observers, who have come to similar conclusions that
most of the reputed antidotes have been powerless, and that
where there has been an appearance of success, it has depended
not on any antidotal or antagonistic action of the remedy so
much as on the fact that the quantity or quality of the poison
was defective ; and how this may be explained, Dr. Fayrer has
endeavoured to prove by showing that the snake may have been
exhausted, that its poison may be deficient in quantity or in
quality, or that it may have wounded without inoculating
sufficient of the poison to cause death, or more than to cause
8 ON THE NATURE AND ACTION OF THE

slight poisoning, and probably that, by a sphincteral arrange-

ment of fibres, as pointed out by Dr. Weir Mitchell to exist in
the rattlesnake, the snake may have the power of imbedding its
fangs without shedding its poison at all.
Much virtue has been recently attributed to one of the oldest
and most trusted of all antidotes — ammonia ; but it was long
ago shown by Fontana by repeated experiments that the injec-
tion of this agent into the veins, as well as its internal
administration and external application, were powerless (as
may be seen by reference to the following* pages of his works),
so it has proved in all the experiments made with it in India.
Any complete and satisfactory means of resisting, antagonising,
or eliminating the poison and of saving life are, we fear, still
unknown ; and it is in the hope that by determining the
physiological action of the poison we may make some advance
in our knowledge of this important subject, that the following
investigations have been undertaken with cobra-virus sent to
us from Bengal, and of which we hope to receive continued
supplies from Mr. Vincent Eichards, of Balasore, who, at our
request, is also carrying on a series of experiments on the
subject.

Appearance and Chemical Characters of Cobra-poison.

The poison when fresh is a transparent, almost colourless
fluid, of a somew^hat syrupy consistence, and not unlike glycerine
in its appearance. When quickly dried it forms a transparent
mass of a yellowish-brown colour, and resembling some kinds
of gum-arabic. The poison may be kept in a fluid state for
some months without undergoing any change, but after a certain
time it decomposes.
During decomposition it gives off a quantity of gas, which
has been ascertained by Dr. Armstrong to be carbonic anhydride,
and at the same time acquires a dark brown colour and a dis-
agreeable odour. The dried poison may be kept for a much
longer time without undergoing any apparent change.
* Traits sur le renin de la Vipere, vol. i, pp. 108, 109, 118, 120, 124, 129;
vol. ii, pp. 5, 6, 7 (Florence, 1781). Opusculi Scientijiei, Letter iv, pp. 125
et sea.
 POISON OF SOME INDIAN VENOMOUS SNAKES. 9

The chemical constitution of the poison has been examined

by Dr. Armstrong. He has not been able to separate from it
any crystalline principle. It is partially coagulated by heat ;
mineral acids produce in it a gelatinous precipitate ; absolute
alcohol throws down a white gelatinous precipitate ; a drop
of it evaporated with a little sulphate of copper solution and
then treated with caustic potash gives a violent coloration.
These reactions show that the chief constituent of the poison
is an albuminoid body. On an ultimate analysis being made,
very little difference was found to exist between the fresh
poison, the alcoholic precipitate, and the alcoholic extract.
This is the only ultimate analysis of the poison of any snake
which has yet been made, so far as we know. We quote the
results of it, and give the composition of albumen for com-
parison.*
- Crude poison. Alcoholic precipitate. Alcoholic extract, j Albumen.

Carbon, 43*55 45-76 43-04

Nitrogen, 43-30 6-60
14-30 12-45
Hydrogen
Sulphur 53-5
15-7
Ash 2-5
traces 7-0
7-1

We have recently received from Bengal some cobra-poison

dried and in appearance resembling dried gum. On this we
hope to report on a future occasion.
Although there is little difference between the composition
of the alcoholic precipitate and extract, there is an immense
difference between their physiological actions, the extract being
a virulent poison and the precipitate almost inert. It is to be

* Dr. Armstrong in his analysis does not appear to have arrived at the same
conclusions as the Prince of Canino (L, Buonaparte), who detected the presence
of a peculiar principle perhaps allied to ptyaline, to which he gave the name
Echidnine or Yiperine, in addition to fatty matter, salts, albuminous and
mucous substance. It has been suggested by Prof. Busk (vide Holmes's
System of Surgery, vol. v, p. 941) that the venom may reside in a principle
analogous to, though differing from, ptyaline. We would not, however, regard
Dr. Armstrong's analysis as conclusive, but hope to have the result of further
examination of larger quantities of the virus.
10 ON THE NATURE AND ACTION OF THE

observed that the poison examined by Dr. Armstrong had

already begun to undergo decomposition ; l)ut if it should be
found by further experiments that the properties of the extract
and precipitate from perfectly fresh cobra-poison are the same
as tliose of the poison he used, it will form a notable distinction
between the poison of the cobra and that of the rattlesnake.
The precipitate thrown down by alcohol from the poison of the
rattlesnake has been ascertained to be active, while the alcoholic
extract is inert {vide Weir Mitchell, Physiology and Toxicology
of the Venom of the Rattlesnake, Smithsonian Contributions,
1860, p. 36).
We have experimented on four different samples of poison
sent from Bengal, The first was originally a clear transparent
fluid ; but after keeping it decomposed and became almost
black, as already described. It retained its fluidity and activity
to the last. The third sample was of a light-brown colour,
quite solid, and resembling dry hard cheese in its consistency.
The second and fourth consisted of a clear, thin, transparent
fluid and a white curdy precipitate. ]N"one of these specimens
had the same activity as the first ; they produced similar
symptoms, but much less marked.

Effects of the Poison. — The local effects of the poison are

partial paralysis of the bitten part, occasionally pain in it,
ecchymosis around the spot where the poison has been intro-
duced, and sometimes in other and distant parts, and, if the
animal survives for some hours, infiltration and perhaps incipient
decomposition of the tissues and haemorrhagic discharge.
The general symptoms are depression, faintness, hurried
respiration and exhaustion, lethargy, nausea, and vomiting. In
guinea-pigs and rabbits peculiar twitching movements occur,
which seem to represent vomiting in them, and occasionally, in
fact, guinea-pigs do vomit. Dogs vom.it, are salivated, and
present an appearance as if the hair had all been rubbed the
wrong way, " staring." As the poisoning proceeds, paralysis
appears, sometimes affecting the hind legs first and seeming to
creep up the body, and sometimes affecting the whole animal
 POISON OF SOME INDIAN VENOMOUS SNAKES. 11

nearly at the same time. There is loss of co-ordinating power

of the muscles of locomotion.
Haemorrhage, relaxation of the sphincters, and involuntary
evacuations, not unfrequently of a sanguineous or muco-
sanguineous character, often precede death, and it is generally
accompanied by convulsions.
In fowls the appearance is one of extreme drowsiness ; the
head falls forwards, rests on the beak, and gradually the bird,
no longer able to support itself, rolls over on its side. There
are frequent startings, as if of sudden awaking from the drowsy
state.*
The effects of the poison upon dogs, guinea-pigs, and rabbits
are illustrated by the following experiments.
The poison which was first sent home and still remained
perfectly liquid, but had become of a dark brown, almost black
colour, and somewhat inspissated, was used.

Experiment I.
1.30. Three drops of this, diluted with water, were injected
into the flank of a small dog. Immediately after the injection
the corresponding leg was drawn up, partially paralysed.
1.32. He walks less steadily. Tail rigidly held out.
1.35. Is restless and whining. Walks about and then sits
down again. Walks unsteadily.
1.45. There are distinct muscular twitches in the shoulder.
General tremor.
1.47. There are twitching movements of the back.
2.8. Has been standing perfectly still. Is now pawing and
licking his lips. Vomits.
2.10. Vomits again, but licks up part of what he had ejected.
2.22. Has been continually vomiting. The ejection consisted
at first of food, afterwards of tenacious mucus. He now lies
down apparently exhausted. He is still trying to vomit, but
* In cases where the quantity of poi?on injected is large, and it is at tlie same
time very active, the bitten animal small and weak, or if inoculation has taken
place into a large vein, death is almost sudden, as if it were from shook. In
such cases the cardiac ganglia are also probably paralysed; at all events the
heart suddenly ceases to beat.
12 ON THE NATURE AND ACTION OF THE

can bring nothing up. He tries to rise, but cannot. Convulsive

struggles occur.
2.25. Breathing has ceased, but the cornea is still sensitive.
Convulsive attempts to vomit.
2.27. Cornea insensible. Heart is still beating strongly.
Death soon followed.

Experiment II.
A young rabbit, weighing 900 grammes, was used. An
incision had been previously made through the skin of the
neck and the wound again sown up, but the animal was
otherwise uninjured. Two drops of cobra-poison, weighing
12 centigrammes, were diluted with 1 c.c. of water.
At 4.6 the diluted poison was ejected under the skin of the
left hip.
4.7. Washed out the watcli-glass in which the poison had
been placed with water, and injected it under the skin of the
back. The animal sat quiet after the injection, occasionally
licking its fore paws.
8' 30''. Eespiration seems hurried. The rabbit occasionally
makes a jerking motion with its hind feet.
10'. Has been restless, running about, occasionally lickiug its
fore feet.
13' 30". Still very restless, and when held makes convulsive
efforts to get away. Ears are much congested.
17'. The animal is now quiet. Its ears are no longer
congested.
About 20'. Quiet, with occasional starts. Disinclined to
move, but can walk quite well.
25'. Movements seem difficult, and hind legs seem weak when
it tries to walk.
26'. Paralysis of hind feet is increasing.
26' 15". The rabbit lays its head down on the table.
28'. When laid on its side it merely makes a few slight
movements with its fore paws and then lies still. The eyes
remain in a half-closed condition, and have done so for some
time. When the cornea is touched the head gives a jerk, but
the eyelids move very little. Eespiration slow and laboured.
 POISON OF SOME INDIAN VENOMOUS SNAKES. 13

4.30. The chin is twitched towards the sternum once or

twice, the hind feet at the same time being twitched back-
wards. The eyes open widely, Slight convulsive extension
of limbs.
4.31. Eespiration has stopped, cornea is insensible; thorax
opened immediately. There were large extravasations of blood
under the skin of abdomen and thorax, and under the skin of
the left hip. Heart beating vigorously.
The muscles contracted on direct irritation. The foot
twitched when the sciatic nerve was exposed and irritated by
an interrupted current. The peristaltic movements of the
intestine were active after the abdomen was opened.

Experiment ITI.

Dissolved 5 milligrammes of dried cobra-poison which had

collected round the stopper of the bottle containing it in
14 c.c. of water, and injected it under the skin of the left hip
of a guinea-pig, weighing 790 grammes.
In f of a minute after the injection the animal became
restless and uneasy and began to cry.
1|- minute it began to give little starts.
3J'. The starting motions became greater, the hind quarters
of the animal being jerked upwards, and the chin drawn in
towards the body ; continues to cry.
4J'. Passes water.
7'. Less restless.
15'. Washed out the watch-glass in which the cobra-poison
had been placed with about -|- c.c. of water, and injected it as
before. Immediately afterwards the restlessness increased.
24'. Seems to be trying to vomit.
27'. It cannot walk rightly.
28'. The hind legs are paralysed and spread out laterally
from beneath it.
29'. Eespiration very slow and deep. The animal lies
quiet, but convulsive twitches of the limb follow almost every
respiration.
Eespiration 8 in J a minute.
14 ON THE NATURE AND ACTION OF THE

30'. Cornea insensible. Eespiration has ceased. Post-

mortem examination made immediately. The left ventricle was
much dilated, the right ventricle empty. There were two
beats of the left auricle for every one of the ventricle, and the
ventricular beat was weak and imperfect.

Experiment IV.
Dissolved 1 centigramme of a substance like gum, and
labelled " alcoholic extract of cobra-poison," in 1 c.c. of water.
It dissolved easily and formed a somewhat opalescent solution.
Injected about one-third of this (equal to 3-J milligrammes
of the dried extract) under the skin of the thigh of a rabbit
weighing about a kilogramme.
Four minutes after the injection there was no apparent
effect; so a similar quantity was again injected, making the
total amount received by the rabbit 7 milligrammes of
extract : 5J minutes after the first injection the animal became
very restless.
7'. Eespiration rapid. The vessels of the ears were noticed
to be much injected. On continuing to observe them the
injection disappeared and then returned again. The alternate
filling and emptying of the vessels was much more perceptible
than in the normal condition. The rabbit sits quietly, but
every now and then gives a start.
22'. The condition of the ears has continued the same. The
eyes are becoming half shut and the eyeballs turned up.
The animal now begins to tremble. The head is laid down
on the table and then raised again : this is succeeded by a
nodding motion of the head. The head is next laid down on
the table,
Eespirations 22 in 15 seconds.
24'. The animal has sunk down on its face and paws, as if
its fore legs would no longer support it. The hind legs,
however, still support the posterior part of the body. Eespira-
tions 11 in 10 seconds. It seems to be trying in vain to raise
its head.
26'. Eespirations 8 in 10 [seconds. Convulsions. The
 POISON OF SOxME INDIAN VENOMOUS SNAKES. 15

cornea is sensitive. The rabbit is now lying on its side.

Eespirations 5 in 15 seconds. Pulse 12 in 18 seconds.
31'. Cornea is nearly but not quite insensible. The eyeball
is protruding.
About olY respiration has stopped. The heart is still
beating vigorously.
32'. Cornea insensible. The animal opened immediately
The heart was beating vigorously ; 21 beats in 10 seconds.
An attempt was made to insert electrodes into the spinal
cord and pass an interrupted current through them. No effect
followed ; but it is not certain that they were well in the
cord. Irritation of the nerves going to the hind legs by
uninterrupted current had but a slight effect. Direct irritation
of the muscles caused them to contract. After the irritation
was discontinued, a fibrillary twitching was observed in one of
the extensions of the thigh.
42'. Heart still feebly pulsating. Irritation of the brachial,
sciatic, and crural nerves has very little effect.
45'. Heart still feebly pulsating.

Experiment V.

Two drops of cobra-poison were injected under the skin of

the thigh of a guinea-pig.
One or two minutes after the injection the legs of the
animal began to twitch. It was then covered with a glass
bell-jar.
Six minutes after injection. The legs are again twitching.
This is a peculiar motion of the hind legs, in which they seem
to make an abortive attempt to kick involuntarily.
7'. Eespirations are deeper than usual.
9'. Legs again twitching.
10'. The animal is restless and moves round and round inside
the bell-jar. Grunts occasionally and grinds its teeth. The
hind-quarters are twitched upwards, and the nose is drawn in
towards the chin at the same time.
13'. Bites at the spot where the injection was made and
passes water.
16 ON THE NATURE AND ACTION OF THE

22'. It can no longer walk.

23'. It has sunk down and lies flat on the table, leaning
rather to one side. Respirations are deep. There are occasional
twitches of the legs.
25'. Cornea is sensitive. Occasional convulsive stretches.
27'. Cornea almost insensible. Respiratory movement of
nostrils continues.
28'. Cornea completely insensible. Post-mortem examination
made immediately. The muscles of the abdomen were dark-
coloured. Peristaltic movements of the intestines occurred
when the abdominal cavity was opened. The heart was dark
and slightly dilated ; all its cavities were contracting, though
feebly. There were three beats of the auricles to each one of
the ventricles. Irritation of the nerves in the pelvis caused
contractions of the legs.
35' after injection. The heart is still feebly contracting.

Experiment YI.
October 28th- — Injected about a grain and a half, or two
grains, of the precipitate, which was thrown down from cobra-
poison by alcohol, into the thigh of a guinea-pig.
2.30. Injection made. A few minutes afterwards it passed
fiome milky-looking water, and then remained perfectly quiet.
3.8J. Passed water, which was quite clear.
3.33. Injected about two grains into the right femoral vein.
It passed clear water almost at once.
3.35. Its nose gave a jerk inwards. Wounded leg drawn up.
3.38. ISTose twitches frequently and the animal emits a faint
barking sound.
3.40, Slight tremors.
3.50. Begins to eat a piece of bread placed near it.
3.58. Still twitches.
4.8. Is still sluggish, but seems nearly well. Recovered.

Experiment VII.
October 29th, 1872.— About half a grain of fresh but
-coagulated and cheese-like cobra-poison was suspended in
 POISON OF SOME INDIAN VENOMOUS SNAKES. 17

distilled water and injected into the back of a guinea-pig,

weighing about a pound and a quarter.
2.23. Injection made.
2.26. The animal looks scared and is twitching. This
guinea-pig is very active.
2.30. Another dose injected. The animal is twitching much.
It jumped out of the deep box in which it had been placed for
observation. Breathing is hurried.
2.36. It seems better. Another dose injected into the thigh.
2.45. Not much effect. Another dose injected.
2.46. Twitching continues ; animal remains active. It
recovered.

Means of preventing the Effects of the Poison.

There are three ways in which the toxic effects of a poison
may be entirely prevented or greatly diminished. These are :—
1st, by preventing its admission into the blood ; 2nd, by
counteracting the effects it produces while it is circulating in
the body and sustaining life by artificial respiration; 3rd, by
quickening its elimination. Tlie first of these methods is the
only one which has hitherto been of any great service in cases
of poisoning by the bite of cobras. Various attempts have
been made to counteract the effects of cobra-poison by means of
antidotes ; but the advantage derived from their use is still, to
say the least, doubtful. Xo special attempts, so far as we
know, have been made to hasten the elimination of the poison,
or at least none have been made avowedly for this purpose,
though it is possible that some of the antidotes may have had
that effect. This part of the subject we will treat in a future
paper.
The subject of prevention of entry of the virus by ligature or
other mechanical measures has been fully discussed in the
ThanatopJiidia ; it is unnecessary to recur to it here, for the
present at all events.
For the purpose of attempting to counteract the effects of the
cobra-poison while it is circulating in the blood, it is necessary
to have some idea of its mode of action.
(95) c
 18 ON THE NATURE AND ACTION OF THE

Mode of Actum of the Poison.

Snake-poison probably produces its fatal or deleterious effects
either by completely paralysing the nerve-centres or other
portion of the nervous apparatus, and thus causing arrest of
respiration, or by partially paralysing them and also poisoning
the blood, thereby inducing pathological conditions of a
secondary nature, which may, according to circumstances, cause
the slightest or the most dangerous symptoms.
The effect produced depends on two sets of conditions : —
first, the species of the snake, its actual state at the time, the
quantity and quality of its poison, and the circumstances
under which it inflicts the bite ; second, the species, size, and
vigour of the living creature, and the circumstances under
which it is bitten.
Snake-poison is essentially a neurotic, and, when it takes full
effect, it appears to kill by annihilating, in some unknown way,
the source or distribution of nerve-force. It is also an irritant ;
for if applied to a mucous membrane or to the conjunctiva, it
soon induces violent inflammation ; absorption at the same
time takes place, and symptoms of poisoning are produced. It
is also, to a certain extent, a septic ; for if the bitten creature
survive, the wound and the parts about it are apt to slough and
to induce septiciemia. The poison acts by absorption — that is,
by entering the circulation, and so reaching the nerve-centres,
it produces, according to the quantity or intensity of the
venom, either death or severe local and constitutional symptoms.
If it find entry by a large ^•ein, such as the femoral or jugular,
life may be destroyed in a few seconds.
The blood itself is affected by tlie poison.
Dr. Fayrer has not been a])le to detect any corpuscular
changes, nor has he any exact information on the chemical
changes it undergoes, or may have undergone; but that it is
altered there can be little doubt ; and in poisoning of the
lower animals, at all events by the Yiperidae, its coagulability
after death is generally destroyed, whilst after death by poison-
ing by tlie colubrine snakes the blood generally coagulates.*
* Our expeiiments in England have not confirmed these observations made in
Ind.a. The blood of animals dead from Daiota-poisoning has been found to
 POISON OF SOME INDIAN VENOMOUS SNAKES. 19

As the blood is tlie channel through which the poison acts, it

is obvious that the first object should be to arrest, destroy, or
prevent its entry into the circulation; or if it has already
entered, to neutralise or counteract its action, or to procure its
elimination by the agency of the natural depurating organs and
their secretions, and to treat local, consecutive, and constitutional
symptoms by such remedial measures as may be required by
the patient's condition.
Absorption takes place with extreme rapidity, so fast, indeed,
that it was formerly supposed, in the case of some of the more
active poisons, that they acted by transmission of a shock
through the nervous system ; and, so far as we know at present,
it is not improbable that such, in some instances, may be the
case. But rapid as the effect of snake-bite sometimes is, there
is no reason to believe that generally it operates on the nerve-
centres through any other channel that that of the vascular
system. The experiments of Blake, Hering, and, later, of
Claude Bernard show that absorption takes place with such
rapidity as to explain the most rapid deaths from such cause.
Blake {vide Guy's Forensic Medicine, 3rd edition, p. 388) found
that a poison passed from the jugular vein to the lungs of a
dog in from four to six seconds, from the jugular vein to the
coronary arteries of the heart in seven seconds; a poison
injected into the jugular vein was distributed throughout the
circulation in nine seconds. Claude Bernard found that a
saturated solution of sulphuretted hydrogen introduced into
the jugular vein of a dog began to be eliminated from the lungs
in three seconds, and when injected into the femoral vein of the
same dog in six seconds.
We have neither seen nor heard of any case of snake-
poisoning, in man or the lower animals, so rapid (though in
some Dr. Fayrer has observed the first symptoms in a few
seconds) as to justify the conclusion that poisoning had
occurred otherwise than through the medium of the circulation.
Some preliminary experiments made in England by one of
coagulate. This is a point tliat needs muoli further and repeated observation,
as, indeed, does the question of the chemistry of the blood of animals affected
b}- snake-poison, and we hope to report further on it.
(95) c 2
20 ON THE NATURE AND ACTION OF THE

US (Dr. Brunton) with the poison before it had undergone

decomposition seemed to show that it produced paralysis of
the spinal cord, of the ends of the motor nerves, and of the
muscles themselves. The experiments which we made together
with the same poison a few months afterwards, as well as with
other samples of poison sent from India, have not given con-
cordant results. We therefore propose to postpone the
consideration of this subject to a future paper, and to confine
ourselves at present to the mode in which death is produced by
the poison, especially in mammals.
Somatic death, according to Bichat, may commence in the
brain, lungs, or heart; but the experiments of Fontana and
Legallois show that so long as circulation and respiration are
kept up, the body remains alive although the head be absent.
The brain is only necessary to life, inasmuch as the respiratory
movements cease when it is removed or destroyed, either
mechanically or by the action of a poison upon it. The causes
of somatic death are thus limited to failure of the circulation
and failure of the respiration.
The long continuance of the cardiac pulsations after apparent
death (Experiments I, III, IV, V, IX, X) excludes failure of
the circulation as the usual cause of death ; and we are thus-
brought by exclusion to regard death caused by the bite of
a cobra, or by its poison introduced into the body in any other
way, as death from failure of the respiration, or, in other words,
death by asphyxia. The truth of this view is well illustrated by
the following experiments,* which show that the vitality of the
heart may be retained for a considerable time if the respiration
is kept up. It shows also that the convulsions which have
been remarked by Eussell and all subsequent observers as
almost always preceding death are not due so much to the
action of the poison itself on the nervous centres, as that they
depend on the irritation which is produced in them by the
venosity of the blood.

* Excepting those cases in which the poison is injected into a large vein, such
as the jugular, and causes sudden arrest of the heart's action.
 POISON OF SOME INDIAN VENOMOUS SNAKES. 21

Experiment VIII.
July, 1872. — A drop or two of cobra-poison diluted with
water was injected into the thigh of a strong fowl. Shortly-
after it began to droop. It then seemed drowsy, and crouched
down with the beak resting on the ground ; it then fell over on
its side. The comb and wattles lost their bright red colour, and
became dusky. Almost simultaneously convulsions occurred,
A cannula was quickly inserted into the trachea, and artificial
respiration commenced. The comb rapidly regained its bright
colour, and the convulsions ceased. On the artificial respiration
being discontinued the lividity of the comb reappeared, and con-
vulsions again began. The experiment was repeated about ten
times, and on each occasion the convulsions disappeared when-
ever the blood became arterial, as shown by the bright colour of
the comb, and reappeared when the blood became venous. After
discontinuing artificial respiration, the convulsions returned and
the fowl died.

Experiment IX.
November 7th, 1872. — A cannula was placed in the trachea
of a rabbit.
12.57. A small quantity of cobra-poison w^as injected into the
hip. Symptoms of poisoning came on slowly.
1.25. The animal is still breathing, but the limbs are almost
completely paralysed. Artificial respiration begun. Tempera-
ture in the rectum, 101°' 8.
1.37. Paralysis is now complete. The animal is perfectly
motionless, and not the slightest movement of the eyelids occurs
when the cornea is touched. Temperature in rectum, 100°'8.
1.55. The animal appears quite dead, but the heart pulsates
vigorously.
2.30. Cardiac pulsations as before. Temperature, 98°"6 E.
2.32. Heart as before. Temperature, 97°.
4.10. Heart still beats vigorously. Temperature, 95°'4. The
continuance of the artificial respiration was now entrusted to an
assistant.
5. Heart beating well.
22 ON THE NATURE AND ACTION OF SNAKE-POISON.

5.20. Heart beating feebly, and its action jumping.

5.30. Heart beating slowly.
6.30. Heart beating a little quicker.
7.30. Heart as before.
8. Heart beating more slowly.
8.30. Cardiac pulsations are very feeble.
9.30. Very feeble and slow.
• The hour was now late; the rabbit was still completely
motionless, and its body felt cold to the touch. The artificial
respiration was therefore discontinued, although the cardiac
pulsations had not ceased. Life was evidently prolonged for
some hours in this case by artificial respiration.

Experiment X.
November 28th, 1872. — One-fifth of a drop of cobra-poison
(the first supply), diluted with about 2 cub. centims. of ^-per-
cent. salt, was injected into the external jugular of a rabbit.
12.5. Injection made.
12.20. The animal has been convulsed and paralysed.
Sensibility of the cornea has disappeared ; cannula placed in
trachea and artificial respiration commenced. Temperature,
100°.
1.15. Temperature, 96°'3. Heart is beating vigorously.
3.13. Heart is beating as before.
3.20. In order to try if possible to quicken elimination, milk
was injected into the stomach.
4.5. Heart is beating as well as ever.
4.40. Heart still beating vigorously. Respiration discontinued.
Death soon followed. In this case also life was prolonged by
artificial respiration.
ON THE NATURE AND PHYSIOLOGICAL
ACTION OF THE POISON OF NAJA
TRIPUDIANS AND OTHER INDIAN
VENOMOUS SNAKES.— Part IL

By T. Lauder Brunton, M.D., Sc.D., M.E.C.P., and J. Fayrer,

C.S.I., M.D., F.R.C.P.Lond., FRS.E., Surgeon-Major Bengal
Army.

(Reprinted from the Proceedings of the Royal Society, No. 149, 1874.)

The effects of the poison of Naja tripiidians are probably the

same as those of Opliioi^hagus elaps, Bwngarns, Hydrophidse, and
other poisonous colubrine snakes, whilst that of Dctboia Bussellii
is similar to that of Echis carinata, and also of the Trimerestiri,
which represent the viperine snakes in India.
Just as the JVaja may be regarded as among the most virulent
of the colubrine, the Bahoia is probably as venomous as any of
the viperine snakes, it being very deadly ; whilst the Crotalidse
are but feebly represented in India by the Trim^resuri.
The venomous colubrine snakes in India are represented by
the Naja tripudians, Ophiophagus elaps, Bungarus fasdatui,
B. coeruleus, Xenurelaps hmgaroides, and the various species of
Callophis and Hydrophidse ; whilst among the viperine snakes
the Viperidse, or vipers, are represented in India by only two
genera, each with a single species, Bahoia Bussellii, Bchis
carinata ; the Crotalidse, or pit-vipers, by the various Trimere-
suri, Beltopelor, Halys, Hyynale, though these are much less
active than their American congeners.
The Bahoia, however, may be considered as virulent as the
most deadly form of the Yiperidse of Africa, or probably as the
Crotalus or Craspedocephalus of the pit-vipers of America and
the West Indies.
24 ON THE NATURE AND ACTION OF THE

In a previous communication we have described the effect of

the poison of Naja tripudians upon warm-blooded animals, and
have illustrated it by experiments on the dog, raljbit, guinea-
pig, and fowl.
We purpose in the present paper to compare its action with
that of the poison of the Dahoia Russdlii, a viperine snake, to
describe its effects upon cold-blooded animals and invertebrata,
and to examine in detail its action upon the various organs of
the body.
In our former paper we stated that the general symptoms of
poisoning by cobra-venom are depression, faintness, hurried
respiration and exhaustion, lethargy, unconsciousness, nausea,
and vomiting. In dogs, guinea-pigs, and rabbits, peculiar
twitching movements occur, which seem to represent vomiting
in them ; occasionally, in fact, dogs and guinea-pigs (Experi-
ment XX) do vomit, and dogs are profusely salivated. As the
poisoning proceeds, paralysis appears, sometimes affecting the
hind legs first and seeming to creep up the body, and sometimes
affecting the whole animal nearly at the same sime. There is
loss of co-ordinating power of the muscles of locomotion.
Haemorrhage, relaxation of the sphincters, and involuntary
evacuations, not unfrequently of a sanguineous or muco-
sanguineous character, often precede death, and are generally
accompanied by convulsions.
In fowls, the appearance is one of extreme drowsiness : the
head falls forward, rests on the beak, and gradually the bird,
no longer able to support itself, crouches, then rolls over on its
side. There are frequent startiugs, as if of sudden awaking from
the drowsy state.
The following experiments upon pigeons and guinea-pigs show
that the general symptoms produced by the poison of the
Dahoia are nearly the same as by that of the Naja. The local
symptoms are greater extravasation of blood and effusion into
areolar tissue. In Experiment III it was noted that greater
lethargy and less violent convulsions occurred in the pigeon
poisoned by cobra-venom than in that poisoned by Dahoia ; but
this might readily be due to individual difference in the bird ;
and an opposite result is noted in Experiment YII upon a
 POISON OF SOME INDIAN' VENOMOUS SNAKES. 25

guinea-pig. In one pigeon, killed by Baboia-w enom, the blood

remained permanently fluid after death ; but in the other, and
also in the guinea-pigs, it coagulated firmly. This is an excep-
tion to the rule which has been noticed in experiments made
in India, that the blood after Bahoia-'poisomng remains fluid —
in marked contradistinction to death from cobra-venom, in
which the blood almost invariably coagulates. Coagulation,
however, of the blood of a fowl after death from the bite of a
Dahoia has also been noticed by one of us (Dr. Fayrer) in India ;
and therefore the coagulation in our experiments was not due
to the lower temperature of the atmosphere.

Experiment I.
August 27th, 1873. — Three milligrammes of dried Dahoia-
poison, received some weeks ago from Balasore, were injected
into the thigh of an old and vigorous pigeon at 2.48.
2.53. No apparent effect, except that the bird is lame on that
leg.
3.2. The bird is sluggish. Eespirations hurried. Lameness
continues.
3.18. Still sluggish, but it is not deejjly affected.
3.30. Disinclined to move. When placed on the table it sank
on its breast. No nodding of the head,
3.45. Sudden and violent convulsions.
3.46. Dead in 58 minutes from the time of injection.
Electrodes inserted into the spinal cord soon after death
caused movements of the wings, but not of the legs. Blood
taken from the bird just before death partially coagulated
after death. Blood taken from it after death coagulated more
firmly, but less firmly than some taken from another pigeon
poisoned with cobra venom.

Experiment II.
A young full-grown pigeon had 3 milligrammes of dried
Baboia-i^oison injected into the peritoneum at 3.5 p.m.
At 3.13 it was observed to pass suddenly into violent con-
vulsions, flapping its wings strongly. It continued in this state
26 ON THE NATURE AND ACTION OF THE

for a minute; and at 3.14 it died, 9 minutes after the

injection.
Electrodes inserted into the spinal cord, in the neck, caused
violent muscular contractions all over the wings and legs. The
cord was thus evidently not paralysed ; but its irritability soon
ceased. The blood remained permanently fluid, and became
bright red on exposure to air ; under the microscope (400 dia-
meters) the corpuscles seemed normal. Rigor mortis came on.

'Experiment III.
A full-grown young pigeon had 3 milligrammes of dried
cobra-poison injected into the thigh at 2.49 P.M,
2.53. The respiration is very hurried ; the bird presents a
sluggish appearance and begins to droop.
3.2. The eyes are now closed and the bird is crouching ; legs
extended.
3.6. Convulsions ; head and back resting on the ground; legs
extended and paralysed.
3.10. Dead in 21 minutes from the injection.
Electrodes inserted into the cord soon after death caused
general contractions of the extremities, showing that the cord
was not paralysed. Its irritability soon disappeared. The
symptoms in this bird are different from those in the one
poisoned by Z>a&om- virus ; there is more lethargy, nodding of
the head, and apparent drowsiness before the convulsions, which
are not so sudden or so violent.

Experiment IV.
A full-grown pigeon had 3 milligrammes of dried cobra-poison
injected into the peritoneum at 3.5 p.m
3.15. The bird is sluggish, nodding its head.
3.17. Gaping ; the head is twitching, and the bird can hardly
stand.
3.22. Convulsions. Several grains of Indian corn are
vomited,
3.25. Quite paralysed. Convulsions.
3.26. Dead in 21 minutes from the injection.
Electrodes in the cord soon after death caused movements in
 POISON OF SOME INDIAN VENOMOUS SNAKES. 27

the limbs. The irritability rapidly disappeared, and at 3.33 was

entirely gone.
The blood coagulated firmly after death.
When examined after death with a magnifying power of
400 diameters, crenation of some of the red corpuscles was
observed, but no other change was noticed.

Experiment V.
February 11th. — About J— 1 c.c. of a mixture of Dahoia-
poison with alcohol (1 part poison with 4 of alcohol) was
injected into the left thigh of a small guinea-pig at 1.45 p.m.
Immediately afterwards it became very restless, and the nose
began to be twitched inwards towards the breast.
1.48. The left lecj drags somewhat.
1.54. The hind legs are jerked backwards regularly every few
seconds.
1.55. It bites at its left leg.
1.58. It has drawn itself together almost into a ball.
2.2. The twitching still continues.
2.23. Its hind quarters have become nearly paralysed. It
lies on its side, and convulsive movements occur from time to
time.
2.28|. It is apparently dead. The heart continues to beat
strongly. On opening it, the lungs were slightly congested.
Peristaltic movements of intestine active. The blood from the
heart was allowed to run into a clean beaker. It was of a dark
colour, but became red on exposure to air. It shortly after-
wards coagulated and formed a firm clot.

Experiment YI.
February 11th. — About 1 c.c. of Z^a&om-poison (1 part poison
mixed with 4 parts of alcohol) was injected under the skin of
the left thigh of a guinea-pig at 1.13.
1.17. Animal rubbing its mouth with its fore-paws. It is
restless and moves about. There are slight twitchings, and it
sits on its hind legs like a cat.
1.22. Very restless.
28 ON THE NATURE AND ACTION OF THE

1.27. Head is drawn towards legs in a twitching fashion.

Animal bites at the left leg. When it moves about, the left
leg drags somewhat.
1.45. Has been very quiet and disinclined to move for some
time.
1.55. About 1 c.c. more was injected into the right thigh.
1.56. Both hind legs drag slightly.
1.58. The animal is very unsteady and tottering on its legs.
2.2. Both hind legs completely paralysed, and, when the
animal draws itself forward with its fore-paws, the hind legs
trail out behind it. There are twitchings of the fore part of
the body.
2.17. Hind legs and loins quite paralysed. The posterior
part of the body lies flat on the ground, the abdomen being
flattened out upon it. Paralysis seems gradually extending to
the fore limbs. There is general twitching. It tries to crawl,
but cannot drag itself forward, though it can still move the
fore legs. Gnaws the bottom of the box in which it lies.
2.20. Almost motionless. Eye is still sensitive. Fluid has
issued from the mouth. The animal can still move its head.
2.23. Convulsive movements.
2.24. Cornea insensible. Weak twitches of the trunk still
occasionally occur ; they seem to be of the nature of respiratory
movements. Heart beats strongly.
In a minute or two afterwards the animal was opened. The
heart was irritable and contracted when touched. The ventricle
did not contract unless touched. The auricles were beating.
The lungs were (I think) slightly congested. Blood from the
large trunks in the thorax was collected in a vessel : it was of
a dark colour; on exposure to air it became bright red and
formed a firm coagulum. Peristaltic movements of the intestine
were observed.
Experiment VII.
February 11th. — About J c.c. of milky -looking cobra-poison
was injected into the right thigh of a guinea-pig of moderate
size at 2.20. It became restless immediately, and the hind
legs began to twitch backwards. Shortly afterwards it again
became quiet and sat quite still.
 POISON OF SOME INDIAN VENOMOUS SNAKES. 29

3.12. The animal did not seem to be much affected by the

poison. Some more injected into left thigh.
4. Both hind legs became paralysed, and the animal lay with
them spread out behind it. The hind part of the body also sank
down, so that the abdomen became flattened on the floor, just
as with the Dahoia-^oison.
4.23. Convulsive twitches occur. The animal lies on its
side. It is more convulsed than the one killed with Daboict-
poison.
Action of Cohra-jmson on Frogs.
When cobra-poison is injected under the skin of frogs they
occasionally become very restless immediately after the
injection. This, however, is by no means always the case ;
and as similar agitation occurs, often to a much greater extent,
after the injection of other substances, it is to be attributed
rather to the insertion of the needle than to the action of the
venom. A gradually increasing torpor then comes over the
animal, sometimes beginning some time after the injection, and
then proceeding uninterruptedly, at other times being inter-
rupted byoccasional movements. The limbs are drawn close
up to the body, and the head gradually sinks down between
the hands in most instances ; but sometimes, as in Experi-
ment VIII, the head is held at first much more erect than
usual. The power of motion is lost before that of sensation ;
for the movements caused by painful stimuli become weaker
and weaker, although they may still follow each application of
the irritant. The progressive weakness is well shown in the
movements of the hind legs. After the frog has sunk down
and is lying flat upon the table, pinching the toes causes it to
kick vigorously ; but by-and-by, instead of kicking, it merely
draws away tlie foot from the irritant with a slow wriggling
motion. If it is then lifted up from the table, so as to remove
the resistance occasioned by friction, the wriggling entirely
disappears, and the foot is promptly and easily drawn up to the
body when pinched. This weakness seems to depend on the
nervous system rather than on the muscles ; for, even in this
state of apparent paralysis, the animal occasionally displays
considerable muscular pov/er, and is able to spring to a
30 ON THE NATURE AND ACTION OF THK

considerable height, aa in the following experiment. A similar

condition is sometimes observed in warm-blooded animals, as in
Experiment LX. Tlie motor paralysis increases, no motion
follows the application of any irritant, however powerful ; but
even then sensation exists, as is seen from Experiment LXXVI.
The heart continues to beat after all motion in the body has
ceased ; but its pulsations become gradually slower, and at last
cease altogether.
Experiment YIII.
September 12th, 1873. — Three frogs of nearly equal size were
selected, and a dose of dried cobra-poison dissolved in water
was injected into the dorsal lymph-sac of each. The quantity
injected into No. 1 was estimated to be equal to three or four
■drops of the fresh poison, that into No. 2 about a drop, and into
No. 3 about half a drop. These estimates, however, are not to
be absolutely depended on.
The injection was made into all three about 3 p.m.
3.17. Nos. 1 and 2 are sitting with the head much more erect
than usual and the belly depressed. No. 3 has the head
depressed between the fore paws.
3.22. No. 3 is now sitting up in the normal posture.
4. No. 1 lies quite quiet ; when moved its limbs give a slight
wriggle. Applied strong acetic acid to its legs ; after many
iseconds it gave a faint wriggle. No. 2 also lies quiet. When
its legs are pulled back it can still wriggle them up towards its
body. When held up it can kick well. After being placed on
the table it suddenly, and without any apparent reason, sprung
up to a considerable height. No. 3 presents tlie same appearance
as No. 2, but seems more paralysed.
4.5. No. 1 does not react at all to any painful stimulus.
Nos. 2 and 3 wriggle their legs when pinched. The observation
was now discontinued. Next morning all three were dead.

A ciion on Lizards.

The action of cobra-pijison ujon lizards seems very similar

to that which it has upon ficgs ; the animal becomes sluggish
end diflicult to rouse ; and tbe bitten part is affected by
 POISON OF SOME INDIAN VENOMOUS SNAKES. 31

paralysis, so that, if a limb has been thus wounded, it is dragged

by the animal. The paralysis afterwards extends to the rest of
the body, and death ensues. Experiments on this subject have
been recorded by one of us (Dr. Fayrer) in the ThaiiatopJiidia
of India.
Effect of Serpent-venom on SnaJiCS.
The bite of venomous serpents, such as the cobra, Bahoia,
and Bnngarus, generally proves fatal to innocuous serpents,
but not always. The occasional escape of the latter is probably
due to the quantity of poison absorbed having been small,
either absolutely, or relatively to the size of the bitten snake.
The effect of the size of the innocuous snake upon the time
required by the poison to produce a fatal effect is illustrated by
Experiment/, in which a small rat-snake was killed by the bite
of a Btingarus ccerideus (less poisonous than a cobra) in 7 hours
17 minutes, while a large snake of the same species was not
killed by the bite of a cobra till after about 36 hours (Experi-
ment a); and another still larger one was unaffected by the
cobra-venom (Experiment g). Venomous snakes are not
generally affected either by their own poison or that of another
sort of snake, no less than 15 drops of venom having been
injected hypodermically into a cobra (Experiment r) without
effect ; but small ones are occasionally killed by large indi-
viduals belonging either to the same or to a different species.*
The symptoms caused by the poison were the same in both
the innocuous and the venomous snakes killed by it, and
consisted chiefly of sluggishness and indisposition to move,
which probably signifies in the snake, as it does in the frog,
a progressive paralysis. Only in Experiment l were convulsive
movements noticed. The movements of the tail in Experi-
ment c,after motion had ceased in every other part of the body,
are remarkable.
The poisonous action of the venom of the cobra, Dahoia, and
Bungarns upon innocuous snakes is shown in the following

* It is probable death may be due to other causes, especially in the case of a

Daboia-hitQ, where the fangs are so large that the wound and internal haemor-
rhage might cause death.
82 ON THE NATURE AND ACTION OF THE

experiments selected from a number recorded in tlie Thanato-

phidia of India :—
Experiment a. — March 10th, 1868. — A rat-snake (Ftyas
mucosa), about 6 feet in length, was bitten by a large cobra at
12.54. Before closing the snake's jaws on the part the scales
were scraped off. Blood was freely drawn by the snake's fangs
from bites inflicted in two places.
1.8 P.M. Appears sluggish ; wound bleeding freely.
1.16. Perfectly active, and moves about rapidly in the cage.
1.35. No change.
There was no apparent change in the snake all that day or
the next, except that it may have been a little more sluggish.
It died in the night of the 11th, being found dead on the
morning of the 12tli.
Experiment h.— A small grass-snake (Tropidomhcs quincun-
ciatus) was bitten by a cobra at 1.12 p.m.
1.11. Very sluggish.
1.20. Tosses its head about in a convulsive manner.
1.25. Dead 13 minutes after the bite.

Experiment c. — Two tree-snakes (Bendrophis picta), one about

3 feet 4 inches long, and the other somewhat smaller, were
bitten by a cobra.
1.7. The larger snake bitten.
1.8. The smaller one bitten.
1.12. Both sluggish.
1.15. The smaller snake dead 7 minutes after the bite.
1.16. The larger one dead 9 minutes after the bite.
They simply seemed to become sluggish and powerless ; there
were no convulsions, no writhings or contortions. After they
had appeared quite dead for a moment or two, the tail of each
moved slightly.
Experiment d. — A green whip-snake (Fasserita myderizans)
more than 3 feet long, was bitten by a cobra about 10 inches
from the head, at 12.37 p.m.
12.38. Sluggish, moves less actively ; gapes, keeping the
mouth wide open.
 POISON OF SOME INDIAN VENOMOUS SNAKES. 33

12.39. Almost paralysed ; mouth now closed ; head lying on

the side. The body is swollen where bitten.
12.43. Dead 7 minutes after the bite. This snake was
peculiarly active and vigorous though innocuous.

Experiment e. — A green whip-snake {Passerita mycterizaTis),

somewhat smaller than the former one, was bitten in the body
by a Dahoia at 1.40.
1.45. Almost powerless. It gradually became more and
more exhausted, gaped like the one bitten by the cobra, and at
2.2 it was dead, 22 minutes after the bite. The Daboia had
been in confinement for some time and was probably exhausted.
Experiment /.— A small rat-snake {Ptyas mucosa), about
2 feet long, was bitten by a Bungarus cceruleus, 42 J inches long,
in the muscles of the back at 1.8 p.m. ; blood drawn.
2.30. Sluggish ; has lost all its vivacity.
8.25. Found dead 7 hours and 17 minutes after the bite.

The occasional escape of an innocuous snake after the bite of

a poisonous one is illustrated by Experiment g. Several others
were made with a like result.

Experiment g. — A full-grown rat-snake {Ptyas mucosa), about

8 feet long, was bitten by a fresh cobra about two-thirds grown
and about half its own size. About 13 minutes after the bite
it seemed restless and uneasy, but remained perfectly active,
and was perfectly well on the third day after the bite.

The power of one venomous snake to kill another appears

from the following experiments :—
Experiment h, — A Bungarus fasciatus, nearly full grown, was
bitten by a very large and powerful cobra, 5 feet 8 inches in
length. It was bitten twice, about 8 inches from the head,
at 12.22 P.M. The cobra took firm hold and implanted the
fangs deeply. It seemed to be unaffected ; and 22-|- hours after
the bite it still seemed well ; but it died about the 29th hour.

Experiment i.— A Bungarus cceruleus, 28 inches long, was

bitten by a very large and powerful cobra. It died in 40 minutes,
(95) D
34 ON THEl NATURE AND ACTION OF THE

presenting the same symptoms as those of an innocuous snake

killed by a cobra-bite.
Experiment j.— A young and very small, though lively,
cobra, 14 inches long, was bitten in the muscular part of the
body by a large krail {Bimgarus cceruleus), 48 inches long,
at 12.50.
At 1 P.M. the cobra is very sluggish.
1.8. So sluggish that it moves with difficulty and can be easily
handled ; it makes no effort at resistance.
1.20. Apparently dying ; movements scarcely perceptible.
1.22. Dead 32 minutes after the bite.

Experiment k. — July lOth, 1869. A young cobra, about

10 inches long, was bitten at 3.45 p.m. by a fresh full-grown
cobra near the tail, so that the viscera might not be injured.
The fangs were seen to penetrate ; and no doubt could exist
that the poison was fairly inserted. Being put on the ground,
it crawled away vigorously, and seemed unaffected by the bite.
On the 13th it seemed well ; but on the 17th it was found dead,
and had apparently been so for about 12 hours.
As this snake was young it may have died partly from want
of food and partly from the wound, as well as from the effects
of the poison.

Though small snakes of a venomous species may be killed by

large ones, either of the same or of another species, full-grown
individuals are rarely injured by the bite of another, either of
their own or another species. This is illustrated by the follow-
ing experiments, which are taken from numerous others of the
same sort.

Experiment /.— A Bimgarus fasciatus was fairly and deeply

bitten by a fresh cobra near the tail ; there was no doubt of the
penetration of the fangs and inoculation of the poison. No
effect was produced, and the Bungarus was alive and well five
days after the bite.

Experiment m. — A Bimgarus fasciatus was thoroughly bitten

in a similar manner by a fresh Daboia. The bite produced no
 POISON OF SOME INDIAN VENOMOUS SNAKES. 35

effect, and live days afterwards the snake was in its normal
condition.

Experiment n. — A Dahoia was bitten by a fresh cobra near

the tail, the scales having been previously scraped off. The
snake bit fiercely and repeatedly. Two days afterwards no
effect could be noticed.

Experiment o. — A large black cobra was bitten in two places

1 foot 6 inches from the head, and also on the head, by a large
and vicious Daboia. Blood was slightly drawn ; and there could
be no doubt that the fangs had penetrated and the poison been
inoculated. Six days after the bite there was no change in the
snake.

Experiment p. — A full-grown cobra was bitten by another

full-grown, fresh, and vigorous cobra in two places about
6 inches from the head, and also in the mouth. They both bit
each other freely in this situation, and blood was freely drawn.
They were both well a week afterwards.
Experiment q. — A cobra had 15 drops of his own venom
injected hypodermically about 8 inches from the head. • A week
afterwards it seemed sluggish ; but this might be from other
causes.

Experiment r. — A cobra had 15 drops of the venom from

another cobra injected hypodermically in the same situation as
the last. A week afterwards he was perfectly well.

Effects on Fish.
Cobra-poison seems to produce paralysis, indicated by the fish
turning on its side in the water — and also great excitement, the
fish struggling and plunging violently.

Experiment IX.
A fish (Ophiocephahcs marulius), about 10 inches in length,
was bitten by a fresh cobra at 11.20 a.m. in two places on the
dorsal and ventral surfaces.
11.22.- It turned over on its side in the water.
11.23. Struggling and plunging violently in the water.
(95) D 2
36 ON THE NATURE AND ACTION OF THE

11.25. Turned over on its side.

11.26. On being roused it plunges violently.
11.40. Dead in 20 minutes from the bite.
For the purpose of comparison the following experiment with
curare was made. It will be seen that there was no plunging.
The failure of muscular action, except when a more than
ordinarily powerful stimulus from the nerve-centres called it
into play, is very evident.

Experiment X.
November, 1873. — Injected a solution of curare under the
skin of a carp near the tail. A great part of the solution came
out on withdrawing the needle of the syringe.
11.25. Injection made.
11.26. The fish lies obliquely in the water, inclining to the
opposite side from the injection. It can move when irritated,
and can remain perfectly upright in the water ; but in a very
short time its position becomes oblique again.
1 1.35. Injected some more curare. A great part of this also
returned.
11.50. Lies obliquely, but can move tolerably vigorously
when roused.
11.55. Moves more feebly when roused.
12.10. Seemed dead, but did not lie fiat on its side, and
still preserved the oblique position.
12.20. It suddenly started up without any apparent cause,
swam across the vessel, a distance of several inches, and then
relapsed into its former state.

Action on Snails.
Cobra-venom seems to destroy their irritability. It first
causes them to shrink within their shells, and finally lessens
their movements when stimulated.

Effect of Reagents, etc., on the Action of the Poison,

The activity of the poison is not destroyed, and scarcely
impaired, by drying. We have made no comparative experi-
 POISON OF SOME INDIAN VENOMOUS SNAKES. 37

ments with perfectly fresli poison and the dried residue of

a similar quantity; but there are few, if any, instances on
record of death from the fresh poison in less than half a minute,
the time in which the dried poison killed a guinea-pig in
Experiment XXVIII.
The local action of the poison, however, seems to be altered
by drying ; for extravasation of blood around the part where
a snake has inserted its fangs, or venom has been injected, is
one of the most prominent effects produced by the fresh poison,
whereas it is very slight, or absent altogether, when the dried
venom has been employed, except in occasional instances, such
as Experiment LYII.
Dilution seems also to have no effect in lessening the activity
of the venom, except so far as it retards absorption ; for it is
evident that a drop of pure poison, injected subcutaneously, is
likely to find its way into the circulation more quickly than the
same quantity diluted with a hundred times its bulk of water.
Coagulation of the venom by alcohol does not destroy its
activity, as we have shown in our former communication. The
coagulum thrown down by the alcohol is innocuous, or nearly
so ; but the poisonous principle remains in solution, and the
alcoholic extract possesses similar properties to tlie poison
itself. A specimen of poison was received from India in a
coagulated state ; but we are uncertain whether this occurred
spontaneously or was produced by the action of reagents. It is
probable, however, that it was due to its having been mixed, in
order to preserve it, with alcohol, which had evaporated before
we received it. It was active, as Experiment XI shows.
Coagulation by boiling does not destroy the activity of the
poison (Experiment XII) ; but a portion which was boiled for
more than half an hour under pressure corresponding to a
temperature of 102° C, had no effect when injected under the
thigh of a lark. The notes of this experiment have unfor-
tunately been lost. Admixture with liquor ammonise and liquor
potassse does not alter the effects of the poison. This appears
from Experiment XIII, and from several made by Dr. Fayrer
in India.
38 ON THE NATURE AND ACTION OF THE

Experiment XL
October 28th, 1872. — A fresh supply of poison was received
from India. It was of a yellowish colour, and was hard and
dry, like tough cheese. About half a grain diluted with alcohol
(in which it was only imperfectly soluble), was injected into the
thigh of the same guinea-pig at 4h. 14' 30".
4.15. Twitchings of an emprosthotonic character. The animal
is apparently attempting to vomit.
4.20. The twitchings continue. The animal throws up its
head. It seems sluggish, and will not walk.
4.22. A mixture of 5 minims of liquor ammonise with 10 of
water was injected into the animal. Almost immediately after-
wards itbecame convulsed, and fell over on its side, paralysed.
4.25. It is dying.
4.26. Quite dead.
4.27. The cardiac pulsations and peristaltic action of the
bowels still continue. The blood, when collected in a vessel,
formed a firm coagulum.
4.32. Peristaltic action diminished. The muscles of the leg
contract when the sciatic nerve is stimulated by an induced
current. Electrodes were then placed in the cord. The muscles
of the legs contracted readily when an induced current was
passed through the cord. One cell was employed, and the
distance of the secondary from the primary coil was 44 cm.

Experiment XII.
May 19th. — A full dose of dried cobra-poison was diluted
with distilled water, and heated until it was filled with white
flocculent coagula.
The solution was injected into a guinea-pig's hip at 3.25.
Twitching began almost immediately.
3.30. Eestless. Hind leg paralysed.
4. Twitching acute in hind leg.
4.10. Active hip-twiching, but hind leg still paralysed.
4.15. Making efforts to vomit.
4.25. Vomiting repeatedly.
4.30. Distinct repeated convulsive attempts to vomit. Limbs
 POISON OF SOME INDIAN VENOMOUS SNAKES. 39

becoming weaker ; began to be convulsed ; gradually becoming

more and more paralysed.
4.45. In convulsions. Dead.

Experiment XIIT.
May 19th. — Dried cobra-poison, dissolved in liquor ammonise,
injected into a guinea-pig's hip at 3.42.
Twitching at 3.43. Eestless.
4. Twitching ; restless ; weak in hind leg.
4.8. A little more injected with a full quantity of ammonia.
The guinea-pig becomes immediately very restless.
4.15. Paralysed. Going into convulsions. Pinching foot at
once causes reflex action; marked reflex actions all over the
body.
4.20. Nearly dead. Heart disturbed ; continued to beat
regularly for some minutes after death. Lungs much congested.

Influence of Constitution on the Action of the Poison. Su2)posed

immunity of the Mongoose.
With cobra-venom, as with other poisons, there is a general
correspondence between the size of the animal and the intensity
of the effects of a given quantity of poison, a small animal being
more readily afiected by it than a large one. There are, how-
ever, some exceptions to this rule; for a cat will resist the
action of cobra-poison as much as, or more than, a dog five or
six times its size. (Compare Experiment LVII with Experi-
ment XLIV.)
The mongoose {Herpestes griseus) has long been supposed to
be unaffected by the poison of venomous snakes, either on
account of some peculiarity in the constitution of the animal,
or, as the story used to run, on account of its knowledge of some
herb which it used to eat as an antidote ; but such is not the
case. If fairly bitten, it succumbs like any other creature, as
proved by experiments in India {Thanatophidia, pp. 68, 69, and
134). Its great activity and vigour enable it to elude the
snake ; and generally, when it is wounded, it is merely
40 ON THE NATURE AND ACTION OF THE

scratched, not pierced by the fangs. If the poison is inoculated,

it dies.
The same is true of the pig, which escapes probably by
receiving the wound in the foot, where absorption is not rapid
or vigorous. This animal, like others, yields to the poison
when the fangs are embedded and the virus thoroughly
inoculated {vide Thanatojphidia, p. 134).

Action on Germination.

In order to see whether cobra-poison had any effect on the

germination of seeds, the following experiments were made. It
will be seen from them that the venom does not prevent
germination, but interferes with it, especially when strong. In
this it agrees with rattlesnake-poison. (Weir-Mitchell On
Rattlesnake Venom, p. 52.)

Experiment XIV.
A piece of flannel was doubled, and, 12 cress-seeds being laid
between the folds, it was placed in a small beaker with 10 c.c. of
water. Another piece, treated in the same way, was laid in
9 c.c. of water and 1 of a 2-per-cent. solution of dried cobra-
poison.
Some time after the water had evaporated, so as to leave the
flannel soaked with water but not covered, nine of those seeds
which had been treated with water and poison had germinated
and grown to about half-an-incii in length, while seven of those
treated by water alone had germinated and had grown somewhat
larger than the others.

Experiment XV.
The preceding experiment was repeated with lettuce-seeds.
Seven of those treated with water alone had germinated, but
only one of those treated with water and poison.

Experiment XVI.
A small piece of cotton- wool was placed in the bottom of
each of two short test-tubes, and 10 lettuce- and 10 cress-seeds
 POISON OF SOME INDIAN VENOMOUS SNAKES. 41

were dropped into each. Ten drops of a solution of dried cobra-

poison, containing 0'0355 gramme in 3 c.c. of water, were then
used to moisten those in one tube, and as nearly as possible the
same quantity of pure water for those in the other. The seeds
were then covered with a few fibres of cotton- wool; the tubes
were stopped with a plug of the same substance, and placed in
a warm room.
Three days afterwards, all the cress-seeds which had been
moistened with water had sprouted and sent out a radicle,
varying from |- to ^ an inch in length. Eight out of the
10 lettuce-seeds had sprouted and sent out a radicle more than
^ of an inch long. All the cress- seeds moistened with poison
had also sprouted, but the radicles were only about yV of an
inch long. Five lettuce-seeds had begun to sprout, but the
radicles were barely visible.
It is not improbable that the delay caused by the poison in
the germination of the seeds, in this experiment, is not to be
attributed entirely to its poisonous action ; and it may be due
in great measure to the solution of the poison having matted
the fibres of cotton- wool more closely than the water, and tlius
rendered the conditions of air and moisture less favourable to
the seeds placed in it.

Uffed of the Poison when introdticed through different channels.

The action of the poison is most rapid when it is introduced
directly into the circulation, as by injection into the jugular
vein ; and in such instances death may occur in less than a
minute. When injected into the thoracic cavity, as in
Experiment XXVIII, death occurred almost as quickly; but
this ma,y have been due to puncture of the lung and intro-
duction ofthe poison directly into some of the pulmonary vessels.
Injection into the peritoneal cavity comes next in order of
rapidity, but a good deal behind the last ; and it is followed by
subcutaneous injection.
Whatever may be the effect of the venom of the viper or
crotalus, the cobra-virus produces its poisonous effects tolerably
rapidly when swallowed, both in the frog and in warm-blooded
animals, as is seen from Experiments XVII and XIX.
42 ON THE NATURE AND ACTION OF THE

It is also absorbed from the conjunctiva, and produces the

characteristic symptoms of poisoning. In Experiment XX the
animal, though affected by the poison, recovered; but in
several experiments made by one of us in India, death rapidly
occurred after the application of the fresh poison to the
conjunctiva {TlmnatoiMdia of India, pp. 108, 115, 127, 128,
135).
Experiment XVII.
May 21st, 1873. — 2.23 p.m. A small bit of dried cobra-poison
put into a frog's mouth and swallowed.
3.25. Frog not much, if at all, affected.
4.5. Frog not so vigorous. Appears to be paralysed in fore
legs, but moves his hind legs freely. On irritating his fore legs
there are vigorous contractions in his hind legs, but none in the
fore legs.
4.10. The anterior part of the body and fore legs seem to be
quite paralysed. No reaction is noticed in the eyelids when
the cornea is irritated. Hind legs are still vigorous.
4.20. Hind legs vigorous. All the fore part of the body quite
paralysed. Mouth gaping. Tongue swollen.
4.25. Hind legs now becoming weaker.
4.30. The application of acid causes slight reflex movements
in the hind legs.
4.35. Acid causes no reflex action. Complete paralysis and
death have thus occurred in two hours and a quarter.
4.40. Thorax opened. Heart still contracting rhythmically
and steadily.
4.55. Heart still contracting, but less vigorously. There is
no movement apparent in the intestines.
5.5. Heart still contracting slowly.
5.25. Heart still contracting. The heart and liver were now
removed and given to another frog.

Experiment XVIII.
The heart and liver of the former frog were given to a large
and strong frog. It was kept under observation for many days,
but did not seem in the least affected.
 POISON OF SOME INDIAN YENOxMOUS SNAKES. 43

Experiment XIX.
A small quantity of dried cobra-poison dissolved in water
was given to a young rabbit at 2.53 p.m. It was readily
swallowed. In seven minutes all the symptoms of poisoning
were developed. The rabbit died in convulsions in 11 minutes,
just as when the poison is injected hypodermically. The thorax
was opened a few minutes afterwards. The heart had ceased to
beat. Rigor Qiioriis came on very rapidly.

Experiment XX.
November 28th, 1872. — 1.49. One-quarter of a drop of cobra-
poison put into a guinea-pig's eye.
3.12. The eye is much congested. The animal has twitchings.
3.14. Has been making efTorts to vomit, and now vomits
frothy clear fluid. Has been purged also.
4.5. Still retching, but not vomiting.
November 29th. — Found to have recovered.

Local Action of the Poison,

Cobra-poison acts as a local irritant, and produces chemosis
of the conjunctiva and swelling of the eyelids when applied to
the eye, and occasionally congestion of the peritoneal vessels
when injected into the abdominal cavity (Experiments XX and
XLIV).
It paralyses the ends of the motor nerves, and also the
muscles of the part into which it has been injected (Experi-
ment XXV). The muscles are not only deprived of their
irritability, but become prone to putrefy (Experiment LVII).
The fresh cobra-poison produces great extravasation of blood
aromid the wound through which it has been introduced ; but
this is not so marked when dried poison is used.
If death do not rapidly follow, great swelling from infiltration
of the areolar tissue may occur, or, in some cases, gangrene of
the skin and subjacent cellular tissue and subsequent changes
indicative of general blood-poisoning.
The local action of viperine is probably more active than that
of colubrine virus.
44 . ON THE NATURE AND ACTION OF THE

Action of Cohra-poison ujooii the Blood.

The blood of animals killed by cobra-poison generally
presents a dark colour, as death is due to failure of the
respiration and not of the circulation ; but it readily assumes a
florid colour when exposed to air. The same is the case with
the blood of animals poisoned by Dahoia-venom (Experiments II,
V, and Vt).
Coagulation usually occurs readily and firmly in the blood of
animals killed by cobra-poison, while it is frequently absent
from the blood of those killed by that of the Daboia. In
experiments made in India, this occurred almost invariably :
and it is illustrated by Experiments II and IV. In Experi-
ments I,V, and VI, however, coagulation occurred in the blood
of a pigeon and guinea-pig poisoned by Daboia-Y QMom. ; and a
similar occurrence has been sometimes observed by one of us
(Dr. Fayrer) in fowls bitten by this snake in India.*
In numerous instances we have been unable to detect any
alteration in the blood-corpuscles after death from cobra-poison ;
but in Experiments XXI and XXII we observed a most
distinct crenation in the corpuscles of rats poisoned by it. This
was probably due in some degree to evaporation, as in
Experiment XXI it was to a great extent prevented by
surrounding the preparation with oil ; but it indicates a change
in the blood, as the corpuscles did not present this appearance
before the injection of the poison — although they were prepared
for observation in exactly the same way, and were as much
exposed to evaporation in the one case as in the other.

Experiment XXI.
A drop of blood from the tail of a white rat was exanained
microscopically. The corpuscles did not form rouleaux; but no
trace of crenation could be observed in them.
12.10. P.M. 0*018 gramme of dried cobra-poison, dissolved in
1 c.c. of water, was injected into the flank. Almost immediately
the nose of the animal began to twitch up every few seconds.

* Thanatophidia of India, pp. 80, 100, 101, 101. Vide Mr. Cunningliain's
remarks.
 POISON OF SOME INDIAN VENOMOUS SNAKES. 45

12.15. Head lias sunk down. The breathing was laboured.

The animal made a sudden start forwards. The hind legs
dragged behind. It did not move readily when irritated. The
breathing was laboured ; the expiration convulsive. General
convulsive movements occurred.
12.18. The animal seemed dead. The heart was still beating.
A drop of blood was taken from the tail ; and, the thorax being
opened, another was taken from the right ventricle. On being
examined microscopically, the corpuscles in both w^ere seen to
be very much crenated. They did not form rouleaux. Another
drop was taken from the right ventricle, and surrounded with
oil to prevent evaporation. Hardly a trace of crenation could
be observed in this drop ; but several branching crystals of a
reddish colour were observed, and some of them appeared to
grow while under observation. Numerous granular masses
were also seen.

Experiment XXII.
August 27th. — Injected 1 c.c. of a 2-per-cent. solution of
cobra-poison under the skin of the hip of a white rat.
1.35. Injection made.
1.37. Eespiration quick. The end of the tail snipped off, and
a drop of blood examined by Dr. Klein. The red corpuscles
are much crenated, and have no tendency to form rouleaux, but
adhere together in flat masses. The plasma contains numerous
lumps of a granular material, probably coagula of some sort.
2.5. The animal lies stretched out. Makes a curious squeak-
ing noise. It does not rise when the tail is pinched.
2.13. Lies with nose on ground. Convulsive movements of
hind legs.
2.15. Head sinks to one side. Convulsive movements.
2.18. Breathing slow. Marked interval between inspiration
and expiration.
2.19. Stopped breathing. Heart still beating.
2.20. The animal lay on its back. A few weak respirations
were made, and then ceased. The heart was beating steadily.
Thorax opened and heart exposed. A little blood drawn from
the ventricles by a fine pipette was examined microscopically
46 ON THE NATURE AND ACTION OF THE

by Dr. Klein. It presented exactly the same characters as

those of the former specimen. Blood from another healthy rat
showed numerous rouleaux, and the corpuscles were not
crenated.
Action on Muscles.

Cobra-poison has the power of destroying the irritability of

voluntary muscular fibre when applied directly to it, either in
a concentrated or diluted condition. It does not produce any
quivering of the fibres ; and in this particular it differs from
the poison of the rattlesnake as described by Dr. Weir
Mitchell.
The local action of cobra-poison on muscle is illustrated by
Experiments XXIII, XXIV, XXV, and XXVI.

Experiment XXIII.

September 4th. — A frog was decapitated, and the skin

removed from both hind legs. A longitudinal cut was then
made in the muscle of both thighs. A strong solution of dried
cobra-poison in distilled water, of such a strength as to
resemble the fresh poison closely in appearance, was then
applied to the cut in one thigh, while the other was moistened
with distilled water. Immediately after the application an
almost imperceptible trembling in the muscles occurred equally
in both thighs ; but it ceased after a few^ seconds, and did not
reappear. On testing the muscles soon afterwards, by an
induced current applied directly to them, those of the poisoned
leg contracted feebly, but those of the non-poisoned leg,
forcibly.
In this experiment, the quivering occurred equally in both
thighs, and was therefore obviously due to the water in which
the poison was dissolved, and not to the poison itself. '
As Weir Mitchell found that the quivering produced by the
poison of the rattlesnake was not prevented by paralysis of the
motor nerves by curare, the previous experiment was repeated
on a curarised frog.
 POISOK OF SOME INDIAN VENOMOUS SNAKES. 47

Experiment XXIV.
September 4th. — The motor nerves having been tested and
found to be completely paralysed, a strong solution of cobra-
poison was applied to a cut in the back of the right thigh. No
quivering of the muscles could be observed after its application.
The poison was only applied to the middle of the back of the
right
Do thigh. After a few minutes, those muscles with which it
had come into contact did not contract when irritated by the
direct application of an induced current. Distance of secondary
from the primary coil 0. The muscles of the sides and front of
the poisoned thigh, as well as those of the other thigh, con-
tracted well when irritated in the same way, with the coil at
13 cm.
The poison paralyses the muscles of warm-blooded animals in
much the same way as those of frogs ; and it seems probable
from the following experiment, that the paralysis of the
wounded limb, which is very frequently noticed in cases of
snake-bite, is partly due to the local action of the poison upon
the muscles.

Experiment XXV.
September 4th. — Injected five or six drops of a strong but not-
perfectly concentrated solution of dried cobra-poison into the
muscles of the left thigh of a guinea-pig.
12.43 P.M. Injection made. The animal immediately became
much excited, and rushed about wildly, crying loudly.
12.47. The leg seemed paralysed and dragged behind the
animal.
12.48. It ground its teeth and cried.
12.50. Began to start, and cried more loudly. Took it in my
arms. It then became quiet.
12.52. Shivered.
12.58. Laid the guinea-pig on its side on the table. It lay
still and did not attempt to rise. Eespiration was still going on.
12.59. Cut off the head of this guinea-pig (No. 1), and imme-
diately after decapitated another healthy guinea-pig of nearly
the same size (No. 2).
48 ON THE NATUllE AND ACTION OF THE

1.7. Exposed both sciatics of No. 1, and irritated tlifein by an

induced current.
Left leg. Coil at 0. No contraction.
Right leg. Coil at 17*5. Movement of toes.
The muscles of both legs twitch well when irritated by single
shocks (coil at 17*5), except those in the middle of the inside
of the left thigh, near the place to which the point of the
syringe had penetrated. These muscles contract when the coil
is at 3.
1.13. The muscles of the hip of No. 2 twitch distinctly when
irritated by single shocks, coil at 24.
The toes move distinctly when the sciatic is irritated ; coil
at 37.
1.15. The ventricles of the heart of No. 1 are firmly con-
tracted and motionless. The auricles are still pulsating
vigorously.
The ventricles of the heart of No. 2 are only moderately
contracted, and there is no pulsation either in them or the
auricles.
1.22. The toes of the right leg of No. 1 move when the
sciatic is irritated, coil at 18.
Those of No. 2 do so, coil at 37.
Put the electrodes in the cervical part of the spinal cord of
both guinea-pigs, and irritated it by an induced current, coil
at 0. No contraction took place in the hind legs of either
animal. Contractions occurred in the muscles of the fore legs
with much the same force in both.
1.45. On irritating the muscles by single induced shocks :—
left leg of No. 1, vastus externus contracts, coil at 9*5 ; rectus
femoris, a pale muscle, 12*5.
No. 1. Eight leg, vastus 15'5, rectus 25. No. 2. Eight leg,
vastus 11, rectus 15.
1.53, No. 1. Left leg, vastus at 16 ; right leg, vastus at 20.
No. 2. Left leg, vastus at 20 ; right leg, vastus at 20. The
vastus contracts rather more strongly in the right leg of No. 1
than in those of No. 2.
2.23. No. 1. Left leg, vastus at 4; right leg, vastus at 11.
No. 2. Left leg, vastus at 11 ; right leg, vastus at 11.
 POISON OF SOME INDIAN VENOMOUS SNAKES. 49

This experiment shows that the venom paralyses the motor

nerves when applied to them locally, a strong current applied
to the sciatic causing no contraction in the lett leg of 'No. 1,
while a moderate one caused movement in the right foot, at
a time when the muscles of both were nearly equally irritable.
Its deleterious action on the muscles, when conveyed by the
blood, is also evident in the rapid loss of irritability after
death in both legs of No. 1 as compared with No. 2. The pale
muscles seemed to retain their irritability longer than those
having a deep colour.
The power of cobra-poison to paralyse muscle when applied
to it, even in a diluted condition, is shown by the following
experiment.
Experiment XXVI.

July 18th, 1873. — The legs of a large frog were cut off close
to the body, and the skin removed. Each was then placed in
a glass, and a sufficient quantity of fresh ox-blood serum
poured over it to cover it. In one glass, the serum contained
about 5 centigrams of cobra-poison dissolved in about 20 c.c. of
serum; but, with this exception, all the conditions under which
the two legs were placed were exactly alike.
July 19th. — About 19 hours after the immersion of the legs
in serum their irritability was examined.
The muscles of the leg in the pure serum did not contract at
all when the strongest irritation was applied to the sciatic
nerve, but contracted very vigorously when irritated directly.
The muscles of the leg in the poisoned serum were whiter than
those of the other one. They had a faint yellowish tinge, and
were somewhat stiff. They did not contract in the least when
the strongest irritation by a Du-bois coil was applied either to
them or the sciatic nerve.
When the poison is injected directly into the circulation, or
is very rapidly absorbed, so that the quantity circulating in the
blood is large, it ,destroys the irritability of the voluntary
muscles rapidly, and, occasionally at least, hastens in a most
remarkable manner the occurrence of rigor mortis. This is
well seen in the Experiment XXV, wiiere rigor mortis super-
(95) E
50 ON THE NATURE AND ACTION OF THE

vened in half an hour after the injection of the poison, while

the muscles of another animal killed at the same time by
decapitation retained their irritability for many hours.

Experiment XXVII.
May 8th, 1873. — Eight thigh of a frog ligatured, with the
exception of the sciatic nerve. Animal poisoned by the intro-
duction of some dried cobra-poison dissolved in water into the
lymph-sac of the back. After the animal had become com-
pletely paralysed, the gastrocnemii of the two legs were
irritated by an induced current (1 bichromate cell).
Left leg (poisoned), distance of coil 13'5, contraction ; right,
24*0, contraction.
Experiment XXVII (a).
Another frog prepared in the same way gave at first :— left leg
(poisoned), distance of coil 42*2, contraction ; right (ligatured),
distance 21*0, contraction.
After some time :— left leg, distance 6*0, contraction ; right,
distance 25-0, contraction.
Some time later :— left leg, distance 0, almost no contraction ;
right leg, distance 14'5, contraction.
In this experiment, the poisoned muscle at first responded
more readily to the irritation than the one which had been
deprived of blood by the application of a ligature ; and this
renders more apparent the effect of the poison, in causing rapid
diminution and final extinction of irritability in the muscle to
which it had access, since the other lost its excitability very
slowly.
Experiment XXVIII.
September 5th. — About 2.35 p.m. injected f c.c. of a 2-per-
cent, solution of dried cobra-poison into the thoracic cavity of
a guinea-pig. It was uncertain whether the lung (right one)
was pierced by the point of the needle or not. Within a few
seconds the animal gave several convulsive struggles, and died
in half a minute or so. The head was then cut off. Immedi-
ately afterwards a second guinea-pig was killed by decapitation.
 POISON OF SOME INDIAN VENOMOUS SNAKES. 51

On opening the thorax of No. 1 (the poisoned guinea-pig) the

lungs were found congested. The heart was tetanically con-
tracted and quite still. The heart of No. 2 was contracting
vigorously. The vena cava contained a few hubbies of air.
The lungs were pale.
2.40. Peristaltic movements are going on very actively in the
intestines of both animals.
2.42. The muscles of the abdominal wall irritated by single
induced shocks.

Guinea-pig, 'No. 1. No contraction. Coil at 0.

Guinea-pig, No. 2. Contraction. Coil at 14*5.
Muscles of the hip irritated in the same way :—

^ 1 /"Trace
L Contraction Coil 0. Coil 13.
still slight.of muscle.
of contraction
■XT n r Contraction. Coil 37.
^* ' LPowerfulkick. Coil 0.
2.50. Rigor mortis is coming on in No. 1. The legs are quite
stiff. A trace of peristaltic movement still going on in the
small intestine.
The muscles of No. 2 are quite flexible.
2.55. No. 1. Muscles of back of thigh and of abdominal wall
irritated directly as before. No contraction. Coil at 0.
Muscles of the front of thigh twitch slightly. Coil at 0.
No. 2. Muscles of back of thigh twitch decidedly. Coil
at 37.
3.12. No. 1. No contraction in any muscles. Coil at 0. The
animal is stiff.
No. 2. Muscles are quite limp. Muscles of back of thigh
twitch decidedly. Coil at 25.
All the muscles do not lose their irritability with the same
rapidity, some of them becoming paralysed before others. The
intercostal muscles, serrati, and abdominal muscles seem to lose
their irritability first ; and such muscles of the limbs as have a
dark colour become paralysed sooner than those which are paler
(Experiment XXV).

(95) E 2
52 ON THE NATURE AND ACTION OF THE

Experiment XXIX.
September 4th. — A cannula was placed in the carotid of a
large guinea-pig, and J c.c. of a 2-per-cent. solution of cobra-
poison injected into it towards the heart. The animal was
seized with violent convulsions, passing into complete opistho-
tonos in about 20 seconds after the injection of the poison.
These ceased, and the animal seemed quite dead in rather less
than a minute from the injection. The thorax was then opened.
The lungs were somewhat congested. The heart was quite still
in tetanic contraction. A strong interrupted current applied
to it caused no contraction of any of the fibres. The muscles
lost their irritability very quickly ; the intercostals of both sides,
and the serratus and subscapularis of the right side, seemed to
lose their irritability before the other muscles.
When the poison is more slowly absorbed, so that a less
quantity of it circulates in the blood, its action on the muscles
is much less marked, as is evident from a comparison of the
irritability of those in the poisoned and non-poisoned limbs in
Experiments XXXYIT, XXXVIII, XXXIX, XLVII. If the
poison has undergone such changes as render it less active, it
has no action, or only a feeble one, on the muscles, as seen in
Experiments XI, XXX, XXXI, and XXXII, where poison,
which had undergone partial coagulation was employed.

Experiment XXX.
January 14th. — In order to test the local action of the poison
on the muscles and nerves, a ligature was tied round the base
of a frog's heart so as entirely to arrest the circulation.
12.0. About a drop of cobra-poison was injected into one leg.
1.30. Laid bare the lumbar nerves in the abdomen, and
irritated them by an induced current. Both legs contracted
nearly equally.

Experiment XXXI.
January 14th. — At 12.15. One or two drops of cobra-poison
were injected into the leg of a frog. The wound bled freely.
 POISON OF SOxME INDIAN VENOMOUS SNAKES. 53

Immediately after the injection the frog became very excited

and jumped about very much.
12.20. Frog quiet. Eespiration quick.
2.30. Frog quiet, but jumps when irritated. It seems to use
both legs equally well.
January 15th. — The frog is not dead, but is feeble. On
killing and opening it, both legs contracted nearly equally
when the lumbar nerves were stimulated by an induced
current.

Experiment XXXII.
January 15th. — Tied the heart of a frog, and, 12.55 p.m.,
injected into the right leg a drop of water, and into the left leg
a drop or two of cobra-poison.
1.55. — Irritated the back of the frog by an induced current.
Both legs contracted nearly equally.

Experiment XXXIII.

May 9th. — A frog {Rana temporaria) was poisoned with

curare. After complete paralysis had set in, the right leg was
ligatured, with the exception of the sciatic nerve. The animal
was then poisoned by the introduction of a solution of dried
cobra-poison in water into the lymph-sac on the back, at about
12.30 P.M. The irritability of the muscles was tested by single
induced currents applied to the denuded muscles, about 2.30.
Distance of Coil.

Left leg 7*5 Contraction.

Eight leg 7-2-7-5 do.

Another frog was curarised and similarly prepared, with this

exception — that the vessels of the right leg only were ligatured,
the muscles, as well as the nerve, being left free. This frog was
also examined in the same way ; and the irritability of the
muscles in both legs was found to be almost exactly the same
three to four hours after poisoning. Both contracted with the
coil at about 7'5.
54 ON THE NATURE AND ACTION OF THE

Secondary Action of the Poison on Muscles.

The muscles of the part into which the poison has been
introduced are very apt to undergo rapid decomposition. We
have already shown that their irritability is either lessened, or
completely destroyed, by the action of the venom ; and it seems
very probable that the mere contact of any other foreign body,
containing bacteria or their germs (as the water in which the
cobra-poison was dissolved in our experiments certainly did)
would suffice to explain the decomposition of the muscle without
assuming any special putrefactive action on the part of the
poison ; for the muscle, which has been at least temporarily
killed by the poison, is placed in the body in the most favour-
able conditions of temperature and moisture for the occurrence
of decomposition whenever any germs are brought into contact
with it. However, Weir Mitchell found that the venom of the
rattlesnake had a curious influence upon muscle, which could
hardly be explained without the supposition that the poison
had a peculiar disorganising action upon the muscular tissue.
In every instance the venom softened the muscle in proportion
to the length of time it remained in contact with it ; so that,
even after a few hours, in warm-blooded animals, and after a
rather longer time in the frog, the wounded muscle became
almost diffluent, and assumed a dark colour and somewhat jelly-
like appearance. The structure remained entire until it was
pressed upon or stretched, when it lost all regularity, and
offered, under the miser oscope, the appearance of a minute
granular mass. In order to ascertain whether cobra-poison had
a similar action, the following experiment was tried.

Experiment XXXIY.
September, 1873. — The gastrocnemii of a frog were removed
and laid in two watch-glasses. One was then covered with
several drops of a solution of dried cobra-poison, dissolved in a
sufficient quantity of |-per-cent. salt solution to form a mixture
about the consistence of fresh poison, while the otlier was
covered with a few drops of salt solution alone. They were
then protected from dust by two other watch-glasses inverted
 POISON OF SOME INDIAN VENOMOUS SNAKES. 55

over them. The temperature of the room was moderately warm.

The poisoned muscle underwent no change. Both muscles
gradually dried up ; but at no time could one be distinguished
from the other, except by the label on the watch-glass.
The influence of cobra-poison in causing decomposition within
the body is evident from the following experiment.

Experiment XXXV.
January 17th. — About three drops of cobra-poison were
injected under the skin of the flank of a guinea-pig at 12.48 p.m.
Immediately afterwards the guinea-pig became restless and
cried. In two minutes its head began to twitch. An hour after
the injection the animal was quiet, and little or no effect of the
poison could be observed. Three hours after the injection it
did not seem very well. Next morning it was found dead. On
examining it 22 hours after the injection it had begun to
undergo decomposition. The abdomen was somewhat inflated,
and sulphuretted hydrogen issued from it when opened. The
hair came off readily from all parts of the animal's skin. The
muscles were soft. There was little ecchymosis at the spot
where the injection had been made. The tissues near it were
rather watery. The heart was contracted ; the lungs somewhat
congested.
Action on the Nervous System.
The most prominent symptoms of an affection of the
nervous system after the bite of a cobra, or other venomous
snake, in animals or man, are depression, faintness, lethargy,
and in some cases, somnolence. There is loss of co-ordinating
power, and paralysis, sometimes affecting the hind legs first and
creeping over the body, sometimes affecting the whole body at
once. Death occurs by failure of the respiration, and is
preceded by convulsions.
These symptoms clearly point to paralysis either of the
nervous centres or of the peripheral nerves. It may be
supposed that the mention of the latter alternative is super-
fluous, and that paralysis of the peripheral nerves cannot
produce such symptoms, which must therefore, by exclusion, be
56 ON THE NATURE AND ACTION OF THE

due to an affection of the central ganglia. More especially may

the occurrence of convulsions be thought to exclude the
possibility of death being due to paralysis of the peripheral
terminations of motor nerves ; for if tlieir function is abolished
here, how, it may be said, can general convulsions, which have
their origin in the nervous centres, occur ?
The answer to this is, that although the ends of the motor
nerves are so far deadened that they no longer transmit to the
muscles any ordinary stimulus proceeding from the nerve-
centres, their function is not so thoroughly abolished that they
cannot transmit those which are stronger than usual. This is
shown by the fact that when an animal is slowly poisoned by
curare (as for example when that poison is introduced into the
stomach after ligature of the renal vessels), convulsions occur
just as in death from cobra-poison. Although the motor nerves
have their function so much impaired that they no longer
transmit to the muscles of respiration the ordinary stimuli from
the medulla, which usually keep up the movements of
breathing, they can still transmit those stronger impulses which
proceed from it when greatly stimulated by the increasing
venosity of the blood, and which cause the respiratory as well
as the other muscles of the body to participate in the general
convulsions. The loss of co-ordination which occurs in poisoning
by cobra-venom has also been noticed by Voisin and Liouville
in poisoning by curare.
That the peripheral terminations of the motor nerves are
actually paralysed by cobra-venom is shown by Experi-
ment XXXVI, in which the animal was able to move the leg
which had been protected from the action of the poison for
some time after the rest of the body was perfectly motionless,
as well as by Experiment XXXVII and those succeeding it.
Its occurrence in man is indicated by the symptoms of a case
described by Dr. Hilson {Ind. Med. Gaz., October, 1873, p. 254).
But paralysis of motor nerves is not the only effect of cobra-
poison on the nervous system. The spinal cord is also
paralysed, as is seen from Experiment XLI, where motion
ceased in the frog's leg which remained free from poison,
although it answered with great readiness to a very weak
 POISON OF SOME INDIAN VENOMOUS SNAKES. 57

stimulus applied to its nerve. In some instances paralysis of

the spinal cord appeared to cause death when little or no
affection of the motor nerves could be observed (Experi-
ment XLVII, etc.) ; but in others the peripheral paralysis was
strongly marked. In no case was it more obvious, and in few
was it so distinct as in Experiment XXXV I, made with the
virus itself, which had neither become coagulated nor dried.
In experiments made with the coagulated poison, death seemed
invariably to be caused by paralysis of the spinal cord, the
motor nerves being little affected (Experiment XI); while, in
those made with the dried venom, sometimes the action on the
cord predominated, and sometimes that on the nerves. In this
respect, as well as in some of the symptoms it produces, cobra-
poison agrees very closely with conia. This alkaloid, as Crum-
Brown and Eraser have shown, often contains a mixture of true
conia and methylconia. Conia alone paralyses the motor
nerves without affecting the spinal cord ; but when mixed with
methylconia, sometimes the one is affected first and sometimes
the other. When the dose is small, the motor nerves are
usually paralysed before the reflex function of tlie cord ; but
when the dose is large, the cord is paralysed before the nerves.
Methylconia also affects both ; but a small dose of it paralyses
the cord before the nerves, while a large one paralyses them
first. The paralysis of the hind legs, often observed in snake-
poisoning (Experiments VI and VII), is probably partly due to
the local action of the poison in the nerves and muscles of the
bitten member, and partly to its action on the cord. This
paralysis is noticed in Genesis xlix, 17, where Jacob says,
" Dan is an adder in the path, biting the horse-heels, so that
the rider falleth backward." In this point cobra-venom, when
dried, appears to resemble methylconia rather than its
admixture with conia ; but it exercises numerous other actions
upon the blood, muscles, etc., which neither of these substances
has been shown to do. It is doubtful whether the cerebrum is
directly affected by cobra-poison, as the intelligence both in
man and animals often remains almost unimpaired to the last,
and the stupor and drowsiness which are sometimes noticed
may be caused indirectly by the action of the venom on the
58 ON THE NATURE AND ACTION OF THE

motor and vaso-motor nerves and on the functions of the cord.

The reflex centres, through which irritation of the fifth nerve
acts, remain unaffected after the reflex function of the cord is
nearly gone ; and even then the power of voluntary motion
still exists.

The eff"ect of the poison upon the respiratory and vaso-motor

nerves will be considered under the heads of respiration and
circulation.

Action of Cohra-poisoyi on Motor Nerves.

As the contraction of a muscle, on irritation of the motor
nerve supplying it, is the index by which we judge of the
irritability of the nerve itself, the paralysing effect of cobra-
poison upon muscle renders the exact determination of its
action upon motor nerves much more difficult than in the case
of such a poison as curare, which leaves the muscular
irritability intact. For the failure of a muscle to contract on
irritation of its motor nerve can be due only to paralysis of the
motor nerve in the case of curare ; but in poisoning by cobra-
venom it may be due to enfeeblement of the muscles, as well
as paralysis of the nerve. But if we find instances in which
the muscles still retain their irritability almost unaltered, and
respond readily to direct stimulation after tliey ha\ e ceased to
contract on irritation of their motor nerve, we are justified in
saying that the nerve is paralysed; and such is the case in
Experiment XLI.
In Experiment XXV this action on the ends of motor nerves
is all the more evident from the paralysis being most complete
in the part where the poison was introduced. At this part, it
was brought, in a concentrated state, into contact with the ends
of the motor nerves, while the other parts of the body received
it after dilution with the blood ; and in them the paralysis was
much less marked.
The paralysis of the hind legs, so often noticed in experi-
ments, appears to be due, at least in considerable measure, to
the local action of the poison on the ends of the motor nerves
of the legs, as the injection or bite is often made on the flank
or thigh.
 POISON OF SOME INDIAN VENOMOUS SNAKES. 59

The action of the poison on motor nerves is illustrated by the

following experiments, performed by Bernard's method of
ligaturing one leg of a frog before poisorring it. The poison is
thus carried to every part of the body except the ligatured
limb, the motions of which indicate the state of the nerve-
centres after the other parts of the body have been paralysed.

Experiment XXXVI.
A ligature was placed round the right thigh of a young frog,
excluding the sciatic nerve.
2.42. A drop of dark fluid cobra-poison (the first supply) was
injected into the dorsal lymph-sac. Immediately after the
injection the animal became restless.
3.0. It lies quietly with its eyes shut. It hardly moves when
touched ; but it struggles when laid upon its back.
3*8. It can still drav^ up the ligatured leg. The other one
can be drawn up, but with a wriggling motion. When laid on
its back the animal no longer resists.
3.9.30. It lies quite flat. There is trembling of the leg when
either foot is touched ; and when it is pinched, either leg can
still be drawn up. On suddenly touching the poisoned leg, the
frog gave a jerk with both. Kespiratory movements have
ceased. The exact time when they did so was not noticed.
3.17. The frog has become much lighter in colour, with the
exception of the ligatured leg.
3.45. The eyes no longer shut when touched ; they remain
widely open. Dilute acetic acid of 1 per cent, produces no
effect when applied to the sound leg ; but when the leg is lifted
up, so as to prevent friction against the table, it is drawn in
towards the body.
4.9. On applying a strong interrupted current to the eye of
the frog the unpoisoned leg jerks feebly, the poisoned one not
at all.
4.13. On turning the frog on his back the non-poisoned leg
moved.
4.20. Opened abdomen. The heart was beating, but only
slowly. Irritated the lumbar nerves on the left side (those of
poisoned leg) by an interrupted current. ISTo contraction
60 ox THE NATURE AND ACTION OF THE

occurred in the poisoned leg ; but twitching took place in the

non-poisoned one. Irritated lumbar nerves of right side.
Tetanus occurred in the right (non-poisoned leg). No move-
ment of the poisoned leg. Laid bare the muscles of both legs,
and irritated them by a Faradic current directly applied.
Those of the poisoned leg were paler than those of the other.
The muscles of both legs contracted when irritated directly.
Exposed the sciatic nerves of both sides and irritated them by
an induced current. ]^o contraction in the gastrocnemius of
poisoned leg. Tetanus in the non-poisoned leg.
4.35. The heart is no longer contracting. Electrodes were
placed in the medulla, and an interrupted current applied.
Contractions occurred in the non-poisoned leg. No contractions
in the poisoned one.
The movements which occurred in the non-poisoned leg when
the lumbar nerves of the other side were irritated may have
been due to reflex action through the spinal cord. If this were
the case, it would indicate that the sensory fibres in the lumbar
plexus were not paralysed, and that the reflex power of the
cord was not quite destroyed; but the nerves were not very
carefully isolated, and it is probable that the twitchings were
due to direct irritation of the lumbar nerves of the right side
by conducted currents, especially as irritation of the left sciatic
nerve caused no movement in the right foot.
The continuance of movement in the ligatured leg, after it
had ceased in other parts of the body, indicates that the ends of
the motor nerves have been paralysed ; and this is confirmed
by the production of tetanus in the ligatured and absence of
movement in the poisoned leg when their motor nerves are
stimulated. The slightness of the movements in the ligatured
leg when a strong interrupted current was applied to the eye,
while the motor nerves of the limb still retained their irrita-
bility, indicates that paralysis of the reflex function of the cord
had taken place. The motion of the leg on turning the frog on
his back afterwards shows that the higher nervous centres,
through which the opposition to the change of posture was
manifested, retained their power longer than the cord.
 POISON OF SOME INDIAN VENOMOUS SNAKES. 61

Experiment XXXYII.
TsTovember 29th, 1872. — The sciatic nerve of the left leg of a
frog was exposed ; and a double ligature being passed under it
round the limb, the whole of the tissues except the bone were
then divided and removed between the ligatures. A fraction of
a drop of cobra-poison, diluted with |~-per-cent. salt solution,
was injected into the lymph-sac. After about two hours the
animal seemed paralysed. On irritating either fore leg by
electricity, or by acetic acid, slight movements occurred in the
hind feet, and were fully stronger in the poisoned than the
ligatured limb. Irritation of the poisoned hind foot also occa-
sioned twitches both in it and the non-poisoned foot. Twitches
did not invariably occur. No twitching of the fore paws was
noticed on irritation of the hind feet. A ligature was then
passed round the poisoned hind leg, and the tissues divided, as
in the non-poisoned one, and the animal left a little longer.
Irritations again applied had a similar result to the former, but
the contractions in the non-poisoned limb were sometimes
stronger than in the other. Irritation applied by a strong
interrupted current to the spinal cord, by electrodes inserted in
it, caused very faint twitches in both hind feet. Irritation of
the lumbar nerves in the abdomen caused very faint twitches in
the feet. Irritation of the exposed sciatic nerve of the non-
poisoned limb by an interrupted current caused strong con-
tractions. Similar irritation of the poisoned sciatic caused
much weaker contractions. Direct irritation of the muscles by
interrupting a constant current caused contractions of nearly
equal strength in both.
The dose of poison in this experiment was small, and it was
given in a much diluted form. The fact that an interrupted
current applied to the sciatic nerve of the poisoned limb had a
much slighter effect than the same current applied to the
sciatic nerve of the non-poisoned limb, while the interruptions
of a constant current by opening and shutting a key caused
the poisoned and unpoisoned muscles to contract with apparently
the same force, shows that a small dose of the poison causes a
considerable amount of paralysis of the ends of motor nerves,
while the muscles are but little affected.
62 ON THE NATURE AND ACTION OF THE

Experiment XXXVIIL
May 14tli. — The right leg of a frog was ligatured, with the
exception of the sciatic nerve, and the animal poisoned by a
rather small dose of dried cobra-poison dissolved in water, and
injected into the dorsal lymph-sac at 11.45 a.m.
12.15. The animal paralysed. Acetic acid applied to the left
arm caused movements in it ; but no movements ensued when
the acid was applied to the nose. When applied to both arms
and one leg, it caused movements in the arms and the left leg,
but none in the right leg.
12.33. Acetic acid applied to the left arm causes movement
in it, but in no other part of the body.
12.51. Electrodes were placed in the spine and the cord
irritated by a Faradic current. At 15 cm. distance of the
secondary from the primary coil there is faint twitch in right
arm. At 9, distinct twitch in both arras. At 0, distinct twitch
in both arms, none in legs ; sciatics exposed and irritated.
At 50, right leg contracts distinctly. At 36, right leg becomes
tetanised. At 16, left leg contracts very faintly indeed. At 8
left leg contracts slightly.
The muscles were then irritated by single induced shocks :—
9'8 cm., right leg faint contraction ; 9*8, left (poisoned) leg
contraction is equally strong ; 101, left (poisoned) leg contrac-
tion occurs. 10*1, right (ligatured one) does not contract.
In this experiment, the irritability of the poisoned muscle is
greater than that of the other, the venom having done less
injury to the muscular substance than the deprivation of blood
by the ligature, and consequently the paralysing action of the
poison on the ends of the motor nerves becomes very evident.

Experiment XXXIX.
May 12th, 1873. — A ligature was passed tightly round the
right thigh of a large frog, the sciatic nerve being excluded.
12. Eight leg ligatured.
12.12. Injected a considerable dose of a solution of dried
cobra-poison in water into dorsal lymph-sac.
12.14, The frog has assumed a most peculiar position. The
 POISON OF SOME INDIAN VENOMOUS SNAKES. 63

left hind leg is drawn up, and the two fore legs are held over
head with palms turned forwards.
12.20. Cornea sensible. Left leg is drawn up again if it be
forcibly extended.
12.31. Cornea sensible. When the left hind foot is pressed it
is drawn up very slowly with a wriggling motion. Pressure on
the right foot causes no movement whatever.
12.40. Acetic acid applied to a forearm causes vigorous
movement in it and also in left hind leg.
1. Acetic acid to right hind foot causes no movement.
Applied to left hind foot it causes vigorous movements in both
forearms and left hind leg.
1.12. A ligature was applied to the left thigh in a similar
manner to that on the right, so as to cut off the circulation in
the left leg also, and thus bring the two legs as much as possible
under the same conditions. The general condition of the frog is
much the same as before ; but the reflex action produced by
irritation of the cornea is slighter.
1.24. Acetic acid applied to right forearm. Slight movement
occurs in right hind leg alone. When applied to left forearm it
caused slight movement in that arm alone.
1.35. Acetic acid applied to both feet, both forearms, and to
the nose caused no motion anywhere. Both sciatic nerves were
now laid bare along a considerable portion of their course. It
was found that, although the right sciatic had not been included
in the ligature, it had been tightly constricted by the fascia at
the place of ligature. Sciatic nerves irritated by an induced
current.
Distance of
secondary from
Leg. primary coil.
Left. 0 No contraction of muscles of leg.
Eight. 32*0 Distinct contraction. As this might
possibly have been due to the left
sciatic being injured by the ligature
more than the right, both sciatics were
exposed further, and irritated quite
below the points of ligature and just
above the knee.
64 ox THE NATURE AND ACTION OF THE

Distance of
secondary from
Leg. primary coil.
Eight. 37'5 Distinct contraction.
Left. 7"0 No distinct contraction. When the
muscles were irritated by single in-
duced shocks, applied to them directly,
they contracted almost equally.
Single shock. 7'5
In this experiment, the right sciatic nerve had been injured
by the operation of ligaturing ; and thus the effect of the
poison on the other limb as compared with the right one was
less manifest. ^N'otwithstanding this it moved, and the other
limbs did not, when the right arm was irritated. The difference
between the irritability of the sciatic nerves when the muscles
of the legs themselves were almost equally irritable, shows, in a
marked manner, the influence of the poison on the motor
nerves.
Experiment XL.
May 14th. — Frog ligatured round the middle, excluding
lumbar nerves.
10.57. Ligature applied. A considerable quantity of blood
was lost.
10.58. A considerable quantity of dried cobra-poison dissolved
in water was injected into the dorsal lymph-sac. Immediately
after being released the frog jumped about, but became quiet
in a minute or so.
11.28. Made some voluntary movements.
11.45. Acetic acid to fore feet causes weak reflex movements
in both fore feet ; stronger in hind feet, especially in right.
11.55. Acetic acid to right forearm caused vigorous kicks of
right hind leg. Acetic acid affected right leg in 10 seconds.
No motion in any other part of body. Acetic acid to left
forearm caused kicks in both hind legs, but much more
vigorous in the right. Also movement of left forearm by
itself, but weak.
12.5. Acetic acid to left fore leg caused wriggling motion,
first in right hind leg and then in left fore leg in 16 seconds.
 POISON OF SOME INDIAN VE.\OMOUS SNAKES. 65

Applied to right forearm it caused a weak kick in right hind

leg and wriggling in left hind leg, but no motion in any other
part.
12.27. Acetic acid applied to forearm. Ko reflex action
anywhere.
12.30. No reflex action anywhere on application of acetic acid.
12.30. Distance of coil 8. Electrodes in the spinal cord.
Slight contractions in right hind and left fore legs, and also in
the abdominal muscles, though very weak. It was now
noticed that the cord with which the frog was attached to the
board had been very tightly tied round the left forearm and
left there. The circulation was stopped there, as the cord had
not been removed.
The paralysing effect of the poison on the motor nerves was
here shown by an involuntary experiment. On irritating the
cord the ligatured leg responded as we had expected, but we
were astonished to see movements in the left arm also. An
examination of the limb at once showed the cause of the
phenomenon. The cord attaching it to the board had been
inadvertently drawn so tight as to obstruct the circulation, and
thus prevented the access of the poison to the nerves.

Experiment XLI.
May 15th. — Right thigh of frog ligatured, with exception of
the sciatic nerve.
1.2. Ligature applied.
1.4. A considerable dose of dried cobra-poison dissolved in
water injected into dorsal lymph-sac.
2.26. Acetic acid applied to a limb causes no movement
whatever in 60 ^
Interrupted current. Distance 0, electrodes in spine ; only
weak twitch in muscles of forearms ; no movement in hind leg.
2.30. Both sciatics exposed.
Right sciatic. Distance 50, distinct contraction of gastro-
cnemius.
Left sciatic. Distance 0, no contraction of gastrocnemius.
Single shocks. Both gastrocnemii exposed and irritated
directly.
(,95) F
66 ON THE NATURE AND ACTION OF THE

Distance 9'5, very faint contraction in both tibial and

gastrocnemic muscles. Apparently equal in both legs. Heart
quite still and contracted.
On testing the irritability of several of the frogs used in
these experiments on the ensuing day, the ligatured leg was
found to contract on irritation of the sciatic nerve, or of the
muscles directly. The muscles of the poisoned leg did not
contract, either when irritated directly or through the nerves.

Experiment XLII.
May 20th, 1873. — In order to test the action of cobra-
poison on the ends of the motor nerves, without disturbing the
experiment by ligaturing one leg, two frogs were taken of as
nearly as possible the same size. Both were very small ; but
No. 1 was somewhat larger and stronger than No. 2. The
sciatic nerve was exposed in one thigh of each frog and pJaced
on the hook electrodes used by Marey for his myograph. By
means of a Pohl's commutator, with the cross pieces taken out,
an interrupted current could be sent at will through either
nerve. The distance of the secondary from the primary coil at
which the first faint contraction took place in the muscles of
either nerve was noted.
Distance of primary from
Time secondary coil.
(about). Frog 1. Frog 2.
1.25 17-7 22
1.40 26-3 12-3
1.46 26 18 Injected a solution of dried
cobra-poison in water into
dorsal lymph-sac of frog
No. 1.
2.7 31-2 24
2.27
31 18-5
2.50 24 17-8
3.10 17-5 19-2 Frog 1 moved the fore legs
when the coil was at such a
distance (19 ?) that no move-
ment occurred in leg when
nerve was irritated.
 POISON OF SOME INDIAN VENOMOUS SNAKES. 67

Distance of primary from

secondary coil.
Time. Frog 1. Frog 2.
3.30
12 17-5
3.40
4 10-5
10 33
37
15-5
4.17 9
4.30 11 18
At 37 voluntaiy moveiiient&'
37 occurred in legs of Frog 2.
4.50 8
4.55 ... ...
Thestroyed.
brains of both frogs de-'
4.58 7-5 16-5
May 21st. — The sciatics of the other legs were exposed and
irritated.

0 11*5 Frog 1, no contraction. Frog 2,.

slight contraction. The irri-
tability of the muscles was
now tested by single induced
shocks applied to them.
0 7*5 Frog 1, no contraction. Frog 2^
slight contraction.

The disturbing effects occasioned in the other experiments by

the necessity of comparing a limb acted on by the poison, but
retaining its blood-supply, with one in which the circulation
had been arrested, is here got rid of by employing two frogs of
as nearly as possible the same size. The paralysis of nerves
caused by the poison is evident.

Experiment XLIII.
December 4th. — Right leg ligatured, with the exception of
the sciatic nerve ; a small quantity of alcoholic extract of cobra-
poison dissolved in water injected into the dorsal lymph-sac.
Noon. Injection made.
1.30. The frog lies quite helpless. A spark of electricity
applied to the side causes reflex contraction of both legs. When
the poisoned leg is drawn out, the frog draws it up again with a
(95) F 2
68 ON THE NATUKE AND ACTION OF THE

wriggling motion. The poisonod leg at once reacts when the

toes arc pinched ; the ligatured one does not.
When the sides of the frog are irritated by an electric spark,
all the legs, except the ligatured one, give a twitch.
3.50. On exposing the lumbar nerves in the abdominal cavity
and irritating them by an induced current, the poisoned leg
contracted, the ligatured one did not.
The effect of the alcoholic extract in causing paralysis is
shown by this experiment ; but the insensibility of the ligatured
leg, which was in all probability due to an injury of the sciatic
nerve by the ligature, renders it difficult to say how much of
the paralysis was due to the cord, and how much to the nerves.
That the nerves were affei'-ted, however, seems clear from the
fact thar. the muscles no longer reacted to voluntary stimuli,
but did so when an extraordinary stimulus was occasioned by
pinching.
Experiment XLIV.
August 27th. — A small dog was chloroformed, and both vagi
were exposed.
12.35 P.M. About two grains of dried cobra-poison were
injected into the peritoneal cavity.
12.42. Water was thrown over the animal to revive him more
completely from the chloroform. Bowels acted. He is very
unsteady on his legs. Looks drunk.
12.44 Dog vomits freely.
12.45. Both vagi divided. The vomiting ceased, the breathing
became very slow, and the head was thrown up with the nose in
the air.
12.53. Has become very quiet. Falls down on his side. The
vomiting has not recurred.
12.55. Dead. Artificial respiration commenced.
1.12. On laying bare the skull and trephining, slight reflex
movements occurred in the limbs.
1.17. Micturated. On irritating the exposed cerebrum by a
Paradic current no contractions occurred in the limbs.
1.47. The spinal cord was exposed and irritated by a Faradic
current. No contractions occurred in any of the muscles, except
those to which the current was conducted, even when the
 POISON OF SOME INDIAN VENOMOUS SNAKES. 69

strongest was employed. On exposing the sciatic nerves and

dividing one of them, and applying a Faradic current, no effect
could be perceived when the electrodes were applied to either
the central or the distal end of the nerve. The motor nerves
were thus seen to be paralysed.
The heart continued to beat vigorously all the time. On
laying open the abdominal cavity, the intestines and peritoneum
were found in a state of intense congestion. Electrodes applied
to the lumbar nerves caused no contraction anywhere.
Thorax opened. The heart was beating vigorously. The
lungs were normal. A Faradic current applied to the phrenic
nerve caused no contraction of the diaphragm ; but when
applied to that muscle directly, it caused vigorous contractions.
The left vagus was divided and its peripheral end stimulated
by a Faradic current. The pulsations of the heart were at once
arrested, but again commenced ; and no further irritation of the
vagi had any effect on the heart.
2.2 P.M. Stomach removed. Its coats were intensely congested,
as though some irritant had been swallowed. It contained
much bile. The blood was tiorid, and formed a firm coagulum.
This experiment clearly shows that cobra-poison produces
paralysis of the motor nerves in warm as well as in cold-blooded
animals, the sciatics being so completely paralysed that they
did not respond to the strongest irritation, although respiration
was efficiently kept up and the circulation continued unim-
paired. In almost all the other experiments, when the nerve
was irritated immediately after death, contractions were pro-
duced but
; the same is the case when the animal is poisoned
with curare, and the contractions are due to the poison not
having had sufficient time to exert its full action.
The complete cessation of vomiting after division of the vagi
seems to indicate that the poison produces emesis by acting on
the peripheral terminations of the vagi, and not on any nerve
centre.
Action of Cohra-poiso7i on Secreting Nerves,
A notable symptom of cobra-poison in dogs is great
salivation ; and this might be supposed at first sight to indicate
that the poison acted as an irritant to the secreting nerves of
70 ON THE NATURE AND ACTION OF THE

the salivary gland. Nausea and vomiting being also present,

however, it is by no means improbable that the salivation is
due to the poison stimulating the secreting nerves of the
salivary glands not directly, but by reflex action, through the
gastric branches of the vagus. Unfortunately we are unable to
say in which of these ways salivation is induced, as we have not
noted whether it occurred after division of the vagus or not.
So far as memory serves us, we are inclined to think that it was
much less in these cases ; but on this point we cannot be at all
positive.
Whether cobra-poison has any stimulating action on secreting
nerves at first or not, it seems finally to paralyse them, or at
least greatly to diminish their power.
This is evident from the following experiment.

Experiment XLV.
A dog was etherised and the chorda tympani exposed after
its separation from the lingual nerve. A cannula was then
placed in the duct of the submaxillary gland. On irritating the
chorda by a weak Faradic current, applied at intervals, saliva
flowed freely. Some dried cobra-poison dissolved in water was
then injected into a vein in the leg. Shortly afterwards the
saliva began to flow much less freely than before ; and although
the current was increased in strength, only a small quantity
could be obtained.
Action on Sensory Nerves.
The sensory nerves seem to be little, if at all, affected by
cobra-poison. As appears from Experiment XXXVI, they
retain their power after the motor nerves are paralysed; and
Experiment XLVI shows the comparative effect of the poison
and of want of blood both on the sensory and motor nerves.
The former were so little affected by the poison that they
caused a ready response when those which had been deprived of
blood had nearly ceased to act. The motor nerves of the
poisoned limb, on the contrary, were quickly paralysed, while
those of the ligatured one, although doubtless weakened by the
loss of their vascular supply, long retained their irritability.
In Experiment LX the optic nerve and the aural and buccal
 POISON OF SOME INDIAN VENOMOUS SNAKES. 71

branches of the fifth nerve retained their irritability after the

cord had become nearly paralysed ; and, in several experiments,
reflex actions could be induced by irritation of the cornea after
voluntary motion and respiration had ceased.

Experiment XLVI.
The right leg of a frog was ligatured, excluding the sciatic
nerve, and a concentrated solution of dried cobra-poison
injected into the dorsal lymph-sac at 2.3 p.m.
2.5. Already affected. Much less active. Lies very quiet.
2.34. Paralysed. On touching his body he moves the right,
but not the left leg.
When acetic acid is applied to the hand, he straightens both
the arms and contracts the right, but not the left leg.
Acetic acid applied to left hind leg causes him to straighten
both arms and draw up the right leg ; but there is only feeble
movement in the left leg.
When acetic acid is applied to the right foot, the foot itself is
drawn up ; but there are no movements of any other part of the
body.
When acetic acid is applied to left hand, the left arm is
powerfully straightened, and there are strong contractions of
right hind leg, but none in the left, and little movement in any
other part of the body.
Acetic acid applied to left foot causes powerful extension of
both legs.
Acetic acid applied to the right foot has no effect. Electrodes
were inserted in the spine and the cord irritated by a Earadic
current. Distance of the secondary from the primary coil
18 cm. There was movement of left hand.
At 16 cm. movement of left hand and right leg.
At 12 cm. also faint movement of left leg.
At 15 cm. the interrupted current was kept up for some time,
and the muscular twitchings were more powerful in the left
gastrocnemius than in the right one.
On applying the electrodes to the lumbar nerves, coil at
48 c, the right leg contracts.
Coil at 42 cm. the left leg only twitches.
72 ON THE NATURE AND ACTION OF THE

Action on the Spinal Cord.

The spinal cord has the threefold function of a conductor of
sensory impressions, a conductor of motor impressions, and a
reflex centre ; and in examining into the nature of the action
of cobra-poison upon it, we must consider the manner in which
each of these functions is affected.
Cobra-poison, as has already been intimated, has a powerful
paralysing action upon the reflex function of the cord ; and this
is exemplified in Experiment XLVII, etc.
As a conductor of sensory impressions, the cord is able to
transmit two kinds, viz., tactile and painful, and these have been
stated to pass through different parts of the cord, the former
being conveyed by the posterior and lateral white columns, and
the latter by the grey matter.
From Experiments XXXVI and LX it would appear that
the power to convey tactile impressions is retained, both in
warm-blooded animals and frogs, after the transmission of
painful impressions has almost, or entirely, ceased. Thus, in
Experiment XXXVI the frog's leg moved when the animal was
laid upon its back, although an extremely painful stimulus, the
application of sparks from a coil to the eye, had caused in it
only the feeblest movement. In Experiment LX no response
was elicited by striking, pinching, or pricking the paws of the
animal, but when the ear was tickled the cat shook its head, or
moved its paw to ward off the irritant.
Erom these cases we think we are justified in concluding that
the grey matter of the spinal cord, thiough which painful
impressions are transmitted, is paralysed by cobra-poison ; but
the white sensory columns are little, if at all, affected. The
power of the cord to conduct motor impressions from the
encephalic ganglia appears to be little, if at all, affected, until
the apparent death of the animal; for in Experiment LX we
find that, very shortly before respiration ceased, and when
ordinary reflex action from the cord was nearly gone, purposive
or voluntary movements were still made. The absence of
movements in Experiment L, when the cord was irritated by a
needle, as well as the rapid loss of its power to produce move-
ment in the limbs when irritated by a Earadic current, is, we
 POISON OF SOME INDIAN VENOMOUS SNAKES. 73

think, to be attributed to paralysis of its function as an

originator, and not as a conductor, of motor impressions.

Experiment XLVII.
May 19th. — The kmibar nerves of a frog were exposed and
a ligature tied round the body, excluding these nerves.
12 (noon). Some dried cobra-poison dissolved in water wa&
injected into the dorsal lymph-sac.
1.45. The frog is partially paralysed ; mouth gaping; reflex
action is still marked in all the limbs, but more in the legs
than in the arms.
The heart was exposed when the ligature was applied ; it still
beats, but feebly and slowly.
1.50. Acetic acid causes reflex movements when applied to
either the hind or fore feet.
1.54. Applied to the nose, acetic acid causes movements in
all the extremities, and especially in the arms.
1.56. Applied to the right hind foot it causes movements of
the arms and of the jaw, which otherwise gapes.
2.2. Applied to the left hind foot it causes no reflex action.
2.14. Heart beating very feebly, 18 pulsations per minute,
Keflex movements still occur in all the limbs, and rather more
in the legs than in the arms.
2.30. Acetic acid produces no reflex action anywhere. The
heart has almost ceased to beat, and only contracts faintly at
long intervals.
2.34. All reflex action has ceased.
2.45. Electrodes placed in the spine and the cord irritated by
a Faradic current. At 15*5 cm. distance, faint contractions in
both arms. At 0 cm. distance, no contraction in legs. Sciatic
nerves exposed and irritated. 32*5, slight contraction in left
leg ; slight contraction in right leg.
2.48. Heart is still acting feebly and slowly ; brachial nerves
exposed and irritated.
At 46*5 cm., contractions in both arms.
2.49. The heart has now ceased to beat, except a faint
pulsation in the auricles.
74 ON THE NATUllK AND ACTION OF THE

Muscles of arms and legs exposed and irritated by single

induced shocks. 6*5 cm., muscles of both arms contract ;
muscles of both legs contract, but somewhat more strongly.
In this experiment there is no evidence of paralysis either of
the neives or muscles; death appears due to paralysis of the
spinal cord. This is caused by the action of the poison ; for
the circulation still continued, though feebly, after all reflex
action had ceased.

Experiment XLVIII.
A ligature was passed under the right sciatic nerve of a frog
and tightly tied round the limb, so as to constrict the whole of
the thigh, with the exception of the nerve, and completely
arrest the circulation.
At 1.8 half a drop of cobra-poison (first supply), diluted with
J c.c. of water, was injected into the dorsal lymph-sac.
1.12. The animal is sluggish.
1.15. Crawls about but sluggishly, and keeps the unligatured
limb drawn up close to the body.
1.20. The frog is more sluggish.
1.23. The hind limbs seem paralysed ; the fore limbs still
move, but much less than before.
1.30. Frog almost motionless. Contractions of the fore limbs
still occur ; but they no longer respond when pinched.
1.57. There is a faint motion in the limbs.
2.18. Erog is dead. Much ecchymosed.
On irritating the lumbar nerves in the abdominal cavity by
an induced current, the poisoned leg contracted rather more
than the non-poisoned leg.
On irritating the sciatic nerves in the thigh, below the level
of the ligature, the contractions of the poisoned leg were much
less vigorous than those of the non-poisoned leg.
Electrodes were then placed in the spinal cord, and the cord
irritated by an induced current.
2.34. When irritation is applied in this way the fore limbs
contract, but not the hind limbs.
When the cord is irritated lower down, the non-poisoned leg
responds to the irritation, but the poisoned leg does not.
 POISON OF SOME INDIAN VENOMOUS SNAKES. 75

The muscles of the ligatured leg respond to the direct appli-

cation of electricity more freely than the other muscles.
In this experiment the poison employed had not been
coagulated or dried, and the dose was somewhat small. The
failure of retlex action while voluntary motions still continued
in the nerves shows that the cord in this instance became
paralysed before the motor nerves. It is, indeed, difficult to
say whether the motor nerves were paralysed in this case or
not, as the muscles themselves were distinctly weakened.

Experiment XLIX.
December 1st, 1872. — The right leg of a frog was ligatured,
excluding the sciatic nerve, which was kept covered by a flap of
skin to prevent its becoming dry. A ligature was also put
round the left leg in a similar manner, but not tightened.
2 P.M. Cobra-poison injected into the abdominal vein.
The effect not being marked, the aorta was exposed.
2.27. Some poison injected into the aorta. It seemed to
take effect at once ; all motion ceased immediately.
2.30. The ligature was then tightened round the left leg.
2.48. The frog has since moved, but all motion has now
ceased.
2.52. Even when irritated by acetic acid there is no move-
ment. The heart is still contracting.
No reflex action occurs when a strong interrupted current is
applied to the nose or limbs.
Lumbar nerves exposed and irritated.
Eight. Distinct contraction of thigh. Coil at 58*5.
Left. „ „ „ „ 58-5.
Eight. Distinct contraction of whole leg. Coil 50.
Left. „ „ „ „ 50.
Sciatics exposed and irritated.
Eight. Contraction. Coil 77.
Left. „ „ 52.
3.28. Eight. Contraction. Coil 50.
Left. „ „ 43.
The poisoned leg seems to be losing its irritability more
quickly than the other. Irritability of spinal cord gone.
76 ON THE NATURE AND ACTION OF THE

3.35. The left still contracts, with the coil at 35. The other,
when irritated by a current of the same strength, contracts
more strongly.
The loss of power occasioned by the cessation of the circula-
tion in the ligatured limb (which is used as a standard with
which to compare the other) was diminished in this experiment
by injecting the poison directly into the circulation, so as to
enable it to reach the motor nerve-ends at once. As soon as it
had taken effect, the poisoned leg was likewise deprived of its
circulation, so as to bring the two limbs as nearly as possible
into the same conditions. The cause of death in this experiment
was paralysis of the cord, all reflex action having been almost
immediately abolished by the large dose of the poison injected
into the circulation, though the heart continued to beat. The
motor nerves were not at first affected, but after a little while
paralysis appeared in the poisoned limb. This experiment is
especially interesting in reference to the cause of death when a
considerable quantity of poison enters the arterial system at
once. In warm-blooded animals, as is shown by Experiment
LXYIII, the heart is arrested in many instances, and death
thus occasioned ; but when this is not the case, the appearance
of paralysis is probably due to affection of the nerve-centres.

Experiment L.
September 13th. — A ligature was placed round the middle of
a frog, excluding the lumbar nerves.
3 P.M. Some dried cobra-poison dissolved in water was injected
into the dorsal lymph-sac. Immediately after the injection the
animal could move all its limbs quite well.
3.3. Eestless ; moves all its limbs.
3.17. Can still move vigorously.
3.21. Can kick vigorously with its legs, especially the right.
When it moves it seems to overreach itself and turns over,
apparently from the hind limbs remaining unaffected and the
arms becoming partially paralysed.
3.40. Still moves voluntarily.
3.52. No reflex motion can be produced by touching any of
the extremities with acetic acid.
 POISON OF SOME INDIAN VENOMOUS SNARES. 77

A minute or two afterwards a slight twitch was noticed in

one arm, to which acetic acid had been applied, but whether
this was greatly delayed reflex action caused by the acid, or
whether it was due to something else, is uncertain. A needle
was now run down the spinal cord. It produced no effect.
The legs contracted readily when the lumbar nerves were
irritated.
The absence of motion in the legs when the cord was irritated
by a needle run down the spinal canal shows that the power of
the cord to originate motor impulses had been destroyed, as it
would usually have caused violent contractions in the ex-
tremities. These, having been protected from the action of the
poison either on muscle or nerve, would respond readily, as,
indeed, they did, to voluntary motor impulses shortly before
the death of the animal.
Experiment LI.
May 12th, 1873. — The sacrum of a frog was removed, and a
ligature passed round the body, excluding the lumbar nerves.
There was a good deal of bleeding.
12.30. Ligature tied.
12.33. A good dose of dried cobra-poison dissolved in water
was introduced into dorsal lymph-sac. Immediately afterwards
the frog sprung about once or twice.
1.27. Cornea insensible. On pinching the finger of either
hand it kicks out vigorously with tlie right hind leg. On
squeezing the toes of right hind foot it kicks out vigorously
with it. On squeezing toes of the left hind foot there is no
movement wliatever. On placing acetic acid on either foreaim
the frog kicks out strongly with the right hind leg.
2. Interrupted current, distance 7. Acetic acid applied all
over the frog no longer causes any movement whatever.
Electrodes placed in spinal cord just below occiput. Cord
irritated by an interrupted current. Eight leg ki cksvigorously.
Ko motion in any other part of the body.
Experiment LIl.
May 15th. — Frog ligatured round the middle, the lumbar
nerves excepted. A moderate amount of bleeding.
78 ON THE NATURE AND ACTION OF THE

12.40. Ligature applied.

12.52. Frog springs actively about when touched. A con-
siderable dose of dried cobra-poison dissolved in water injected
into the dorsal lymph-sac.
1.15. Cornea insensible. When either hind foot is pinched,
it is drawn up with a wriggling motion when the frog is lying
on the table. When the frog is suspended the foot is drawn up
at once.
1.20. On applying acetic acid to both fore limbs and nose, the
hind legs were vigorously drawn up to the body, but only after
a long interval.
1.26. Strong acetic acid applied to both fore limbs and nose.
Movements in all four limbs after 8 seconds.
1.36. Weaker acid applied to both fore limbs. Movements
in all the limbs in 37 seconds.
2.20. Applied to both fore legs. Movements in both fore
limbs in 4 seconds. Worse in hind legs.
2.53. Applied to all the limbs and the nose. No motion
anywhere. Divided medulla.
2.58. No reflex at all in 200 seconds after application of acid
to all the limbs and the body.
Abdomen opened. Lumbar nerves irritated just below exit
from spine.
Distance of primary
from secondary coil
Leg. in centims.
6'3 Left gastrocnemius contracts very
slightly ; right not.
0 Left gastrocnemius contracts slightly ;
right not. Both sciatics exposed and
irritated in the thigh some distance
below ligature.
Left 57 Tetanus of leg.
Eight 58 Tetanus. Nerve rather more firmly
applied to electrodes. Viscera re-
moved and brachial nerves irritated.
Eight 47 Contraction of foot.
Left 45 Contraction of foot.
 POISON OF SOME INDIAN VENOMOUS SNAKES. 79

In this experiment the loss of reflex motion was gradual.

It is shown to be due to paralysis of the cord, and not to
excitation of Setschenow's inhibitory centres, by the division of
the medulla having no power to increase the reflex action.
The fact that irritation of the lumbar nerves hardly caused
any contraction in the legs, while irritation of the sciatics below
the ligature caused them to contract readily, indicates either
that the nerves had been injured by the ligature, or that the
part of them lying between the spine and the ligature had been
paralysed by the poison. The latter is possible ; but as the
frog moved its arms and not its legs before death, the former is
more probable.
Several years ago Setschenow showed that the optic lobes in
the frog possess an inhibitory power over the reflex acts
originating in the spinal cord. Irritation of the optic lobes
greatly lengthens the time required for the performance of any
reflex act, and thus produces an effect apparently similar to
that of diminished excitability, or paralysis, of the spinal cord.
A diminution in reflex action may therefore be due to two very
different causes :— (1) Lessened excitability of the cord, and
(2) excitement of Setschenow's inhibitory centres. These can,
however, be readily distinguished from one another by dividing
the cord just below the medulla. It is thus separated from the
inhibitory centres ; and if the diminution in reflex action is due
to excitement in them it will disappear, but will be permanent
if it is caused by paralysis of the cord. The following
experiment, performed by Tiirck's method, shows that in cobra-
poisoning the diminution of reflex action is due to the latter of
these causes.
Experiment LIII.
May 19th, 1873. — The right leg of a frog ligatured, excluding
the sciatic nerve.
3.5. A full dose of dried cobra-poison dissolved in water
injected into the dorsal lymph-sac.
3.54. The animal appears dead. Both hind legs dipped into
dilute acetic acid. Eight arm twitched.
3.57. Keflex action in both arms. None in the legs when
the left leg is dipped in the acid.
80 ON THE NATURK AND ACTION OF THE

4. No reflex action from ligatured leg.

4.2. No reflex action from lel't leg in 60 seconds when it is
dipped in the acid.
4.10. No reflex action from either leg in 250 seconds.
The medulla was now divided in order to separate the cord
from Setschenow's inhibitory centres.
4.35. No reflex action can be observed.
As the operation of dividing the cord somewhat lessens the
excitability, in the following experiment the division was
performed on the previous day, so that its effects should have
passed off before the poison was injected.
The columns headed " left " and " right " indicate the number
of seconds which elapsed before the corresponding leg was
drawn out of the acid.

Experiment LIV.
May 15th. — About 3 p.m. divided the medulla of the frog.
May 16th. — Suspended the frog by a hook in its jaw.
Time. Left. Rig}
11.17 8 8 When touched the frog draws up its legs,
and makes wiping movements on its
flanks.
11.44 5 6
11.59 4 5
12.6 o 7
12.10 5 3
12.25
12.30 8 10
The point of an aneurism needle was
drawn across the spine so as to destroy
any remnant of medulla. The frog
at once passed into a state of opis-
thotonos, but in a few minutes this

12.40 9 passed off.

12
12.48 12 9
12.55 10 8
12.58 10 5
1 8 5
 POISON OF SOME INDIAN VENOMOUS SNAKES. ^1

Time. Left. Eight.

1.2 ... ... Injected a drop of concentrated solution
of dried cobra-poison under skin of
back.
1.5 ... ... It draws up legs and wipes back once or
twice.
1.9 11 10
1.16 10 10
1.18 ... ... Another drop.
2.35 300 300 No reflex action in either foot. The
heart could not be seen beating till
the frog was opened ; then it was
found beating slowly and languidly,
24 in a minute.
2.45 ... ... Half a drop of liquor atropise placed on
heart. Immediately afterwards its
pulsations became more forcible, but
were still 24.

Experiment LV.
May 15th. — Divided the medulla of a frog about 3 P.M.
May loth. — Suspended it by hook through the jaws.
1.3 6 3
1.6 5 5
1.11 6 6
1.16 6 12
1.18 5 6
1.19 ... ... One drop of slightly diluted, but stilF
concentrated solution of cobra-poison
injected under the skin of the back.
2.23 10 14 The foot was twitched up in the acid at
these times, but the leg was not
drawn up.
3.25 150 150 No reflex action. Strong acetic acid
causes none. Thorax opened. Heart
quite still.
These experiments show that the time required for the per-
formance ofa reflex act went on increasing, or, in other words,
(95) G
82 ON THE NATURE AND ACTION OF THE

the excitability of the cord went on diminishing, after the

injection of the poison ; and all communication with the
inhibitory centres having been previously cut off by dividing
the medulla, this effect could only be due to the action of the
poison on the cord.
Experiment LYI.
At 1.16. Half a drop of cobra-poison was injected into the
peritoneal cavity of a guinea-pig.
1.17. The animal is restless and twitching ; runs about.
1.18. Micturates.
1.24. It is getting weak and sluggish. The hind quarters
have assumed a crouching posture. It moves when roused.
1.26. It looks drowsy, is disinclined to move, and is jerking.
The hind legs are almost paralysed. When they are retracted
it draws them up with difficulty.
1.27. Has defecated. Is convulsed generally, but the con-
vulsions are more marked in the hind quarters.
1.32. Convulsions continue. They are not increased or
excited by external stimuli. Cornea insensible.
1.34. Mouth only twitches. Heart acting vigorously.
1.35. The animal is quite dead.
1.36. The spinal cord irritated by an induced current through
electrodes inserted in the vertebral column. The irritability of
the cord seems perfect. (It was judged of by the contractions
of the hind limbs.)
1.40. The heart continues to beat. Thorax laid open. The
vagi isolated and one of them irritated. The cardiac action
seems to be increased by the irritation of the vagus. The
auricles contract very rapidly, the ventricles not so rapidly.
The cord is still irritable.
1,50. The irritability of the spinal cord as affecting the lower
extremities is almost gone ; as affecting the upper limbs it is
still retained. Heart still contracts vigorously.
1.54. The lower limbs are no longer affected by electricity
applied to the spinal cord. The upper limbs are aff'ected.
1.56. The spinal cord is still slightly irritable. The heart is
acting freely.
2. Cord still slightly irritable. Heart acts briskly.
 POISON OF SOME INDIAN VENOMOUS SNAKES. 83

2.2. Heart acts as vigorously as ever. Artificial respiration

was tried.
2.15. Artificial respiration has been kept up, but has been of
no service. The irritability of the cord is much diminished,
though not quite extinct. The strongest current causes a
barely perceptible motion. The heart is still acting. There
are spots of ecchymosis all over the intestines.
2.40. The irritability of the cord is quite gone. The heart is
still acting. The blood collected from the large vessels
coagulated firmly.
Experiment LVII.
August 30th. — A cannula was placed in the trachea of a cat,
and 1^ decigramme of dried cobra-poison was weighed out and
dissolved in a small quantity (about 2 c.c.) of distilled water.
The solution was clear and glairy, hanging in threads from the
stirring- rod.
2.40. Injected about two-thirds of this solution under the
skin of the right hip.
2.50. Kespiration is quicker. The cat lies down and does
not like to rise. When raised it walks toward a dark corner,
dragging the right leg.
2.58. Shivering of right leg and partially of body. No other
symptom than paralysis of right leg being noticed, a further
injection was made.
4.26. The remainder of the solution injected in the same
place. This also seemed to produce little effect.
5.10. Injected 0*02 grain dissolved in a little water, as the
cat did not seem about to die.
6.2. Injected IJ c.c. of a 2-per-cent. solution of cobra-poison
in distilled water, partly into a vein in the back of the left
hind leg, partly into the peritoneum. The left hind leg seems
partially paralysed. The respiration has a peculiar character,
the diaphragm seeming to relax with a jerk. The respiratory
movements are very deep. Peristaltic action of bowels.
6.20. The fore legs are now becoming paralysed.
6.25. Eespiration quick. Entirely diaphragmatic. Cornea
quite sensitive. The animal opens its mouth when the tail is
pinched, but not when the feet are pinched.
(95) G 2
84 ON THE NATURE AND ACTION OF THE

6.37. Sensibility of the cornea seems nearly gone. When

the inside of the ear is tickled the animal shakes its head.
6.43. Altlioiigh respiratory movements still continued,
artificial respiration was begun. The animal was laid in an
apparatus which kept it warm.
6.45. The cat tries in vain to vomit. The cornea is almost
insensible.
About 8.30 the heart-beats ceased. The body of the animal
was examined next day at noon. Rigor mortis well marked.
The body of the animal had a strong odour of decomposition.
The lungs were congested, the right side of the heart gorged,
the left empty and firmly contracted. The pericardium
contained a quantity of dark-red serum. A considerable
quantity of dark-red serum was contained in the abdominal
cavity. Inside of stomach quite normal. The bladder was
firmly contracted and quite empty. Where the injection had
been made, the muscles were infiltrated with blood, soft, and
decomposed. Those of the left thigh were normal.

Experiment LVIII.
July 14th, 1873. — Some dried cobra-poison dissolved in
water was injected into the peritoneum of a cat at 2.15 p.m.
2.20. Vomits.
2.30. Vomits again. The animal can walk perfectly, but it
prefers to lie on its side.
2.40. Can walk, but seems slightly giddy.
2.45. Vomiting and defecation.
3.12. Sensibility of the cornea nearly gone. When the ear
is irritated the cat shakes its head. When the eye is touched
the eyelids do net move ; but when the point of a pair of
forceps is pressed into it, the fore foot is raised to push the
forceps away.
3.20. The animal suddenly got up, walked a few steps, and
then fell.
3.22. It seems as if it wanted to vomit, but is too comatose.
When the ear is tickled it shakes its head.
3.26. There is distinct reflex action on irritation of the hind
feet, but not when the fore paws are pinched.
 POISON OF SOME INDIAN VENOMOUS SNAKES 85

3.32. Breathing is getting deep and slow, and the head is

extended at each inspiration. There is still motion of the
head when the ears or mouth are tickled. A minute or two
ago it got up, stood for a second or two, and then fell.
Kespiration gradually ceased. A cannula was placed in the
trachea, and artificial respiration kept up. The heart ceased to
beat very shortly after. Electrodes were placed in the cord
opposite the seventh and twelfth dorsal vertebra. A Faradic
current passed through them caused contractions in the
adjoining muscles of the back, but none elsewhere. The left
sciatic was exposed and irritated. The limb contracted.
About an hour afterwards curious and somewhat rhythmical
movements took place in the right foot. The sciatic had not
been exposed in the right leg.

Experiment LIX.
July 25 tb. — At 3.34 a little cobra-poison was injected into
the peritoneum of a guinea-pig. Immediately afterwards the
animal became restless and uneasy.
3.38. The animal is quiet. Occasional lifting of head. The
fore legs are spreading out laterally. When made to walk it
staggers, and has difficulty in maintaining its balance. It
rises up and runs when any sound is made. Eespirations
68 per minute.
3.44. Tlie left ear is drooping.
3.58. Passed milky urine.
4.4. Convulsive motions occur, but the animal can still run.
Almost immediately after, when laid on its side, it could not
get up.
4.7. The cornea is now insensible. A cannula placed in the
trachea and artificial respiration commenced.
4.15. A needle placed in heart. Pulsations quick. The
artificial respiration was discontinued. The pulsations became
quicker.
In this experiment the paralysis began in the fore legs.
There was distinct loss of co-ordination ; but the animal could
run up to the last, although it could not walk. This indicates
that the higher co-ordinating centre (probably the cerebellum)
86 ON THE NATURE AND ACTION OF THE

was paralysed before the lower ones, just as in the case of

a man who is drunk.
Experiment LX.
August 29th, 1873. — A cannula was placed in the trachea of
a cat about 5.30 p.m.
5.35. One decigram of dried cobra-poison, dissolved in 2 c.c.
of water, was injected into the peritoneal cavity.
5.39. The animal lies on its side breathing very rapidly and
wagging its tail. Eises, sits with head erect and mouth widely
open.
5.45. The respiratory movements are very rapid and shallow,
with occasional deep ones. The animal sits up. Kespirations
240 per minute. Pulse 148 per minute.
6.3. The animal was lying down and occasionally rising. Is
now lying down. The respiratory movements have an extra-
ordinary vermicular character. Dr. Sanderson ascertained by
palpation that this is due to the diaphragm contracting before
the thoracic walls expand.
6.7. The respirations are feeble, with occasional deep ones.
The cat walks quite well. The bowels act.
6.20. Bowels act again. Tries to vomit several times.
6.37. The cat lay on its side, and stretched itself once or
twice in a sort of convulsive manner.
6.41. Lies quietly. When the cornea is touched or poked
with a pointed instrument, or when the finger is rubbed over
it, the eyelids do not close, nor does the animal give any sign
of feeling. When the hind legs are struck, it moves its fore
legs very faintly. Eespiration is quite regular and apparently
normal. The end of the tail gently moves from side to side.
When the inside of the ear is tickled the animal shakes its
head. It took a deep breath, and moved its head voluntarily.
The pupil is much contracted. When the arms are irritated by
a sharp stick the animal draws its body slightly together.
A minute or two afterwards it moved its tail fi'om side to side
several times voluntarily. The animal was lying on its side.
Lifted it up and laid it on its belly with its feet under it. It
rose up and walked several steps.
6.45. The cat again rises and walks, but staggeringly. It
 POISON OF SOME INDIAN VENOMOUS SNAKES. 87

then falls and lies on its side. The hind legs seem to be
weaker than the fore legs.
6.52. Animal lying on its side. When a bright light is"
brought before its eyes it draws back its head. The cornea is-
quite insensible. When the paws are irritated by striking,
pinching, or pricking there is no response. When the inside
of the ear, nose, or mouth is tickled, the cat shakes its head,
and sometimes moves its paw to put the irritant away.
7.5. On touching the eyes it sometimes draws back its head,
but there is not the slightest motion of the eyelids. It
voluntarily moved its paws and head as if to rise, and then
sank back as if asleep, and lay still on its side.
7*6. Laid it on its belly. It rose and walked a step or two
towards a darker corner and then fell. Immediately after-
wards the muscles of the neck gave a sort of shudder. After
movement the respiration becomes much quicker, and then
rapidly becomes slow. After lying a minute or so its respira-
tions are 27 per minute.
7.25. Moves its paws and tries to get up voluntarily, but
cannot do so. Irritated paws and ear by sparks from a Du-Bois
coil. No reaction. On irritating the inside of the thigh in
a similar manner, it stretched out its fore legs, protruded its
claws, and seemed to be trying to grasp me.
7.33. The respiration ceased without convulsions. The
cannula in the trachea was immediately connected with an
apparatus for artificial respiration, and this was kept up.
While some adjustment was being made on the apparatus the
animal was observed, and its heart was found to have ceased to-
pulsate about five or ten minutes after artificial respiration had
been begun.
On opening the thorax the lungs were found somewhat
congested. The right side of the heart was moderately filled.
The left ventricle was quite empty and firmly contracted. The
surface of the stomach and intestine was much congested.
The interior of the stomach was not congested.
In this experiment, respiration continued for two hours after
the injection of the poison. The most remarkable points as
regards respiration are its great acceleration, with occasional
"9$ ON THE NATURE AND ACTION OF THE

'deep breaths at first, its vermicular character about the middle

of the experiment, and its regularity towards the end. Reflex
action seemed entirely abolished, and sensation very much
impaired ; the mental faculties seemed sluggish ; but voluntary
power was retained, and the movements of the animal were not
indefinite but distinctly purposive.
The motor nerves and muscles were evidently not paralysed ;
but the grey matter of the cord seemed to have lost its power
of inducing reflex actions or of conveying painful impressions.
Tactile impressions, such as laying the animal on its belly, still
caused reaction. The movements thus induced, as well as those
caused by irritating the ears, etc., may all be reasonably ascribed
to the action of the brain.
Closure of the eyelids would seem to be a purely reflex act,
in which, the brain is altogether unconcerned.

Experiment LXI.
October 29th, 1872. — To ascertain if a mixture of strychnia
and woorara produced the same effect as cobra-poison, a guinea-
pig weighing 1 lb. was experimented upon.
2.36.30. 1 c.c. of a solution of woorara (1 in 1000) was
injected under the skin of the side.
2.54. As the first dose seemed to produce little effect, another
cubic centimetre was injected in the same way as before.
2.5 G. A drop or two of Liquor Strychniae (4 grains to 1 fluid
ounce) was injected into the side.
2.57. Twitchings motions of the body begin. (They were
not exactly like those produced by cobra-poison.)
2.58. The animal has fallen over on its side and is paralysed,
but the twitching continues.
3.2. The animal is dead. No convulsions. On opening the
animal the heart was found contracting vigorously.
Electrodes were inserted in the spinal column and the cord
irritated by an induced current. The limbs contracted when
irritation was applied to the cord. The sciatic nerve was
exposed and irritated by an induced current. The muscles of
the limb contracted.
3.9. Heart still contracts feebly. The lungs are congested.
 POISON OF SOME INDIAN VENOMOUS SNAKES. 89

Action of Cohra-poison on the Stomach and Intestines.

One of the most noticeable symptoms of cobra-poisoning in
dogs is vomiting of a violent, repeated, and most distressing
kind ; and it is also present in cats and guinea-pigs, though to
a less degree. Its occurrence in guinea-pigs is somewhat extra-
ordinary, as these animals very rarely vomit, and, according
to Schiff, only do so after their vagi have been divided ; whereas
other animals which vomit under ordinary circumstances are
then unable to vomit at all. The nervous centre by which
the movements of vomiting are originated is closely connected
with the respiratory centre, and it may be set in action by
stimuli conveyed to it by the branches of the vagus distributed
to the stomach and other intestinal organs, and also through
the pharyngeal branches, either of the vagus or, possibly, of
the glosso-pharyngeal nerve. The brain can also excite it ; but
the vomiting it produces is not usually prolonged. The
vomiting which occurs in cobra-poisoning is, in all probability,
due, in part, to irritation of the gastric or abdominal branches
of the vagus — but not altogether ; for the attempts to vomit
continued in Experiment LXY after that nerve had been
divided in the neck ; and the failure to bring anything up is
to be attributed to the cardiac aperture of the stomach failing
to dilate at the proper time — a result which usually occurs
after section of the vagus.
In Experiment XL IV there was intense congestion of the
mucous membrane of the stomach ; but this does not occur in
all cases. It could hardly be due to the division of the vagi
in this instance, as that operation is usually followed by paleness
of the membrane. The intestinal movements are quickened by
the poison, since there is purging, which cannot be due to
increased intestinal secretion, as the stools consist chiefly of
mucus. The movements continue for a considerable time after
death.
Effect of Cohra-poison npon Respiration.
The action of cobra-poison upon respiration is perhaps the
most important of those which it exerts upon the organism ;
for it is through this action that death is generally caused.
90 ON THE NATURE AND ACriOX OF THE

The respiratory movements, besides being frequently altered in

form, are generally quickened after the introduction of the
poison ; then the number sinks to the normal or even below it ;
they become weaker and finally cease altogether. The blood,^
being no longer aerated, becomes more and more venous, and,
by irritating either the respiratory centre itself or some nervous
centre closely associated with it, occasions general convulsions.
These disappear whenever artificial respiration is begun and the
blood again aerated ; while they reappear when the respiration
is discontinued and the blood regains its venous character.
This condition is to be observed in Experiment LXII. The
dependence of the convulsions on the venosity of the blood is
well shown by Experiment VIII of our former communication,
where the condition of the blood was indicated by the colour
of the fowl's comb, and as this became florid, or livid, the
convulsions disappeared or returned. After they have continued
a short while the convulsions cease ; for the venous blood does
not maintain the vitality of the nervous centres sufficiently to
keep them in action ; but if artificial respiration be recom-
menced, the first effect of aerating the blood is to renew the
convulsions, by increasing the vitality of the nervous centres,
and rendering them again susceptible to the action of a
stimulus, though the convulsions disappear as soon as the
arterialisation has proceeded sufficiently far.
Increased rapidity of the respiratory movements may depend
either upon greater excitability of the respiratory centre in the
medulla, or upon stimulation of some of the afferent nerves
which have the power to accelerate it. The chief of these are
the pulmonary branches of the vagus, though there are probably
others proceeding from the cerebrum, through which the
emotions influence the breathing, and others from the general
surface of the body.
In order to ascertain the cause of the acceleration of respira-
tion, several experiments were made. Experiment LXIII shows
that it is not due to the action of the poison on the cerebrum ;
for it occurs after the cerebral lobes have been removed. The
ultimate arrest of respiration is probably due, in part, to
paralysis of the medulla, and, in part, to paralysis of the motor
 POISON OF SOME INDIAN VENOMOUS SNAKES. 91

nerves distributed to the respiratory muscles. The complete

insensibility of the phrenic nerve to the strongest stimuli, while
the sciatics and vagus still retained a considerable amount of
irritability, in Experiments XLIV and LXYI, is very remark-
able. The want of co-ordination between the diaphragm and
the thoracic muscles in Experiment LX is not improbably due
to paralysis of the phrenic nerve, though it may be attributed
to some alteration in the respiratory centre. Brown-Sequard
states that the diaphragm contains ganglia which will keep up
rhythmical movements in it after the central nervous system
has been destroyed ; and if this statement is correct, it seems
probable that paralysis of the phrenic, by interrupting the
connection between the respiratory centres in the medulla and
those in the diaphragm, may allow the movements of the
thoracic respiratory muscles and of the diaphragm to occur one
after the other instead of simultaneously.
It is difficult to say to what extent the stoppage of respiration
depends on paralysis of the medulla, or of the motor nerves, in
each case. Probably the effect of the one preponderates in
some cases, and that of the other in others.

Experiment LXII.
November 29th, 1872. — The vagi of a cat were exposed and
some dilute cobra-poison injected subcutaneously. Little effect
being produced, the dose was repeated, and then a solution of
alcoholic extract of the poison injected subcutaneously and into
the peritoneum. After the last injection the animal became
feebler. No vomiting. Before death slight convulsions occurred.
After they ceased, a cannula was put in the trachea and arti-
ficial respiration begun. Slight convulsions again appeared, but
ceased as respiration was continued. They recommenced when
the respiration was stopped, and disappeared when it was again
begun. On once more stopping respiration and allowing the
convulsions to cease spontaneously, recommencement of the
respiration caused them again to appear.
•92 ON THE NATURE AND ACTION OF THE

Experiment LXIII.
July 21st, 1873. — A rabbit was etherised and the cerebral
lobes were exposed and carefully removed.
3 P.M. Operation finished.
3.7. Respirations 37 per minute.
3.8. A small quantity of cobra-poison injected into the flank.
Active reflex movements occur on pinching the limbs and tail,
■and respiration also becomes more rapid.
3.12. Respirations 96 per minute. Heart's action feeble.
3.23. Breathing hurried. Reflex force continues active.
Another quantity of cobra-poison injected, the two doses
together not making more than a moderate amount.
3.37. Respirations very feeble. The upper part of the spinal
cord, on being irritated by a Faradic current, caused movements
in the limbs. Reflex movements still present, but much
diminished.
3.38. Respirations ceased. Cannula inserted in the trachea,
a,nd artificial respiration commenced.
3.40. Sciatic nerve exposed and irritated by a strong current,
induced twitchings in the limbs, but occasioned no reflex move-
ment in any other part of the body.
3.45. The animal seems perfectly dead. The strongest current
produces no effect either when applied to the cord or to the
sciatics.
The colour of the muscles seems changed when compared
with those of the other rabbit (Experiment LXIV) which had
no poison. They are of a less vivid colour, and altogether have
an altered appearance.
In this experiment the respirations became quickened from
37 to 96 per minute after the injection of the poison, although
the cerebral lobes had been previously removed. The accelera-
tion, therefore, could not be due to emotion, or to the action of
the poison on the cerebrum. A comparison with Experi-
ment LXiV, in which the cerebral lobes were removed without
injecting any poison, shows that in the latter no acceleration
whatever occurred, and the respirations became gradually slower
till they ceased.
 POISON OF SOME INDIAN VENOxMOUS SNAKES. 93

Experiment LXIV.
July 21st, 1873. — A rabbit was etherised, the calvaria
removed, and the cerebral lobes carefully excised. Tlie bleeding
was arrested by cotton-wool steeped in perchloride of iron, and
by the actual cautery.
1.18. The operation concluded.
1.23. Respirations 32 per minute. Keflex movements well
marked on pinching feet or tail.
1.33. Respirations 16 per minute and much deeper ; and each
one ended with a jerk, as if of the diaphragm.
1.35. Fore legs extended in a convulsive manner. Respira-
tion ceased almost entirely; but at long intervals of about
15 and 20 seconds, an inspiration occurred.
On pinching the feet the respiratory movements became more
perfect, though feeble.
1.43. Heart beats rapidly but feebly. Respiration has ceased.
Reflex movements are still well marked.
1.44. Cannula placed in the trachea, and artificial respiration
begun. Reflex movements continued for some minutes ; but
then the heart ceased to beat.
3.55. Sciatics exposed and irritated by a Faradic current, l^o
contractions occurred in the limbs. The muscles contracted
when irritated directly.
From these experiments it was evident that the accelerated
respiration was not of cerebral origin ; and it was therefore
probably due to stimulation of the pulmonary branches of the
pneumogastric by the poison. If this were so, the acceleration
would not appear if the vagi were divided previously to the
injection of the poison, as the stimulation of the terminal
branches of the nerves in the lungs would no longer be con-
ducted to the medulla. The following experiment shows that
our hypothesis is correct, the injection of the poison rendering
the respirations, which had already been greatly diminished in
rapidity by division of the vagi, still slower.

Experiment LXV.
September 15th. — A dog was chloroformed ; both vagi were
divided, and a cannula placed in the trachea. On recovering
94 ON THE NATURE AND ACTION OF THE

from the chloroform, the aniinai became very restless and

retched constantly, but was unable to vomit. A little while
afterwards he became more quiet, and his respirations were
counted.
3.10. Eespirations 7 J per minute.
3.13. Eespirations 7 per minute.
3.15. About 0*01 grain of dried cobra-poison dissolved in
^ c.c. of water was injected into the vein of dog's leg.
Immediately the animal became very restless, and tried in
vain to vomit. Eespirations 7 per minute.
3.21. Constant retching, but no vomiting. Eespirations 7.
3.23. AbouD 0*02 grain more was injected.
3.27. Constant retching. Eespirations 6. The animal now
lay down exhausted, and was killed by a blow on the head.

Experiment LXVI.
July 9th. — About 1 grain of dried cobra-poison dissolved in
water was injected into the flank of a white cat.
3.38. Injection made.
3.43. Cat seems depressed, sits with head drooping and eyes
nearly shut. Licks its lips occasionally. Pupils moderately
dilated.
3.48. Eubs its ear with fore paw, and licks fore paw after-
wards. Isdisinclined to move. Pupils more widely dilated.
4.25. Another dose injected.
4.50. Another dose injected into peritoneum. As yet there
is no symptom except depression and languor.
4.51. Vomiting. Lies crouched down.
5.5. Still vomiting.
5.14. Lies on its side. Movements of vomiting. When the
cornea is touched the eyes move, but the lids do not close.
There is also sometimes a movement of fore foot as if to ward
off the irritant.
5.17. Whining. Pupils much contracted. AVhen the inside
of the ear is tickled the animal scratches at its shoulder with
the hind leg of same side. It cannot stand. It shakes its
head sometimes when its ear is tickled.
 POISON OF SOME INDIAN VENOMOUS SNAKES. 95

5.25. Eeflex movement of leg much fainter when the ear is

irritated.
5.31. Tries to get up voluntarily. Got up, staggered some
steps. Convulsive movements. Death. Immediately a cannula
was placed in the trachea and artificial respiration begun.
Sciatic nerve isolated. Irritated by induced current. Foot
twitched when secondary coil was at 57 cm.
About 6.30. Electrodes screwed into cord about 2nd and 5th
dorsal vertebr£e.
The strongest current of the coil produced contraction of the
muscles of the back, but no contraction of the limbs. The
sciatic nerve, when irritated directly, caused contraction of
foot with the coil at 23.
6.50. Tlie pVirenic nerve irritated ; no contraction of
diaphragm ; vagus irritated ; heart stopped.
In this experiment the continuance of reflex action on
irritation of the ear, and of voluntary movements, after reflex
action on irritation of the eye had disappeared, and almost up
to the time of death, are remarkable ; as is also the paralysis of
the phrenic before the sciatic and vagus nerves.

Action of Cohra-poiso7i on the Circulation.

In most cases of death from cobra-poison, the fatal issue is
not to be attributed to any failure of the circulatory apparatus ;
for the heart continues to pulsate vigorously, long after all
motions have ceased in the voluntary muscles and the strongest
irritation applied to the spinal cord and motor nerves fails to
produce the slightest effect. But this only occurs when the
dose of poison is not excessive ; and when a large quantity of
it is introduced at once into the circulation, the heart is not
exempted from its action, but is, on the contrary, most
seriously affected. This is seen in Experiments LXVIII and
XXVIII, where the poison having been either injected into the
circulation, or absorbed with extreme rapidity, the action of
the heart was at once arrested. But it is to be noted that it is
not paralysis, but tetanic contraction of the heart which is
produced, the poison, in fact, seeming to act as an excessive
stimulus; and this being the case, we feel less surprise on
9b ON THE NATURE AND ACTION OF THE

finding that, in ordinary cases of poisoning, the cardiac action

may be maintained by the use of artificial respiration for more
than thirty hours, as Mr. Eichards has succeeded in doing in
India. The cardiac movements cease much sooner in frogs
poisoned by cobra-venom than in those paralysed by curare —
the pulsations in the latter often continuing for very many
hours, or even for one or two days. They are also arrested by
the direct application of the poison to the heart, as in Experi-
ment LXXII. Its action seems to be somewhat different in
degree, if not in kind, when applied to the outside of the heart,
as in Experiment LXX, and to the inside, as in Experi-
ment LXXII ; for in the former case the pulsations continued
for a considerable time, while in the latter they were instantly
arrested, the heart stopping in partial systole and moderately
contracted.
The action of cobra-poison being exerted on the heart of the
frog after its excision shows that it acts on the heart itself;
and its effect being very much the same without the body as
within it renders it probable that the central nervous system is
little concerned in the arrest of circulation by the poison, at
least in the frog.
The stoppage of the excised heart may be due (1) to irritation
of the inhibitory centres contained within it, or (2) to paralysis
of its motor ganglia, or (3) to excessive stimulation of them
producing tetanus, or (4) to the action of the poison on the
muscular fibre of the organ. It is not due to the first of these
causes ; for atropia, which paralyses the inhibitory ganglia, does
not restore the movements. The second is improbable, as the
heart does not stop in diastole but in systole, and resists
distention by fluid within it. The third seems the most
probable cause, as one does not see why the poison should
arrest the cardiac pulsations at once when applied to the
interior of the organ, and not do so when placed on the
outside, if it acted on the muscular fibre, whereas it may
readily be supposed that the poison may reach the ganglia
more readily from the inner side of the heart — though we do
not venture to assert that this is the true explanation of the
facts we have observed.
 POISON OF SOME INDIAN VENOMOUS SNAKES. 97

The inhibitory branches of the vagus are not always

paralysed (Experiment LXVI) ; but sometimes the cobra-poison
appears to affect them as well as the motor nerves ; and in this
it resembles curare, which in small doses does not impair the
inhibitory action of the vagus, but in large doses completely
destroys it. In Experiment LVI irritation of the vagus
quickened, instead of retarding, the cardiac pulsations —
a circumstance which indicates that the inhibitory fibres of the
vagus were paralysed by the poison, but not the accelerating
ones.
The capillary circulation is not unaffected by the poison. In
Experiment IV of our former paper, the rhythmical contrac-
tions and dilatations, altogether independent of the cardiac
pulsations, which Schiff first observed in the rabbit's ear, and
which were noticed by Ludwig and Brunton in the vessels of
many parts of the body, were greatly increased by the injection
of the poison.
In Experiments LXXIV and LXXV the blood-pressure
remained high after the heart had ceased to beat. This shows
that the arterioles, or capillaries, must have been much con-
tracted, thus opposing a barrier to the exit of blood from the
arteries into the veins.

Experiment LXVI I.
May 21st. — A cannula w^as placed in the trachea of a large
black rabbit ; and some dried cobra-poison dissolved in water
was injected into the hip at 1.25 p.m.
1.50. The animal shows symptoms of poisoning. Limbs
becoming weak. There is trembling, and the body sinks down.
There is starting. The respiration is hurried.
2. Eeflex action is well marked when the animal is touched.
The limbs seem almost paralysed ; but the animal moves the
head and neck freely. It makes efforts to rise, but is unable to
do so. The head falls over ; the respiration is getting feeble.
The animal seems quite conscious, and starts if touched.
2.4. It is now quite feeble. When the cornea is touched the
reflex action is less than before.
2.5. ISTo convulsions. Artificial respiration commenced. The
(95) H
 98 ON THE NATUl^E AND ACTION OF THE

rabbit, wrapped in cotton, was placed in a double tin bath

filled with warm water. Temperature in rectum 98°-8.
2.11. Kespiration discontinued for a space.
2.12. Convulsive twitchings of legs begin. Natural respira-
tion has ceased.
Artificial respiration resumed. Pupils contracted. Reflex
action on irritation of the cornea has ceased.
2.16. Since the artificial respiration has been resumed there
have been no more convulsive twitchings.
2.55. The heart beats rapidly but vigorously.
Temperature 101°. The bath being rather hot, its tempera-
ture was low^ered by a little cold water added to it.
2.57. The animal passed a quantity of urine tinged with
blood.
3.5. Heart beats vigorously.
3.15. The eyeballs are very prominent ; pupils normal.
3.45. Heart beating well, but apparently not so vigorously as
hefore. Temperature 100°'5.
3.55. The bath getting cold ; a little hot water added to it.
Tiie heart beating more vigorously than at 3.30.
4.20. Heart beating well — if anything more vigorously than
ibefore.
4.40. Heart beats steadily, but apparently with less vigour.
Temperature 100°-2.
5. Heart sometimes beats steadily 130-140 times per minute.
Then it gets feeble and intermits, and again beats steadily.
5.5. Heart beats more freely. Added more warm water to
the bath.
5.25. Heart beats rapidly but more feebly.
5.35. The same.
6. Heart beating rapidly, perhaps rather more feebly.
Temperature maintained at 100°'5.
6.10. Heart beating well and more vigorously.
6.30. Heart beating well, rapidly but steadily.
The attendant, being left alone, discontinued artificial
respiration, and the animal died. The fluctuations in the
activity of the pulsations were, in all probability, due to the
more or less perfect maintenance of the artifical respiration.
 POISON OF SOME INDIAN VENOMOUS SNAKES. 99

Experiment LXVIII.
A small rabbit had two drops of diluted cobra-poison injected
into the jugular vein. In 30 seconds he was in convulsions,
and in 60 seconds was dead.
The thorax was opened immediately ; the heart had ceased
to beat, and was firmly contracted.
A large vein entering the auricle on the left side was
pulsating vigorously and rhythmically, though no part of the
heart itself showed the least trace of motion.

Experiment LXTX.
June 26th, 1872. — Half a drop of cobra-poison diluted with
■^ c.c. of water was injected under the skin of a guinea-pig,
weighing about 450 grammes (1 lb.).
At 12.13.15 the injection was made. Immediately the animal
became restless and cried constantly.
At 12.15 twitching movements began in the limbs.
At 12.16 the animal was quiet, and would not move when
touched. It then became restless again, and remained so till
12.44.
12.44. The jugular vein was exposed, and | c.c. of the diluted
poison was injected into it (= -^ drop of poison).
In less than 30 seconds the animal appeared to be dead.
The thorax was opened, and the heart found to be motionless
and the walls of all its cavities firmly contracted. The lungs
were ecchymosed.
12.55. Electrodes w^ere inserted into the spinal cord, and an
interrupted current passed through it. Whenever the current
passed, the legs of the animal jerked vigorously.
The blood which was collected from the large thoracic vessels
formed a firm coagulum.
1.22. The cord was still irritable when excited by the induced
current.
Experiment LXX.
January 14th, 1873. — The heart of a frog was excised. It
beat 20 times in 1 minute. Several drops of cobra-poison were
(95) H 2
100 ON THE NATURE AND ACTION OF THE

then placed upon it, and it beat 24 times in 1 minute. When

seized with forceps and placed in cobra-poison it stopped in
systole ; but this might be due to the e£fect of the compression
by the forceps.
Experiment LXXI.
Frog's heart excised. Beats, 30 in the first minute, 34 in the
second.
Cobra-poison applied to it. It immediately stopped, and then
began again, but slowly and feebly. Then it beat 26 times per
minute, less strongly than before. It gradually recovered and
seemed little affected, but stopped about 10 or 15 minutes
afterwards.
Experiment LXXII.
A cannula was placed in the aorta, and another in the vena
cava of a frog. All branches were tied, the heart excised, and
placed in connection with H. P. Bowditch's apparatus for
keeping a stream of serum circulating through the heart and
recording its pulsation by means of a manometer on a revolving
cylinder. When fed with pure serum, the heart's contractions
were regular and strong ; but whenever serum containing dried
cobra-poison in solution (in the proportion of about 2 grains in
3 fluid drachms) was introduced into the apparatus the heart
stopped almost immediately. As will be seen from the accom-
panying tracing, it became partially contracted and gave one or
two feeble beats, but did not dilate, and then remained still, the
contraction, however, very slowly and gradually increasing.

Experiment LXXIII.
A cat was deprived of consciousness by a severe blow on the
head ; and a cannula being placed in the trachea, artificial
respiration was begun. The thorax was then opened and the
heart exposed. A solution of dried cobra-poison in water was
then injected into the jugular vein. At first the cardiac pulsa-
tions became much quicker, but they were also strong. They
next became very small and rapid. Lastly, the right ventricle
became much distended, and the heart stopped. The lungs
became contracted ; and when force was used to distend them
 POISON OF SOME INDIAN VENOMOUS SNAKES. 101

they did not expand equally, but became emphysematous in

spots, so that the exterior of the lung assumed a nodulated
appearance. When the right ventricle was punctured it con-
tracted firmly. No further contraction took place when it was
irritated by the direct application of a Faradic current. The
blood coao^ulated.

lliese tracings were ubtaiued from a frog s heart hy means of a small

mercurial manometer connected with the aorta. The tracings all read from
right to left,
1. Tracing obtained from the heart supplied with pure serum by means of a
tube in the vena cava.
2. Tracing of the same kind, with the addition of the line A, which indicates
the zero of the mercury. The tracing B, given by the heart, sinks down to zero
during each diastole.
3. Tracing given by the heart after it had been supplied with serum contain-
ing a small quantity of cobra-poison in solution. The heart makes a few
ineffectual attempts, but can neither contract nor relax, and remains still, in a
condition midway between complete systole and complete diastole. The line A
is the zero to which B would sink if the heart relaxed completely during
diastole.

Experiment LXXIV.
A cannula was placed in the carotid of a dog and connected
with a kymographion.
Mean blood-
pressure. Pulse per
Time. millims. minute.
1.36 150 144 Injected some cobra-poison dis-
solved inwater into the sciatic
vein. The pressure rose to
165 165, and then sank in 7
135 seconds to 135.
1.45 50
102 ON THE NATURE AND ACTION OF THE

Mean blood-
Pulse per
Time. pressure.
millims. minute.
1.48 57 ...
Faeces passed. A clot formed
in the cannula and had to be
removed.
1.55
About 1.58 70 ...
1.58i 80 Injected some more poison.
20
1.59 55
1.59-J ... Clot again formed.
70
2.2
2.7 75
85
2.10 85
2.16 80 Legs loosened, but the animal
did not move. Convulsive
movements occurred almost
2.17J immediately afterwards.
Cornea still sensible.
2.18 85 Convulsive movements.
2.19i 90 ... Convulsions.
80 80 No movement.
2.201
2.201 90 64 The pulse here suddenly
changed from 80 to 64, and
at the end of every third beat
the pressure sank 25 millims.,
while at each of the others it
only sank 5 millims.
2.21 J 98 ... Height of each single pulse-
wave is now 10 millims. in-
stead of 5, and every now and
then it sinks 30; but the
number of beats after which
it sinks is not now so regular.
2.21f 100 64 Convulsions.
2.22-J 105 ... There were now 8 pulsations,
and then an interval of 6
seconds, during which the
pressure went down to 43
 POISON OF SOME INDIAN VENOMOUS SNAKES. 103

Mean blood-
pressure. Pulse per
Time. millims. minute.
milliras. Five beats more
raised it to 120. Height of
each pulse- wave about 15
millims.
2.29 30 ... Tlie pulse has been getting
smaller and smaller, and the
intervals longer and longer ;
it is now imperceptible.
2.30 30 ... The pressure still seems at 30,
notwithstanding the imper-
ceptibility of the pulse.
2.45 ... ... The heart was cut out. It still
contracted when irritated.

The injection of cobra-poison here caused a diminution of

the blood-pressure at first ; but a further injection again raised
it. In the latter part of the experiment there is not the
slightest trace of failure of the heart's action, but, on the
contrary, every evidence of powerful action. When the
respirations failed, the heart became slow from irritation of
the roots of the vagus by venous blood ; and the pulsations
were gradually weakened by the same condition. The fact that
the blood-pressure sank slowly and did not fall below 30, even
after the heart had almost entirely ceased, shows that the
arterioles were much contracted.

Experiment LXXY.
A cannula was placed in the carotid artery of a rabbit and
connected with a kymographion.
The blood-pressure was 75 mm. of mercury. One cubic
centimetre of a 2-per-cent. solution of cobra-poison was injected
into the jugular vein. Almost immediately the animal began
to struggle, and the pressure rose to 95. It remained at this
for a minute and then fell. The float unfortunately stuck, and
the curve it should have described in falling v/as consequently
lost. On again getting the instrument to work, the pressure
104 ON THE NATURE AND ACTION OF THE

was found to be 25 ; and this continued, although the heart

had ceased to beat and the thorax was opened. On cutting
across the aorta, the pressure fell to zero, showing that it had
not been due to any clot in the vessel.
In this experiment the poison seems to have caused tetanic
contraction of the heart, and also of the arterioles. The
permanence of the pressure at 25, notwithstanding the stoppage
of the heart's action, can only be ascribed to contraction of the
arterioles preventing the escape of blood from the arterial into
the venous system.

Excretion of Snake-poison.
We have made only one or two experiments, ourselves, on
the excretion of cobra-venom ; but, from the data afforded by
the experiments and observations of others, we consider that it
is excreted by the kidneys and mamijiary glands, and probably
also by the salivary glands and mucous membrane of the
stomach. A case reported by Mr. Shircore, of Calcutta, in
which an infant, suckled by its mother after she had been
bitten by a snake (species unknown), died in two hours after
it had partaken of the milk, shows that the poison is excreted
by the mammary glands, and with considerable rapidity ; for
the child took the breast before any marked symptoms had
occurred in the mother.* Its excretion by the kidneys appears
from an experiment of Mr. Eichards, of Balasore, who found
that some urine from a dog poisoned by the bite of a sea-snake
{Enhydrina hengalcnsis) killed a pigeon in 22 hours after being
hypodermically injected.t Some saliva, which we obtained
from the submaxillary gland of a dog poisoned by cobra- venom,
had no effect when injected under the skin of the thigh of a
lark ; but Mr. Eichards found that 1 drachm of the greenish
liquid which flowed from the mouth of a dog poisoned by
cobra- venom killed a pigeon in two hours. As this fluid
flowed constantly from the mouth, and the animal was paralysed
and motionless, it seems probable that, notwithstanding its
colour, it was saliva and not bile.
* Thanatophidia, p. 43.
t Indian Medical Gazette, May 1, 1873, p. 19,
 POISON OF SOME INDIAN VENOMOUS SNAKES. 10;")

As the poison-glands of the snake are modified parotid

glands, we should naturally expect the poison to he excreted
by the salivary glands ; and we think it possible that the
immunity which poisonous snakes enjoy from the effects of
their own poison or that of another species (an immunity
which is not shared by innocuous serpents, nor even by small
individuals of a venomous species poisoned by a large dose of
venom) may be due, at least in some measure, to their power
of excreting the inoculated venom through their own poison-
glands. We have, however, had no opportunities of trying
whether venomous serpents, after extirpation of their poison-
gland, succumb to the bite of others in the same way as
innocuous ones.

On the Means of preventing Death from the Bites of Venomous

Snakes.
In the case of all poisons, snake-venom included, there is
a dose which is insufficient to kill ; and animals may recover
from it even after the characteristic symptoms of the poison
have been distinctly manifested.
It has been clearly shown by Hermann that the real dose of
any poison, or, in other words, the quantity which is actually
circulating in the fluids and operating on the tissues of the
body, depends on two factors, viz., the rapidity with which it
is absorbed and the rapidity with which it is excreted. If
absorption goes on more rapidly than excretion, the poison
accumulates in the blood and exercises its lethal action ; while
the quantity in actual circulation may be reduced to an infinite-
simal amount and deprived of all power for evil, if the excretion
can keep pace with, or go on more rapidly than, the absorption.
Thus it is that curare kills an animal when introduced into
a wound; for the poison is absorbed from the wound more
rapidly than it can be excreted by the kidneys. If placed in
the stomach, curare has usually no apparent action whatever ;
for it is excreted in the urine as quickly as it is absorbed by
the gastric walls. But if absorption be quickened by increasing
the quantity administered and giving it on an empty stomach,
curare will have the same effect as when it is placed in a wound
106 ON THE NATURE AND ACTION OF THE

or injected into the circulation. A like result is obtained by

arresting its excretion, either by ligaturing the renal vessels or
extirpating the kidneys. Snake-vcnoni is also poisonous when
absorbed by the mucous membrane of the stomach.
On the other hand, when we wish to prevent the accumula-
tion of a poison in the blood and thus to arrest its action, we
must either lessen its* absorption, quicken its excretion, or
combine the two means.
In the case of curare the former of these is sufficient ; and all
the bad effects of the introduction of this poison into a wound
may be prevented by applying a ligature between the wound
and the heart, and only loosening the bandage occasionally, for
an instant or two at a time. The same obtains in snake-
poisoning. In this way only a little of the' poison is absorbed
each time the ligature is slackened, and this is excreted by the
kidneys before another quantity is absorbed. If the poison
can be removed from the wound itself by other means, instead
of making the whole of it pass through the circulation, the
danger it causes will, of course, be sooner over. Our power to
quicken excretion is, in most cases, much less than that to
retard absorption ; and it is therefore on the latter that we
mainly rely in cases of poisoning in general, as well as snake-
bites in particular.
The various methods of mechanically arresting the intro-
duction ofthe virus, by excision, cautery, and chemical agency,
have been fully discussed in the Thanatopliidia of India ; and
we purpose now to consider its excretion or removal from the
organism.
Before doing so, however, we must inquire whether its
removal is likely to be of any service or not ; for, as we have
already pointed out in our previous communication, the action
of the poison may be of two kinds :— 1st. It may resemble
curare in destroying the power of the nervous system so long as
it is present in the blood, but leaving it in a condition to resume
its functions as soon as the poison has been removed. 2nd. Its
action may be identical with, or similar to, that of a ferment,
decomposing or altering the nervous and muscular tissues
in situ (in somewhat the same way as the pancreatic or gastric
 POISON OF SOME INDIAN VENOMOUS SNAKES. 107

ferments would decompose them if they had been placed in the

intestinal canal), and thus rendering them utterly incapable of
ever again performing their functions.
If the action of the poison is of the latter kind, no treatment
can be expected to be of any avail if the dose has been large ;
but if it is of the former, we may still entertain a reasonable
hope of averting a fatal result, even when the dose of venom
has been large.
We have shown in our previous communication, that, by
means of artificial respiration, life may be prolonged for many
hours, and time thus afforded for the excretion of some of the
poison ; but the means at our disposal have not enabled us to
maintain respiration sufficiently long to show us whether the
nervous and muscular systems regain their function after the
excretion of the poison has proceeded far enough. The experi-
ments of Mr. Vincent Eichards, and of a committee appointed
by the Government of India in Calcutta, at our suggestion, to
investigate the use of artificial respiration in death by snake-
bite, being performed under more favourable auspices, have
afforded us the data which we were unable to obtain from our
own. In one instance, a dog was bitten by a sea snake
{Enhydrina hengalensis), and, two hours afterwards, died in
convulsions. Artificial respiration was commenced ; but, four
hours afterwards, the application of a galvanic current caused
no muscular contractions ; the eyes were dry and glazed, and
the body was cold. Next morning, about sixteen hours after
the apparent death of the animal, reaction commenced ; the
application of a galvanic current again caused movements of
the body and expulsion of urine, and the bowels acted spon-
taneously. Infive hours more reaction seemed established and
went on increasing; the animal appeared as if it would
recover ; the eyes lost their glazed appearance, tears were
secreted, and a greenish -looking fluid flowed from the mouth ;
reflex action became re-established, the eyelids closing when
the cornea was touched or when water was poured into the
eye. Attempts to swallow were made when water was poured
into the mouth ; and the application of a pan of hot charcoal
to the chest caused convulsive movements all over the body ;
108 ON THE NATURE AND ACTION OF THE

and these also occurred spontaneously. The animal also

became more or less sensible, and the eyelids twitched when
the finger was merely brought near the eye.
These phenomena show that the muscles, the motor nerves,
the secreting nerves, the spinal cord, and the cerebrum had all
recovered their functions to a certain degree, after it had been
completely abolished for sixteen hours. This, we think, would
not have been the case had the poison acted by decomposing
the tissues in the manner of a ferment ; and we are therefore
inclined to hope that, like curare, it acts only while present
in the system, and that its injurious effects may be arrested by
its removal.
Notwithstanding the fair promise of recovery which the use
of artificial respiration gave in this instance, the heart became
weaker, and the animal died 24 hours and 35 minutes after
its first apparent decease. Nor has the Committee been more
successful in its further experiments, although life has been
prolonged for even 30 hours. This result shows that, although
artificial respiration may still prove useful in sustaining life
and affording time for the use of other measures, it alone is not
likely to be of much service in preventing death from snake-
bite, except in those cases where the quantity of poison is just
enough to kill and no more.
It is evident from the length of time during which life may
be maintained without the animal ultimately recovering, that
the excretion of the poison is very slow ; but we at one time
thought to quicken it by the employment of diuretics and
sialogogues, and to prevent reabsorption by draining off the
urine and saliva constantly. We also proposed to wash out
the stomach from time to time, in order to remove any poison
which might be excreted through the gastric walls, keeping it
partially filled with milk or other nutrient fluid during the
intervals, in order to sustain the strength of the animal.
We are by no means certain that some of these methods
may not prove useful adjuncts ; but as our hope of stimulating
excretion, by the salivary glands at least, has been much
lessened by our discovery that the poison paralyses the nerves
of secretion, we are inclined to think that, perhaps, the readiest
 POISON OF SOME INDIAN VENOMOUS SNAKES. 109

method of removing the poison from the body may be to allow

it to flow out along with the blood in which it is circulating,
and supply the place of the poisoned blood thus withdrawn by
means of transfusion.
The greater part of the poison present in the system is
probably contained in the blood, and only a small proportion
in the tissues ; for one of us (Dr. Fayrer) has found that a few
drops of the blood of a dog killed by the bite of a cobra or
Dahoia caused death in 75 minutes, when injected into the
thigh of a fowl (Thanatophidia, pp. 80, 83, 119, 120). By
removinoj as much blood as could be taken without endangering
the life of the animal, a great part of the poison would be
withdrawn from the system ; and, probably, any harm from
the copious bleeding would be prevented by transfusing fresh
blood immediately afterwards.
We have tried one or two experiments with transfusion ; but
they have hitherto been unsuccessful.
AVe are therefore by no means confident that death may be
prevented by the combined use of artificial respiration and
transfusion ; but we think that they present some chance of
success, and that, at all events, the suggestion is justifiable on
scientific and rational grounds.
The treatment of animals poisoned by cobra- virus by the
hypodermic injection of liquor ammonise has been frequently
tried in India by one of us (Dr. Fayrer) (vide Thcmat., pp. 89
et seq.\ and also by Mr. Eichards, of Balasore, and by ourselves
again in London, on several occasions.
The alkali has been administered internally, injected into the
areolar tissue, and also into the veins, over and over again ;
but no benefit has resulted. The objection has been made that
experiments of this nature, made on animals, are not conclusive
in reference to the probable action of the agent experimented
with on human beings; but this objection can hardly be
considered valid in a physiological point of view.
At any rate the trials that have been made of this mode of
averting the lethal effects of the poison, in India by Dr. Hilson,
Civil Surgeon of Moradabad, do not atibrd any indication that
the intravenous injection of liquor ammonise was followed by
110 ON THE NATURE AND ACTION OF SNAKE- POISON.

any diminution of the effect of the poisons, the man in both

cases having died* {vide Indian Medical Gazette, October, 1873).
The same may be said of other reputed antidotes, such as :—
Tanjore pill and other preparations of arsenic ; the hypodermic
injection of liquor potassse, quinine, ipecacuanha, Aristolochia
indica, and a variety of other drugs, generally of a vegetable
nature, and enjoying a large amount of popular confidence : all,
when brought to the test of carefully conducted experiment,
failed, as might have been expected, to give any favourable
result.
It seems almost unnecessary to allude to the so-called snake-
stones; they are powerless for good or evil. They have also
enjoyed much confidence ; but when submitted to the test of
impartial experiment and observation, their virtues prove as
unreal as those of the antidotes above mentioned.
With reference to the mechanical methods of preventing the
entry of the poison into the circulation after a bite, we think
that the speedy application of an elastic cord (such as is used
in bloodless operations) round the limb, combined with the
application of cups attached to an exhausting-syringe or pump,f
might be of advantage, and that it might be made of general
application in India.
* It is unnecessary to occupy time by describing iu detail the various sub-
stances (animal, vegetable, and mineral) that have been administered as
antidotes. Particulars may be found in the Thanatophidia, where the details of
experiments conducted for the investigation of their actions are recorded.
t Such an apparatus has now been constructed.
ON THE NATURE AND PHYSIOLOGICAL
ACTION OF THE CROTALUS-VOISO^ AS
COMPARED WITH THAT OF NAJA
TRIPUDIANS AND OTHER INDIAN
VENOMOUS SNAKES ;
ALSO

INVESTIGATIONS INTO THE NATURE OF THE INFLUENCE

OF NAJA- AND CROTALUS-POliSO}^ 0"^ CILIARY AND
AMCEBOID ACTION AND ON VALLISNERIA, AND ON
THE INFLUENCE OF INSPIRATION OF PURE OXYGEN
ON POISONED ANIMALS.

By T. Laudee Brunton, M.D, F.E.S., Sc.D., M.RC.P., and

J. Fayrer, C.S.I., M.D., F.RC.P. Lond, F.R.S.E.,
President of the Medical Board at the India Office.

(Reprinted from the Proceedings of the Royal Society, No. 159, 1875.)

In our former papers we described the general phenomena

accompanying the physiological action of cobra- and Daboia-
poisons on warm-blooded animals, reptiles, fishes, and inverte-
brata. We propose in this paper to compare with these the
action of the Crotalus-\\v\x.^ in its general effects on life, on the
functions, organs, and tissues, and especially as it affects the
blood and vessels as regards a marked influence in causing
haemorrhages and extravasations of blood generally and locally ;
and, further, to examine the action of snake-poison generally on
ciliary and amoeboid movements — or that which represents its
action on contractility, apart from that which is caused through
the medium of the nerve-centres and nerve-distribution.
It appears that there is little difference between the physio-
logical effects of the crotaline or vipurine and the colubrine
virus. The mode in which death is brought about is essentially
the same in all ; though there are evidences, even when allowing
for individual peculiarities, that the action is marked by some
112 ON THE NATURE AND ACTION OF THE CROTALUS-POISON

points of difference sufficiently characteristic to require notice

in detail.
We have already expressed our belief that death is caused by
the cobra-, Daboia-, and Hydrophis-^oimn, 1st, through its action
on the cerebro-spinal nerve-centres, especially on the medulla,
inducing paralysis of respiration ; or, 2nd, in some cases (where
the poison has entered the circulation in large quantities and
has been conveyed more directly to the heart) by arrest, tetani-
cally in systole, of cardiac action, probably owing to some action
on the cardiac ganglia ; 3rd, by a combination of the two
previous causes ; 4th, by a septic condition of a secondary
nature, and which, being more essentially pathological in its
bearings, the details were not considered suitable for discussion
here.
There is reason to believe that death is caused in the same
way by the Crotalns-^^oimn also ; and it appears, from the
experiments recently performed in Calcutta, by Dr. Ewart and
the members of the Committee appointed by Government, upon
Fseudechis porphyriacus, or the black snake, and Hojplocephalus
curtus, or the tiger-snake of Australia, that their virus causes
death in the same manner. These reptiles had been sent from
Melbourne to Calcutta for the purpose of investigation and com-
parisonVide
.( Committee's Eeport, pp. 58 et seq., Appendix.)
But though the actual cause of death is essentially the same,
the phenomena which precede and accompany it differ in some
degree according to the nature of the poison, the quantity and
site of the inoculations, and the individual peculiarities of the
creature inoculated, as may be seen in the experiments herewith
recorded.
The condition of an animal poisoned by the rattlesnake-
venom, then, essentially resembles that of one subjected to the
influence of the colubrine or viperine poison of Indian
snakes :—
Depression, hurried respiration, exhaustion, lethargy, un-
consciousness, nausea, retching, and vomiting {vide experiment
on cat. Experiment IX).
Muscular twitchings, ataxy, paralysis, and convulsions (the
latter probably chiefly, though not entirely, due to circulation
 AS COMrARED WITH THAT OF OTHER VENOMOUS SNAKES. 113

of imperfectly oxygenated blood, the result of impeded

respiration), and, finally, death.
Haemorrhages or hsemorrhagic extravasations and effusions,
both local and general, occur m all vaiieties of snake-poisoning.
But we observe (and in this our observations are in accord
with those of Weir Mitchell) that there is a greater tendency to
both local and general liajmorrhage and extravasation of blood
and of the colouring-matter of the blood, especially as observed
in the peritoneum, intestines, and mesentery, and also probably
to a more direct action on the cord (vide Experiments I, III, V,
VI, VII, IX, XI, XIV, XV), than in poisoning by either cobra
or viper {cide Experiments IV, VII, XIII, XVI, XVII, XX).
The viscera and other tissues, after death, are found congested
and ecchymosed, and in some cases to a great extent, seeming
to show that either a preternatural fluidity of blood or some
important change in the vessels, favouring its exudation, has
occurred.
But with regard to tlie blood itself, we have observed that it
does form a coagulum after (ieath, generally, if not invariably ;
as we have noted to be the case, tliough not to the same extent,
in the blood of animals that have succumbed to the Dahoia-
virus.*
With reference to the coagulation or non -coagulation of the
blood in cases of snake-poisoning, we observe that the following
conclusions have been arrived at by Mr. Bichards and the
Calcutta Committee {vide p. 45 of their Beport).
"We now propose to deal with the ph}sical clianges produced
by snake-poisoning on the blood. Erom observations which
have been made by Mr. Bichards and ourselves, we have arrived
at the following conclusions :—
'•' The blood appears to remain fluid after death under the circum-
stances noted below :
" 1st. When a large quantity of the cobra-poison has been
directly injected into the circulation, as, for example, into an
artery or a vein.f

* In Dr. Fayrer's Indian experiments the blood of animals dead from Bahoia-
poison nearly always remained duid after death,
t This is not always so. — J. Fayrer.
(95) 1
114 ON THE NATURE AND ACTION OF THE GROT ALUS-POISON

" 2nd. In cases where animals or men have been poisoned by

the bite of vipers, such as the Russell's viper.
" 3rd. In all cases of snake-bite, whether from the poisonous
colubrine or viperine genera, in the human subject.*
" The blood undergoes either partial or complete coagulation
under the folloiving conditions : —
" 1st, When a small quantity only of the cobra-poison has been
injected into a vein or an artery.
" 2nd. In cases where the lower animals have been bitten by
the cobra.
" Why the admixture of a large and quickly fatal injection of
the cobra-virus into the circulation of animals should produce
comparatively permanent fluidity of the blood or interfere with
its ordinary coagulability soon after removal from the body or
after death, and why the injection of a smaller and more slowly
fatal quantity should interpose no obstacle to its speedy coagu-
lation, are questions extremely difficult to account for or
explain. We can only state the fact that, in the one case,
coagulation occurs speedily, and in the other this coagulation is
retarded or altogether prevented by some cause at present
unknown."
The following experiments were made on the physiological
action of the virus of the rattlesnake, with the view of com-
parison with that of the cobra and Dahoia.
We are indebted to Dr. Weir Mitchell, of Philadelphia, for a
supply of the virus. He was good enough to send about
six grains of the dried poison of Crotalus — the species not
named, but it is believed to be of Crotalus durissus.
The dried poison supplied is said to be about 6 J years old,
and was dried in July or August at the natural temperature,
and has since then been preserved in a phial. It was tried by
Dr. Mitchell, and found active three years ago.
It has the appearance of fractured fragments of dried gum-
arabic, of rather a darker yellow colour, but otherwise resembling
the dried cobra-virus sent from Bengal.

* Not always so. — J. Fayrer.

 AS COMPARED WITH THAT OF OTHER VENOMOUS SNAKES. 115

Experiment I.
June 9th, 1874. — 0*015 gramme of the dried Crotalus-'poison
diluted with 1 c.c. of distilled water was hypodermically injected
into the thigh of a full-grown guinea-pig at 11.30 a.m.
Restlessness and muscular twitchings of the body generally
soon commenced ; these passed away, but the animal became
sluggish, in which condition it remained all night, and died at
about 9 A.M. the next morning.
The injected limb became much swollen, infiltrated, and
discoloured with sanguineo-serous effusion.
The intestines were not ecchymosed ; there was much san-
guinolent fluid and also blood effused into the abdominal areolar
tissue.
No convulsions were observed ; but as the animal was not seen
during a short time previous to death, they cannot be said
positively not to have occurred ; nor is it known if the heart
ceased to beat at the moment when apparent death took place.

Experiment II.
A few drops of watery solution of Crotalus-i^oison, of same
strength, were injected under the skin of a guinea-pig's thigh at
12.16 noon.
12.17. Marked twitchings of head and hind legs, very similar
to those produced in some of the cases of cobra-poisoning.
12.18. Hind leg (poisoned one) weak.
12.20. Twitchings much increased, now mainly in head and
neck, not so much in hind legs.
12.28. Guinea-pig quiet, but with occasional twitchings ;
sluggish and disinclined to move.
1.30. Sluggish in moving; can still move about, though
disinclined to do so. The punctured thigh is very blue.
The rest of the notes of this experiment were lost.
The animal died.
Experiment III.
June 10th, 1874. — J of a grain of Crotalus- and J of a grain of
cobra-poison were carefully weighed and diluted, each with
(95) I 2
116 ON THE NATURE AND ACTION OF THE OROTALUS-POISON

10 drops of distilled water. Two full-grown guinea-pigs of equal

weight were then selected.
The solution of Crotalus-'j^oison was injected into the peritoneal
cavity of guinea-pig No. 1 at 1.52 p.m.
1.55. Muscular twitchings of head and neck.
2 P.M. Startings and twitchings continue.
It gives faint squeaks occasionally, as though the sudden
startings, which occur at intervals of 5 or 6 seconds, cause pain.
2.5. Twitcliings continue.
2.8. Very restless ; twitchings going on, but no paralysis yet.
2.17. The same.
2.25. Eestless and weaker, but still moves freely on being
roused.
2.42. Sluggish ; drags the hind legs.
2.58. Weaker ; rolls partially over on one side, but can run
when roused.
3.3. Lying on side, but can be roused ; is partially paralysed
in hind legs. Eespiration abdominal and hurried.
3.5. Nearly quite paralysed; is roused with difficulty.
3.7. Can still be roused. Abdomen distended and painful;
cries out when it is touched, as though peritonitis were
setting in.
3.12. Can be roused with difficulty; respiration hurried;
convulsive movements of fore legs and neck. Can still stagger
for a few paces, but co-ordination of muscular power much
diminished.
3.30. In violent convulsions.
3.38. Convulsions continue.
3.45. Quiet. Paralysed ; but reflex action still continues.
3.55. Dead in 2 hours and 3 minutes.
3.56. Electrodes in cord cause twitching of muscles of the
back, and very slightly in those of the legs; the cord was
evidently all but paralysed. Muscular fibre contracts freely to
direct stimulus of current. The intestines were ecchymosed
and congested. There were effusions of red serum into the
peritoneal cavity, and much ecchymosis of peritoneum and
subperitoneal and intra-muscular areolar tissue. Peristaltic
action continued faintly.
 AS COMPAEED WITH THAT OF OTHER VENOMOUS SNAKES. 117

4 P.M. The heart has ceased to contract 4 minutes after

apparent death ; it continued to contract, especially the auricles,
for part (not the whole) of the time.
The blood removed from the heart-cavities and vena cava
rapidly formed a firm coagulum in a glass receiver.
The electrodes applied to the sciatic showed that the nerve-
trunk, as well as the spinal cord, was paralysed.

Experiment IV.
Guinea-pig No. 2, an albino, had the |-grain cobra-virus
solution injected into its peritoneal cavity at 1.56 p.m.
It immediately became much excited.
1.57. Is now quite tranquil.
2. Sluggish. Does not twitch as guinea-pig No. 1 did.
2.4. Started and squeaked slightly, as though in pain, but no
twitching.
2.5. Slight twitching generally. Paralysis and ataxy com-
mencingdrags
; its legs with difficulty.
2.9. Sharp twitchings of head and neck.
2.12. Subsided on to the belly; head fallen over; crawls
with difficulty ; is very feeble, almost paralysed. The albino
eyes have a heavy dull look ; lost their bright pink.
2.14. Convulsed.
2.15. Pteflex action ceased. Apparently dead, but heart can
still be felt beating. Occasional convulsive twitchings of
lower lip.
2.16. Dead in 21 minutes.
2.17. All movements have ceased. Heart had ceased to
contract, except slight flickering movements of auricles.
2.20. Electrodes in cord. Spinal cord and nerves paralysed ;
muscles contract freely to direct stimulus of current. Heart
distended with blood. Blood, when removed, formed rapidly a
firm coagulum. Intestines, peritoneum, and subperitoneal
areolar tissue congested and ecchymosed. Sanguinolent effu-
sions into peritoneum, but not so well marked as in the Crotahcs-
poJsoning. Peristaltic action of bowels ceased rapidly.
The results of these two experiments show, so*far, that the
action of the cobra-poison is more energetic than that of the
118 ON THE NATURE AND ACTION OF THE CROTALUS-POISON

rattlesnake. Both were watery solutions of exactly the same

quantity of the dried virus ; but it is to be borne in mind that
that of the rattlesnake was 6J years old, wliile that of the
cobra was only 1 year old.
The guinea-pigs were both full-grown and of the same size,
yet one succumbed in 20 minutes to the cobra-poison, while the
other survived the inoculation of the rattlesnake-poison for
2 hours and 3 minutes.

There were- no very marked differences' in the action of the

poison in these two cases, except in the energy with which the
cobra exceeded the Crotalus.

Crotalus. Cohra.
Twitchings ; restless ; squeaks ; Twitchings ; excitement ;
sluggish ; ataxy ; paralysis. squeaks ; sluggish ; ataxy ;
Hurried respiration. Peri- weakness; paralysis. Con-
tonitis. Convulsions. Death vulsions. Death in 20
in 2 hours 3 minutes. Co- minutes. Spinal cord and
agulated blood. Ecchymosis nerves paralysed. Muscles
and extravasation of serous irritable. Heart distended,
effusion well marked. Cord Blood coagulated. Ecchy-
paralysed. Muscles retain mosis. Congestion less than
irritability. in Crotalus.

Experiment V.
June 10th. — A grain of Crotalns-^oimn diluted with water
was injected into the peritoneum of a full-grown guinea-pig at
2.40 P.M. Twitchings began almost immediately.
3.3. Eestless ; startings ; staggers on hind legs.
3.20. Very weak, especially in hind quarters. General
paralysis setting in. Abdomen distended and very tender.
3.30. In convulsions. Still feels when the abdomen is
touched.
3.37. Paralysed, but feels the touch. Eeflex well marked.
3.45. Apparently dead in 65 minutes.
3.48. Cavities opened. Auricles flickering. Blood from heart
and great vessels coagulated firmly. Abdominal cavity and
areolar tissue and subperitoneal tissue inliltiated with bloody
 AS COMPARED WITH THAT OF OTHER VENOHtOUS SNAKES. 119

serum. Much ecchymosis of peritoneum and intestines, but not

of lungs. Cord and nerves paralysed. Muscles contract
vigorously to induced current.

Actio7i of Cvoi'dX\\s,-poison on Babbit,

Experiment VI.
i of a grain (0*0 15 gramme) of the same Croialus-i^oison,
dissolved in 1 c.c. of water.
The jugular vein of a large white rabbit was exposed and the
above solution was injected into it at 1.50 p.m.
At 1.51 violent convulsions, with opisthotonos.
At 1.53 apparently quite dead. Artificial respiration com-
menced innnediately. Heart acting still, though feebly and with
irregular flickering contractions. Spinal cord exposed. Elec-
trodes applied ; no reaction.
2.12. Heart still contracting feebly.
2.15. Faint contractions of heart still observable. Ventricles
punctured, and blood withdrawn. Peristaltic action has ceased.
2.20. Feeble cardiac movements continue.
2.21. Heart has now ceased. Muscles react to direct current.
Death caused by rapid paralysis of medulla and cord. The
blood taken from the heart and great vessels did not coagulate.
At 4 P.M. it was still fluid, though very florid in colour.
Examined under the microscope nearly two hours after
apparent death, the white corpuscles appeared natural ; the red
corpuscles not in rouleaux, and very much crenated, though a
few retained their natural contour.
The blood was neutral to test-paper.

Experiment VII.
June 17th. — I of a grain (0'015 gramme) of dried cobra-poison,
dissolved in 1 c.c. of water, was injected into the jugular vein
of a large white rabbit, of the same size as in the previous
experiment, at 2.55 p.m.
The rabbit passed at once into violent convulsions, and was
apparently dead before it could be removed from the board,
within one minute. The cord was immediately exposed,
120 ON THE NATURE AND ACTION OF THE CROTALUS-POISON

artificial respiration having also been begun. Electrodes

applied, with strong current ; no reaction ; the cord was
perfectly paralysed.
Thorax examined at 2.59. Heart had ceased to contract.
Ventricles moderately contracted. Auricles distended with
blood. Phrenic irritated, quite paralysed. Diaphragm, when
directly irritated by current, contracts very faintly, whilst the
neighbouring muscles contract vigorously. Peristaltic action
goes on. Electrodes applied to vagus appear to accelerate
peristaltic action ; applied to splanchnic, they diminish it.
3.7. Ventricles of heart have now contracted firmly.
3.15. Blood taken from heart and great vessels has coagulated,
but not firmly. The clot is small, and the serum very red.
3.15. Electrodes to sciatic; no reaction. Blood examined
under microscope ; no aggregation in rouleaux, no crenation of
corpuscles. Blood neutral to test-paper.
We have in former papers remarked that when the cobra-
poison was injected into the jugular vein directly and caused
almost immediate death, that the fatal result was due to
cessation of the heart's action by arrest in systole, and such was
partially the case in the last experiment (VII), made for the
purpose of comparison with Crotalus ; but in Experiment VI
death was not so caused, for the heart continued to contract
for about 28 minutes after apparent death, which was probably
due to the sudden and total annihilation of the functions of
the medulla and cord, no reaction to a strong current occurring
when the electrodes were applied immediately after apparent
death.
In this instance of Crotalus-^oi^oTimg it is also to be remarked
that the coagulability of the blood was destroyed, whilst in that
by cobra- virus it was only partially so.
It appears from the results of this experiment that the direct
inoculation of large doses of the virus, whether viperine or
colubrine, into the circulation have the power in some cases
of annihilating almost instantaneously the irritability of the
cord and medulla, as in others they have of arresting the heart's
action.
 AS COMPARED WITH THAT OF OTHER VENOMOUS SNAKES 121

Experiment VIII.
June 17th. — Ten drops of the blood of the rabbit described
in the last experiment, poisoned by Crotalus-virns, were injected
into a guinea-pig's thigh at 3.40 p.m.
The guinea-pig was not apparently affected constitutionally
by the poisoned blood. It was alive the next morning ; but
the leg was swollen and discoloured. It ultimately recovered.

Experiment IX.
June 24th, 1874. — A full-grown cat was chloralised at
1.20 P.M. J of a grain of Crotahos-i^oison, diluted with 1 c.c. of
water, was injected into the jugular vein. The respirations
were immediately quickened.
1.21. Twitching of muscles generally.
1.22. Efforts to vomit. Forcible extension of limbs.
1.24. Hurried respiration and retching. Eeflex action
perfect.
1 30. Muscular twitching and tetanic stretching of limbs.
Efforts to vomit continue. Micturition. Kolls over on the
ground. ^
1.34. Ataxy. Staggers when walking, which it can only do
for a few paces. Peculiar twitching of diaphragm ; not syn-
chronous with respiratory movements. Kolls over on its side.
2 P.M. In the same state.
2.8. Injected |- of a grain more of the poison into the same
jugular vein. The animal immediately got up and walked,
comparatively steadily, for several paces, as though it had been
stimulated, and then rolled over.
2.16. Twitching of diaphragm continues at the rate of 150
per minute.
2.18. Again got up and walked for a few paces ; but it is
gradually becoming more paralysed.
2.44. Violent tetanic spasms of limbs. Eeflex action
diminished.
2.46. Eeflex action gone from eyes. Deep sighing respiration.
2.47. Convulsions. Death. Body opened immediately.
Lungs deeply congested and much ecchymosed. Deep red
122 ON THE NATURE AND ACTION OF THE CROTALUS-POISON

gelatinous effusion all about the roots of the lungs. Heart

contracting. Electrodes applied to phrenic caused vigorous
contraction of diaphragm.
2.50. Heart ceased to contract three minutes after respiration
had ceased.
2.52. Electrodes in cord ; do not cause contraction of limbs.
2.54. The sciatic nerve, when irritated, conveys impressions ;
muscles of legs contract. Blood from the heart and great vessels
did not form a coagulum, and remained permanently fluid. Red
corpuscles of blood were much crenated.
Death in this case appeared to be caused through the
medulla.
Experiment X.
June loth, 1874. — Action of Co^otalus-'poiBon on the frog.
A frog's hind leg was ligatured, excluding the sciatic nerve.
A solution of Crotalus-ipoisoii was injected into the lymph-sac
at 12.32 P.M.
2.30. Sluggish, but not otherwise affected.
3.15. In the same condition.
June 16th. — 12.3, noon. Sluggish, but can still move.
June 17th. — Found dead this morning early ; pupils con-
tracted.
Electrodes applied ; no reaction in either cord or nerves on
either side to the strongest current.
The frog may have been dead some hours.

Experiment XI.
June 15th. — At 3 p.m. same day a solution of Crotalus-Yenom
was injected into the dorsal lymph-sac of a frog, the aorta
having been previously ligatured, so as to prevent the poison
from affecting the trunks or peripheral extremities of the
sciatic nerves.
3.40. The frog seems quite unaffected.
June 16th. — 12.30, noon. Frog dead ; not rigid; mouth open.
Irritation of cord with strongest current does not cause
contraction of legs. Irritation of sciatic with coil at 24 causes
twitchings of gastrocnemius.
 AS COMPARED WITH THAT OF OTHER VENOMOUS SNAKES. 123

Neither of these two experiments give any definite results,

as the period intervening between death and examination of the
condition of the nerve-centres was not determined exactly.
The results of the following experiments show that the local
as well as the general effect of the cobra- and Crotahcs-i>oisons,
i.e. colubrine and viperine, is to cause hsemorrhage, ecchymosis,
and sanguinolent effusions into the areolar tissue, not only at
the seat of inoculation and its neighbourhood, but also in the
mucous membranes and other vascular parts. It is obvious
also that the Crotalus-ipoison acts more energetically in this
respect than the cobra-poison, and that this is perhaps one of
the most marked distinctions between them.

Experiment XII.
August 6th, 1874. — A cat was chloralised, and part of the
mesentery placed under the microscope on the warm stage.
Crotalus-i^oison, diluted with water, was then applied to the
mesentery, and its eff'ects watched. The white coipuscles were
observed to cling in quantities to the walls of the vessels, and
as the current of blood hurried through them, some masses of
pale matter like aggregation of white corpuscles were observed
to pass with the stream ; very soon, marked extravasation of
red corpuscles took place, and to the naked eye the mesentery
became discoloured by patches of ecchymosis in the course of
the small blood-vessels, like the foliage on the branches of a
tree.
There could be no doubt that the local action of the poison
had a marked effect in producing extravasation of blood.

Experiment XIII.
A similar experiment was repeated on another part of the
mesentery of the same cat with cobra-poison, exactly as the
Crotahis-^oison had been applied in the previous experiment.
This was carefully watched, but no extravasation took place;
there was a marked difference in the result of the application
of the two poisons, at all events as far as these two experiments
were concerned.
124 ON THE NATURE AND ACTION OF THE CROTALUS-POISON

Experiment XIV.
August 12th, 1874— A cat was chloralised at 2.30 p.m.
Mesentery exposed and placed under microscope on warm
stage.
Crotahcs-^oison applied to mesentery ; circulation soon dimi-
nished in some vessels but continued vigorously in others
Isolated extravasated patches soon made their appearance, of a
triangular form ; others followed and coalesced with these until
a network was formed in the course of the vessels all over the
field. The extravasation soon became general, the circulation
still continuing slowly.

Experiment XV.
A fresh portion of mesentery of same cat exposed. Intestines
becoming cold and circulation now very languid.
Cobra-poison applied.
No apparent effect produced, but the circulation is very
languid, indeed has almost ceased, so that the results of this
experiment are not conclusive.

Experiment XVI.
August 14th, 1874. — A cat was chloralised, part of mesentery
withdrawn, and placed under microscope on warm stage.
Dried cobra-poison dissolved in a salt solution, 0*75 per cent.,
applied to the mesentery at 4.10 p.m.
4.14. Circulation is languid, almost ceased in some vessels.
4.18. Slight extravasation taking place where the poison has
been in contact.
4.20. Extravasation rather more obvious.
4.35. Exposed another part of the mesentery ; examined the
state of the circulation before applying the poison. Blood
flowing languidly.
Poison applied at 4.37 ; at first it seemed rather to accelerate
the movement of the blood.
4.38. Circulation continues at same rate.
4.42. Same rate.
 AS COMPARED WITH THAT OF OTHER VENOMOUS SNAKES. 125

4.45. It becomes more languid.

4.48. Circulation has ceased, but yet there is no marked
extravasation.
Experiment XVII.
Another portion of the same mesentery had cobra-poison
applied, but after half an hour there was no sign of extravasation.

Experiment XVIII.
A fresh piece of mesentery exposed of same cat, and diluted
Crotalus-ipoison applied at 4.52 p.m.
The circulation was rather languid at the time, and apparently
became more languid.
At 4.58 no extravasation had taken place, the blood flowing
very languidly.
5.15. Circulation still going on, but very slowly; no extrava-
sation it
: soon after ceased.

Experiment XIX.
At 5.2Q p.m. a fresh portion of the mesentery was exposed ;
to one part cobra- and to the other Crotalus-'poison was applied,
and the effect was watched with the naked eye.
5.45. No extravasation visible.
At 6.15 p.m. slight extravasation equally visible on both.

Experiment XX.
August 25th, 1874. — At 2 p.m. a young cat was chloralised.
The mesentery was drawn out and a part treated with cobra-
poison, another part with Crotalus-'poison.
At 5 P.M. On examination, that under the influence of the
Crotcdus-\)oisoii was found deeply congested and reddened with
blood, extra vasated in the course of the small vessels, forming a
well-marked redness to the naked eye. Under the microscope
the red corpuscles were seen in numbers outside the vessels.
Circulation still going on vigorously. That part treated with
cobra-poison was barely altered, but on close examination slight
patches of extravasation were seen in the course of the vessels.
The difference was well marked between the two — the
126 ON THE NATURE AND ACTION OF THE CROTALUS-POISON

extravasation produced by Crotalus-wenom being well marked,

that by cobra-venom scarcely perceptible. In both cases the
microscope showed red corpuscles outside the vessels.
These experiments show that Crotalus-'poison causes haemor-
rhage and haemorrhagic effusions more than the cobra-poison
does.

The following experiments were made, at the suggestion of

Mr. Darwin, with the object of testing the influence of snake-
poison on ciliary action, especially in reference to its compara-
tive action on vegetable protoplasm, as will be seen by his
remarks.

Experiment XXI.

Influence of Cohra-poison on Ciliary Action.

June 29th, 1874. — Ciliated epithelium from the frog's mouth

was treated with a solution of cobra-poison and examined under
the microscope.
At 1.35 P.M., when examined, the action of the cilia was
vigorous.
At 1.45 it was much diminished.
At 1.55 it had entirely ceased.

Experiment XXII.
Ciliated epithelium placed under microscope ; one part was
treated with water, the other with the poisoned solution.
At 2.10 P.M. ciliary motion vigorous in both, perhaps more so
in that subjected to the poisoned solution.
2.18. Non-poisoned cilia active. Poisoned cilia very feeble.
2.20. Non-poisoned cilia still active. Poisoned cilia very
feeble.
2.24. Non-poisoned cilia active. Poisoned cilia very languid.
2.30 Non-poisoned cilia still active. Poisoned cilia have
entirely ceased to act.
It is evident from this that the poison first stimulates and
then destroys the activity of the ciliary action.
 AS COMPARED WITH THAT OF OTHER VENOMOUS SNAKES. 127

Experiment XXIII.
August 14th. — Frog's blood placed in salt solution, 0*75 per
cent., at 1.25 p.m. on warm stage, and then subjected to the
action of cobra-poison.
At first the amceboid movements of white corpuscles went on
vigorously. At 2 p.m. they had ceased, or very nearly so, in all
that appeared in the field.
2.30. All movement had entirely ceased. The red corpuscles
seemed more flattened, the nucleus more visible, and the edges
better defined, assuming a pointed and more oval form than
usual.
P^xperiment XXIV.
August 25th, 1874. — Newts' blood examined under ^ object-
glass on hot stage, white corpuscles moving slowly. Cobra-
poison applied, but no perceptible change observed.
The following communications were received from Mr. C.
Darwin on the action of some of the same cobra-poison ou
vegetable protoplasm :—
" You will perhaps like to hear how it acted on Brosera. 1
made a solution of ^ gr. to 5ij. of water. A minute drop on a
small pin's head acted powerfully on several glands, more power-
fully than the fresh poison from an adder's fang.
" I also immersed three leaves in 90 minims of the solution ;
the tentacles soon became inflated and the glands quite white,
as if they had been placed in boiling water. I felt sure that
the leaves were killed ; but after eight hours' immersion they
were placed in water, and after about 48 hours re-expanded,
showing that they were by no means killed. The most sur-
prising circumstance is that, after an immersion of 48 hours,
the protoplasm in the cells was in unusually active movement.
Now, can you inform me whether this poison, if diluted, arrests
the movement of vibratile cilia ? "
" I dissolved J gr. [of cobra-poison] in 5J of water, so that I
was able to immerse two leaves. It acted as before, but more
energetically ; and I observed more clearly this time that the
solution makes the secretion round the glands cloudy, which I
have never before observed. But here comes the remarkable
128 ON THE NATURE AND ACTION OF THE CROTALUS-POISON

point : after an immersion of 48 hours, the protoplasm within

the cells incessantly changes form, and I never saw it on any-
other occasion so active. Hence I cannot doubt that this poison
is a stimulant to the protoplasm ; and I shall be very curious
to find out in your papers whether you have tried its action on
the cilia and on the colourless corpuscles of the blood. If the
poison does arrest their movement, it will show that there is a
profound difference between the protoplasm of animals and of
this plant. Therefore if you try any further experiments I hope
that you will be so kind as to inform me of the results. I may
add that I tried at first 1 gr. to the 5J> as that is my standard
strength for all substances.
" It is certainly very remarkable that the poison should act
so differently on the cilia and on the protoplasm of Drosera.
After the 48 hours' immersion, T placed the two leaves in water
and they partially re-expanded. I thought that the whitened
glands were perhaps killed ; but those of one leaf which I tried
with carbonate of ammonia absorbed it, and the protoplasm was
affected in the usual manner. I am very much surprised at the
action of the poison on the viscid secretion from the glands,
which it coagulates into threads and bits of membrane, with
much granular matter. Have you observed whether the poison
affects in any marked manner mucus or other such secretions ? "

Experiment XXY.
Action of Cohra-poison on Muscle.
June 29th, 1874. — A standard solution of cobra-poison,
0'03 gramme to 4'6 c.c. of water, was prepared.
1.25 P.M. The gastrocnemius of a frog was separated and
immersed in this solution in a watch-glass ; it immediately con-
tracted considerably.
1.30. The muscle contracts with current at 11.
1.45. The muscle has lost its irritability ; does not respond to
the strongest current.
Experiment XXVI.
At the same time (1.25 p.m.) the gastrocnemius from the
other leg of the same frog immersed in water. Did not
 AS COMPARED WITH THAT OF OTHER VENOMOUS SNAKES. 129

immediately contract like that placed in the poisoned

solution.
1.30. Contracts strongly to current at 15 cm. of Du Bois
Eeymond's coil, more than the poisoned muscle at 11, at the
same moment.
1.45. Contracts distinctly at 11, whilst the poisoned muscle
has lost all irritability.
From this it is evident that the poison first stimulates the
muscular fibre to contract, but rapidly afterwards destroys its
irritability.
Experiment XXVII.
The gastrocnemii of a frog were again treated in the same
way as in the previous experiment, with precisely the same
results.
June 28th. — Made several experiments with cobra-poison on
ciliated epithelium of frog's mouth, and found that it at first
accelerated, then destroyed, the action of the cilia.

Experiment XXVIII.
To Test the Effects of Cohra-poison, when swallowed, on the Frog.
June 24th, 1874. — At 2.25 p.m. about |- of a grain of dried
cobra-poison was passed down a frog's throat.
2.30. Frog making violent efforts to vomit. Gaping. Head
thrown back tetanically.
2.34. Bloody mucus vomited with violent efforts.*
2.50. Moves with difficulty ; is becoming paralysed. Efforts
to vomit continue.
3. Much the same.
3.5. Very weak ; still tries to vomit.
3.10. Keflex action still well marked.
3.15. Motor nerves apparently quite paralysed.
3.20. Apparent death.

Artificial Respiration with Pure Oxygen.

As life had been prolonged for many hours in snake-poisoning
by artificial respiration with atmospheric air, it was thought
* This experiment is especially interesting, as showing that frogs do occa-
sionally vomit, a fact which has been denied by some physiologists.
(95) K
130 ON THE NATURE AND ACTION OF THE CROTALUS- POISON

expedient to ascertain if the more complete oxygenation by the

undihited gas would be more efficacious, as it seemed might be
possible ; accordingly the following experiment was made on
the 24th April, 1874.

ICxperiment XXIX.
J of a grain of dried cobra-poison dissolved in distilled water
was injected into a rabbit with the hypodermic syringe.
Symptoms of poisoning were rapidly manifested. A tube
had been previously introduced into the trachea, and respiration
was commenced as soon as poisoning was manifest.
Artificial respiration, with oxygen contained in a large bag,
was steadily continued for 2 hours, but with no better effect
than in other similar cases where atmospheric air was used for
the same purpose. At the expiration of 2 hours, apparent
death had occurred ; the heart continued to beat for about
2 minutes after the respiration ceased.
Beyond a very florid condition of the blood, there was no
obvious difference between the effect of oxygen and that of
common air. It did not indeed appear that, as far as the effects
produced by the poison were concerned, it differed in its action
from common air.

Experiment XXX.
November, 1874. — A little cobra-poison, dissolved in water,
was added to water containing some cells scraped from the
mantle of a freshwater mussel. Among these was a large
ciliated cell, which, before the addition of the poison, had been
moving slowly, although its cilia were moving actively. Imme-
diately after the addition of the poison the cell began to spin
round on its own axis with extraordinary rapidity. In about
three or four minutes its motions began to be languid, the
ciliary motion ceased, the cell itself elongated, contracted, and
then slowly resumed its former shape and became perfectly
motionless.
Experiment XXXI.
Water from the interior of a freshwater mussel, and
 AS COMPAKED WITH THAT OF OTHER VENOMOUS SNAKES. 131

containing two specimens of Paramcecmm in active motion, was

examined. They were rotating with great rapidity. A lit tile
cobra-poison dihited with water was added. Three minutes
after the addition one was discovered with both the ciha and
cell-body perfectly still. The cilia of the other were still, but
the cell-body was contracted. In about half a minute more it
expanded to its normal size and then remained perfectly still.

Experiment XXXII.
A piece taken from the mantle of a freshwater mussel was
placed on the slide and examined at the end of about half an
hour. Active ciliary motion could be observed in the fringe of
the mantle itself and in several specimens of Paramcecmm. A
little dilute poison was added. At first the ciliary motion
seemed increased, but in about 2 minutes it became slower, and
in 6 had become very languid, and in 10 minutes stopped
altogether in the specimens of Paramcecmm, but still continued
in some of the cilia of the mantle.

Experiment XXXIII.
A little dilute cobra-poison was added to a piece of the mantle
of a freshwater mussel. The cilia began immediately to move
much more rapidly. This was watched for some time. Ciliary
motion not affected, or at all events not arrested, after more
than half an hour.

Experiment XXXIV.
December 10th, 1874. — A piece of the gills of a freshwater
mussel placed under the microscope and a little cobra-poison
added at 10.40 p.m. The cilia vv^ere extremely active.
At 10.55 still active.
11.5. Several ciliated amoeboid masses are now quiet instead
of rolling over and over as they did, but the cilia on their
surface are still moving.
11.15. The cilia on these Infusoria have now nearly all
stopped. A few are moving slowly, whilst those on the gills
are but little affected.
(95) K 2
132 ON THE NATURE AND ACTION OF THE CROTALUS-POISON

11.55. Cilia on the gills are still quite active. Those on the
ciliated bodies still moving, rather more actively than before.
1.30. Cilia on gills have become much more sharply outlined.
Many are standing still, though many still move briskly.

Experiment XXXV.
To another specimen a strong solution of cobra-poison was
added at 10.50.
1.30. Cilia still moving.

Experiment XXXVI.
A third specimen was laid in an almost syrupy solution of
dried cobra-poison at 11.28.
At 11.40 no effect observable.
1.30. Some have stopped, but numbers are still moving quite
briskly.
In this case the poison seemed not to have any action on the
ciliary motion.
Experiment XXXVII.
January 6th, 1875. — At 3.40 some diluted cobra-poison added
to Vallisneria. Circulation going on vigorously. About -^ grain
in 3 drops of water.
3.58. The movements are unchanged.
5 P.M. Movements going on as before.

Experiment XXXVIII.
Added some solution of cobra-poison at 4 P.M. to another
specimen of Vallisneria.
4.10. No change.
4.45. Circulation goes on vigorously.
4.55. Perhaps rather less brisk in their movements.
The results of these experiments show that cobra- virus must
be regarded as, to a certain extent, a poison to protoplasm,
seeing that it arrested with rapidity the movements in
Infusoria* {vide Experiments XXX, XXXI, and following).
* Is this accounted for by the existence of a rudimentary nervous system
diffused throughout these two forms of life, and on which the poison could act ?
 AS COMPARED WITH THAT OF OTHER VENOMOUS SNAKES. 133

Still it cannot be regarded certainly as a very powerful one, for

the cilia of the freshwater mussel continued to move for many
hours in a strong solution of cobra-poison ; though in other
experiments the action was apparently arrested even in weaker
solutions of the poison. In the case of cilia from the frog's
mouth, the results were more definite, but action was not
invariably destroyed. The results of the action of the poison
on the amoeboid movements of the blood-corpuscles are not
very definite. In the case of Vallisneria, the circulation in the
cells went on with undiminished vigour after the application of
the poison for 2 hours.
NOTE ON INDEPENDENT PULSATION OF
THE PULMONARY VEINS AND VENA
CAVA.

By T. Lauder Brunton, M.D., F.R.S., and Sir J. Fayrer,

M.D, K.C.S.r.

(Eeprinted from the Proceedings of the Eoyal Society, No. 172, 1876.)
Keceived June 15th, 1876.

In a former communication* we incidentally mentioned that in

a rabbit killed by the injection of cobra-poison into the jugular
vein we had observed the pulmonary vein pulsating after all
motion had ceased in the cavities of the heart. We have since
observed the same phenomenon three or four times under con-
ditions which show that this pulsation is not due to the action
of the cobra-poison with which the animal in which we first
observed it had been killed. The following example will show
the changes in rhythm observed in these pulsations.
A cat was chloroformed, and the vagi exposed and irritated
by an interrupted current. Artificial respiration was kept up
by air containing chloroform vapour, and the thorax was then
opened, and a solution of atropia injected directly into the heart
by means of a Wood's syringe. The vagi were again irritated,
but without any effect being produced on the heart, the inhibi-
tory apparatus in it being evidently paralysed by the atropia.
A solution of glycerine extract of physostigma was now injected
into the heart in a similar way. The vagi were now irritated
again, and the heart stood still, the effect of the atropia having
been counteracted by the physostigma. After the irritation
ceased the heart again commenced to pulsate.
Artificial respiration was now discontinued, but all the cavi-
ties of the heart continued to beat for a considerable time. The
* Proceedings of the Royal Society, 1874, vol. xxii, j^. 125.
 INDEPENDENT PULSATION OF PULMONARY VEINS, ETC. 135

ventricles then stopped, but the auricles continued to beat. It

was then noticed that the pulmonary veins in the right lung,
which was exposed to view, were pulsating. The veins, as well
as both auricles, pulsated at the rate of 119 per minute, but the
contractions of the veins were not synchronous with those of
the auricles. Both auricles next ceased to beat, but the pul-
mtmary veins in both lungs continued to pulsate. The ventricles
now began to beat again, while the auricles remained still. The
ventricles pulsated at the rate of 8 per minute, while the pul-
monary veins pulsated at the rate of 46 per minute ; and no
motion was perceptible in any part of the auricles.
One hour and twenty minutes after the thorax had been
opened, and about an hour and ten minutes after artificial
respiration had been discontinued, the ventricle was still pul-
sating. Its rhythm was very irregular. After one beat a pause
of half a minute followed, and then 37 pulsations all together.
One hour and forty minutes after opening the thorax the inferior
vena cava was noticed to be pulsating close to its entrance into
the auricle. A contraction spread like a wave from the vena
cava over the right auricle, and the appendix contracted after
the auricle itself. The superior vena cava also pulsated close to
the heart. The left auricle had ceased to pulsate a considerable
time previously, and the ventricles had also stopped. After the
auricles had pulsated for a while the ventricles again began.
At one hour and fifty minutes after opening the thorax the
inferior vena cava was still pulsating. In ten minutes more all
movement had nearly ceased, and the observation was discon-
tinued.
At one hour and fifty minutes after opening the thorax slight
contractions of the diaphragm were noticed.
The striking points in this experiment are the contractions of
the pulmonary veins and the vena cava independently of the
heart, the long time during which they retained their irrita-
bility, and the continuance of their pulsations after the other
parts of the heart had ceased. The pulsation of the pulmonary
veins and of the ventricles at the same time, while the auricles
were motionless, is also deserving of attention.
In another experiment we found the pulmonary veins
136 INDEPENDENT PULSATION OF PULMONARY VEINS, ETC.

pulsating in a cat killed by a blow on the head. We have

also seen pulsation in animals killed in other ways ; but the
proportion of cases in which we have seen it to those in which
we have not seen it is very small. On looking through several
modern text-books of physiology, we have failed to find any
mention of the rhythmical contractile power of the pulmonary
veins and vena cava ; but the earlier anatomists were well
acquainted with it, and Haller* states that he has seen the
pulmonary veins continue to pulsate for two hours, and that
others had seen the vena cava pulsate for three hours while all
motion in the other cavities of the heart had already ceased.
Johannes Miillerf has also observed contractions of the vena
cava and pulmonary veins ; and in young animals the contrac-
tions of the pulmonary veins extend as far as they can be
followed into the lungs.
The importance of contraction of the vena cava and pul-
monary veins in preventing reflux of blood into them during the
contraction of the auricle, under circumstances when any
hindrance is opposed to the free flow of its contents into the
ventricle, is self-evident. Indeed, Hallerj: says that it was sup-
posed to exist by Senac, although he had not seen it. Especially
in cases of valvular disease of the heart is it likely to be of
great service ; and we think it advisable to bring again before
the notice of physiologists and physicians this power of the
veins, which, although so long known, appears of late years to
have been overlooked.
* Elementa Physiologia, 1757, torn, i, pp. 410 and 399; and Memoires sur la
Nature sensible et irritable des parties du corpes animal, 1756, torn, iy, p. 4.
t Miiller's Physiology, translated by Baly, 2nd edit., vol. i, p. 182.
X Op. cit., p. 410.
NOTE ON THE EFFECT OF VARIOUS SUB-
STANCES IN DESTROYING THE ACTIVITY
OF COBRA-POISON.
By T. Lauder Beunton, M.D., P.E.S., and Sir Joseph Fayree,
KC.S.I., M.D., F.E.S.
(Eeprinted from the Proceedings of the Royal Society, vol. xxvii, p. 465, 1878.)
Received June 20tli, 1878.

In a paper read some time ago before this Society by Mr. Pedler,
he mentioned his discovery of the fact that the activity of cobra-
poison was completely destroyed by admixture with perchloride
of platinum. This substance, however, could only be regarded
as a chemical and not as a physiological antidote to the poison,
inasmuch as it had no power to modify or prevent the action of
the venom after its absorption into the blood. Mr. Pedler
expressed his opinion that the proper method of pursuing the
investigation was to investigate separately the action of platinum
salts and of cobra-poison upon the animal body. In the discus-
sion which followed we stated that the method proposed by
Mr. Pedler was, in the present instance, not likely to lead to
any results, and that, as the action of the substance employed
by him was in all probability due to its simply forming an
insoluble compound with the cobra-poison and not to any action
of the platinum 'per se, certain other metallic salts would have
a similar action to the perchloride of platinum. Experiments
have confirmed the opinion we then expressed,* and we find
the action of chloride of gold is precisely similar to that of
perchloride of platinum, the cobra-venom being rendered
entirely inert by admixture with the gold salt before its
injection into the body. Chloride of gold, however, like per-
chloride of platinum, is merely a chemical antidote, and does
not modify the action of the venom after its absorption into

* A. W. Blyth, M.R.C.S., "The Poison of the Cobra," The Analyst, Feb-

ruary 28th, 1877, p. 204.
138 ON THE EFFECT OF VARIOUS SUBSTANCES IN

the circulation. Permanganate of potash, which has been

recommended as an antidote, also destroys its activity com-
pletely. Chloride of zinc, chloride of mercury, nitrate of silver,
and carbolic acid all diminish the activity of the poison, and
prolong life when mixed with it before its injection ; but they
do not prevent death, nor do they prolong life to any great
extent. Perchloride of iron has very much less action upon the
poison than one would expect, and it prolongs life to a very
slight extent. Liquor potassai impairs the activity of the poison
very considerably, and prolongs life for several hours. When
a large dose of cobra-poison is injected, none of these substances
prevent death, even when applied immediately to the wound.
The reason of this probably is that they do not come into such
perfect contact with the poison as to destroy the whole of it,
and the portion which escapes destruction is sufficient to kill.
It is possible, however, that when minimum doses only are
injected, the local application of one or other substance may
turn the balance between life and death, but this point we
must reserve for a future paper.
Our first experiment was made in order to compare the
action of chloride of platinum alone with that of cobra-poison
alone, and of chloride of platinum injected after cobra-poison.

Experiment I.
February 25th, 1878. — A cat weighing 4 lbs. had about
1 c.c. of the chloride of platinum solution of the British
Pharmacopoeia injected into its flank.
3.44 P.M. Injection completed.
3.55 „ No apparent effect. The cat well and playful.
No symptoms whatever were observed, but after some days a
slough formed at the point of injection. Chloride of platinum
thus appears to have no physiological action whatever when
injected subcutaneously, beyond its effect as a local irritant.
In Experiments II and III similar* doses of cobra-poison
were subcutaneously injected into two cats; but in Experi-
ment III the injection of the poison was followed immediately
by the injection of a solution of chloride of platinum, into the
same spot, so as, if possible, to destroy the venom which had
 DESTROYING THE ACTIVITY OF COBRA-POISON. 139

not yet been absorbed. In this case death was delayed, but
not to a very great extent, as it occurred in an hour and fifty
minutes after the injection of the venom and chloride of
platinum, and in an hour and two minutes in the animal which
received the poison alone.

Experiment II.
Black cat, weight 5 lbs.
25 milligrammes of cobra-poison dissolved in 1 c.c. of
distilled water injected into skin of flank at 3.26 p.m. of
February 25th.
3.28 P.M. Vomiting. It had taken chloroform to keep it quiet
whilst being weighed, and was recovering from the chloroform.
Micturated. Drooping head on one side.
3.32 P.M. Vomiting again. Looks much depressed. Defecated.
3.40 „ Breathincr slow. Shalloiv.
3.41 Vomiting again.
3.45 Twitching of muscles.
3.52 The same state.
3.58 Defecating. Micturating.
3.59 Eetching. Vomiting. Moves about in a restless
manner
4.6 Moving hackwards with staggering gait.
4.12 Staggers, and head droops.
4.13 Falls over on its side. The respiration is slow.
Keflex from eye and ear almost gone.
4.14 P.M. Eeflex from head and legs when irritated. None
from tail.
4.15 P.M. Attempts to rise.
4.18 „ Got up, but fell over again. Tried to walk. No
reflex from the head.
4.20 P.M. Head raised and fell over again. Touching the eye
seems to rouse the cat, but no reflex of lids. Tries to get up,
but cannot. Fell over on the opposite side.
4.24 P.M. Again a struggle to rise.
4.27 „ Touching the eye produces no reflex. Breathing
very slow. Convulsive twitching of limbs.
140 ON THE EFFECT OF VARIOUS SUBSTANCES IN

4.28 P.M. Apparently dead. Heart still beating one hour and
two minutes alter injection of poison.
The blood, after death, formed affirm coagulum.
Intestines much congested. Patch of congestion in stomach.
Bed serum effused into the peritoneal cavity.
No local symptoms or changes.
A good deal of food in the stomach, notwithstanding the
vomiting. Digestion was in full action.
In these and other experiments the dose of cobra-poison was
regulated according to the weight of the animal, the same
proportion per pound weight being given in each case.

Experiment III.

A grey cat, 4 lbs. weight, had 20 milligrammes of cobra-poison,

dissolved in 1 c.c. of distilled water, injected under skin of flank
at 3.39 P.M. of February 25th.
At 3.40 P.M. a solution of chloride of platinum injected at the
same spot.
3.42 P.M. Very restless.
3.43 „ Drinks water.
3.52 „ Vomiting.
3.55 „ Dull and depressed.
4.10 „ Same condition.
4.20 „ Sluggish.
430 „ Eestless. Moving about.
4.45 „ Staggers. Getting very sluggish. Walks with
difficulty. Head drooping.
4.54 P.M. Shivering. Head fallen over.
5 P.M. Fallen over. Slow paralysis creeping over limbs.
Eespiration slow. Gets up, rolls over again. On touching the
eye the eyelid moves. Keflex not gone from the ear.
5.6 P.M. Fallen over. Paralysed. Keflex nearly gone, still
slight from ear. Pupils dilated.
5.9 P.M. Convulsions. Pupils become normal again. Eespira-
tion very slow — 13 per minute.
5.20 P.M. Tries to rise. Very feebly.
5.22 „ Fallen quite over.
 DESTROYING THE ACTIVITY OF COBRA-POISON. 141

5.23 P.M. In same condition. Makes feeble efforts to rise.

Pupils dilated again.
5.27 P.M. Again tries to rise. Micturition.
5.29 „ Convulsions.
5.30 „ Dead.

I
No local symptoms, i.e., no extravasation about the puncture.
No congestion of stomach or bowels. Stomach empty. Blood
coagulated after death.
Injected at 3.39 p.m.
Died at 5.30 p.m.
Death in one hour and fifty-one minutes.
The following experiments show the effect of chloride of gold
in completely destroying the cobra-poison.

Experiment IV.
March 7th, 1878. — 3 milligrammes of cobra-poison, mixed
with 1 grain of chloride of gold, dissolved in 40 grain measures
of water, injected into the hip of a white guinea-pig, weighing
18 oz., at 3.50 p.m.
4.10 P.M. Crouching quietly in corner of box. Tremor,
perhaps fright.
4.15 P.M. Seems uneasy; crouching in corner. No other
change. Recovered without any bad symptoms.

Experiment V.
March 14th. — In this experiment a very large dose of poison
was used.
30 milligrammes of cobra-poison, mixed with 1^ c.c. of a
10-per-cent. solution of chloride of gold, were injected into a
guinea-pig weighing 20 oz., at 3.30 p.m.
75 c.c. of water was used to wash out glass, and then injected.
The poison and the chloride form a yellow creamy precipitate.
3.30 P.M. Began to jerk and twitch immediately, excited,
running about the box.
3.35 P.M. Crouching in corner, twitching, but not otherwise
affected.
3.42 P.M. Not apparently affected.
142 ON THE EFFECT OF VARIOUS SUBSTANCES IN

3.52 P.M. Crouching ; does not appear affected, but is weak in

the hind legs when he runs.
4.10 P.M. Very little affected ; hind legs weaker, but he is
very active otherwise.
4.20 P.M. Much the same ; active, except that hind legs seem
rather weak.
4.55 P.M. Eemains the same. Eecovered perfectly without
any further symptoms.
In order to make sure that the dose of cobra-poison would
certainly prove fatal if administered alone, the animal, after its
recovery, was injected with a quantity of pure cobra-poison,
fifteen times less than that from which it had recovered, and,
as will be seen from Experiment VI, death rapidly occurred.

Experiment VI.
March 14th. — White guinea-pig that recovered from 30 milli-
grammes ofcobra-poison, mixed with chloride of gold.
At 4.45 P.M., 2 milligrammes of cobra-poison were injected
into the hip.
4.50 P.M. Very restless ; scratching his skin.
4.52 „ Twitching; very restless.
4.59 „ Squeaking ; very restless.
5.10 „ Injected leg weak ; not so restless.
5.15 „ Trying to vomit; twitching movement of head,
jerking upwards; violent efforts to vomit; a sort of cough;
flows from nostrils and mouth ; getting gradually paralysed, he
still crawls ; nearly violent efforts to vomit.
5.24 P.M. The animal creeps along, putting his head along the
ground.
5.26 P.M. Apparently dead ; heart still beats.
At 4.45 P.M. the injection was made, and at 5.26 p.m. the
animal was dead. Death in 41 minutes.
Experiment VII shows that chloride of gold is a chemical,
and not a physiological antidote, and does not prevent the
action of the poison after its absorption.
 DESTROYING THE ACTIVITY OF COBRA-POISON. 143

Experiment VII.
March 14th. — Guinea-pig, weight 16 oz.
5 miUigrammes of cobra-poison dissolved in 1 c.c. of water and
injected into the right hip at 3.39 p.m.
In three minutes afterwards 1^ c.c. of a 10-per-cent. sokition
of chloride of gold were injected into another spot (the left hip,
at 3.42).
3.43 P.M. Very restless.
3.45 „ Very restless ; head twitching ; drops the left leg.
3.53 „ Kestless.
3.55 „ Weak, dropping both hind legs ; left appears quite
paralysed.
4.5 P.M. Getting weaker ; paralysis creeping over him.
4.10 ,, Barely moves ; hind quarters completely paralysed.
4.12 „ Convulsions.
4.19 „ Heart still beats feebly.
4.20 „ Dead.
Experiment VIII shows that permanganate of potash destroys
the action of the venom.

Experiment VIII.
5 milligrammes of poison were dissolved in 1 c.c. of water, and
mixed with 1 c.c. of liquor potassae permanganatis of the
British Pharmacopoeia, and injected under the skin of a guinea-
pig. No symptoms were produced, and the animal remained
quite unaffected.
Experiment IX.
Two rabbits of the same litter, each weighing exactly 2 lbs.,
were taken. 5 centigrammes of cobra-poison dissolved in 1 c.c.
of distilled water were mixed with 1 c.c. of liquor potassae
permanganatis (B.P.), and allowed to stand for about 8 minutes.
The mixture was then injected under the skin of the flank of
one rabbit. No symptoms whatever were produced, and the
animal, though kept under observation for some weeks, remained
quite unaffected by the poison. 5 milligrammes of cobra-poison
dissolved in 2 c.c. of water were injected into the other rabbit
at the same time. During the injection a little of the poison
144 ON THE EFFECT OF VARIOUS SUBSTANCES IN

was lost, SO that the animal did not receive the full dose, yet it
died in 30 minutes.
Chloride of zinc delays the action of the cobra poison, but
does not prevent it, as appears from Experiments X and XI, in
which a guinea-pig that had received 3 milligrammes of pure
cobra-poison (Experiment X) died in 45 minutes, whereas one
that had received a similar dose, previously mixed with chloride
of zinc, lived for about 3 hours (Experiment XI).

Experiment X.
March 1st, 1878.— 3 milligrammes of cobra-poison, dissolved
in 2 c.c. of distilled water, injected at 3.43 p.m. into a guinea-
pig's hip. Weight of guinea-pig, 20 oz.
3.46 P.M. Twitching.
3.50 „ Eestless twitching.
3.53 „ The same. Irritable; squeals; quarrels with the
other guinea-pigs ; respiration jerky.
4 P.M. The same,
4.5 P.M. Drags the injected leg, which is nearly paralysed.
4.15 P.M. Much the same.
4.19 „ Paralysed, and crawls with difficulty; all hind
quarters invaded by poison's influence.
4.21 P.M. Paralysis extending; struggles to rise; can only
move the head. Eeflex from eye diminished.
4.25 P.M. Convulsive movements.
4.28 „ Dead in 45 minutes. Heart continued to beat after
apparent death.
Experiment XI.
Eed guinea-pig, weight 16 oz. At 3.46 p.m., March 1st,
3 milligrammes of cobra-poison, dissolved in 2 c.c. of distilled
water and mixed with 0*01067 chloride of zinc, were injected
subcutaneously. The poison and the chloride were mixed
5 minutes before injection.
5.53 P.M. Guinea-pig restless; twitching; grunting; keeps
licking the puncture ; irritable with other guinea-pigs.
4 P.M. Very restless. Puncture seems irritable ; leg partially
paralysed.
 DESTROYING THE ACTIVITY OF COBRA-POISON. 145

4.15 P.M. Much the same.

4.29 „ Much the same.
4.35 „ Not quite so restless.
4.45 „ Active ; runs about.
4.45 „ Eestless ; not worse.
5 „ Seems pretty well now.
Died about 7 o'clock.
Liquor potassse impairs the activity of the poison, but does
not destroy it, as will be seen from Experiment XIT, in which
the dose of the poison, which had usually proved fatal consider-
ably within an hour, did not cause death until 8 hours had
elapsed.
Experiment XII.
March 14th. — Guinea-pig, weight 16 oz.
5 milligrammes of cobra-poison, dissolved in 1 c.c. water
mixed with 1 c.c. of liquor potassse, injected into hip at 3.52 p.m.
3.53 P.M. Twitching.
4.0 „ Leg paralysed.
4.20 „ It seems much the same.
4.35 „ Appears much the same.
4.55 „ Appears much the same.
5.35 „ Much the same.
At 11.30 it was lying with left hind leg paralysed, could walk
when irritated, mouth opened, head twitching back frequently.
11.45 P.M. its respiration ceased, but when the skin of the
belly was pinched the animal took a breath and respiration
continued for about a minute afterwards. The heart continued
to beat until 11.50.
Liquor ferri perchloridi fortior (B.P.) has much less action
upon the cobra-poison than one would have expected, as will be
seen from Experiment XIII, in which death occurred in an
hour and a half.
Experiment XIII.
March 14th. — Guinea-pig, weight 16 oz.
5 milligrammes cobra-poison, dissolved in 1 c.c. water mixed
with 1 c.c. of liquor ferri perchloridi fortior (B.P.), injected into
the left hip of the guinea-pig at 4.4 p.m.
(95) L
146 ON THE EFFECT OF VARIOUS SUBSTANCES IN

4.20 P.M. Dropped the hind leg, but otherwise seems active
and well.
4.35 P.M. Very active, but leg drops.
5.10 „ Hind leg paralysed.
5.15 „ Tries to crawl, cannot, struggles, convulsed.
5.16 „ Jerking convulsions.
5.17 „ Almost dead.
5.18 „ Dead.
Carbolic acid likewise delays the action of the poison, but to
a very much smaller extent than liquor potassse, as is proved by
Experiment XIV.
Experiment XIV.
March 14th. — Carbolic acid, ^ c.c. mixed with 5 milligrammes
of cobra-poison in 1 c.c. of water, injected into hip of a guinea-
pig weighing 14 oz. at 4.55 p.m.
5.55 P.M. Much the same.
7.50 „ Very slight convulsions.
7.55 „ Much the same.
8.0 „ Dead.
Experiment XV.
March 7th. — 3 milligrammes cobra-poison mixed with 1 grain
of nitrate of silver dissolved in 40 grains of water, injected into
hip of black guinea-pig, weighing 14 oz., at 3 p.m.
4.10 P.M. Twitching ; crouching in corner, crying out slightly
as guinea-pigs are wont to do when restless.
4.15 P.M. Kestless, crouched in corner of box, twitching of
muscles, cries as before.
4.25 P.M. Eestless, crying fretfully.
Died in about IJ hours afterwards, about 2 hours after the
injection of the poison.

Experiment XVI.
March 7th. — 3 milligrammes of cobra-poison mixed with
10 -grain measures of a saturated solution of chloride of mercury
(corrosive sublimate), injected into hip of guinea-pig (black and
white), weighing 14 oz., at 3.56 p.m.
4.10 P.M. Twitching, uneasy, tremors.
 DESTROYING THE ACTIVITY OF COBRA-POlSON. ' 147

4.15 P.M. Quiet, crouching in the corner.

4.24 „ Eestless, but does not seem to twitch ; cries like
the other guinea-pigs occasionally.
About 6.0 the animal lay quiet, with occasional twitches, and
about 6.30 it died.
In order to ascertain which substance would be most likely
to save the life of the animal by local application to the point
of injection, either by destroying the poison itself or by pre-
venting its absorption by the tissues, we applied chloride of
gold, permanganate of potash, chloride of platinum, and carbolic
acid locally, the method adopted being to inject . the poison
imder the skin of the leg, immediately afterwards to apply a
ligature tightly above that point, and then to make an incision
and apply the substance in just the same manner as we would
have done if the animal had actually been bitten. From the
following experiments, however, it will be seen that the absorp-
tion of the poison is so rapid that all local applications Were
useless. It should be noted that the quantity of poison we
employed was large, and it still remains to be seen whether
these local applications may turn the balance between life and
death when the quantity of the poison would be just sufficient
to kill in case of no remedy being applied.

Experiment XVII.
Guinea-pig weighing 1 lb.
4.27J P.M. Injected 2 gr. 9 centigrammes of cobra-poison into
thigh, ligature applied immediately.
4.29^ P.M. Solution of chloride of platinum applied.
. 4.31 „ Twitching violently.
4.33 „ Twitching violently ; ligature remains on limb.
4.40 „ Not worse; the ligature is evidently delaying the
action of the poison.
4.47 P.M. Getting weaker.
4.50 „ Convulsed.
4.52 „ Dead.
Death delayed in this instance.

(95) L 2
148 ON DESTROYING THE ACTIVITY OF COBRA-POISON.

Experiment XVIII.
April 4th, 1878. — Guinea-pig weighing IJ lb.
4.11' 10" P.M. Injected 3 centigrammes of cobra-poison.
4.13 P.M. Chloride of gold solution, 1 in 10 ; ligature kept on
until chloride of gold was applied.
4.15 P.M. Twitching.
4.16 „ Leg paralysed.
4.17 „ Animal nearly paralysed.
4.19 „ Animal dying.
4.20 „ Convulsions.
4.22 „ Dead.
Experiment XIX.
April 4th, 1878. — Guinea-pig weighing IJ lb. Injected
4 centigrammes of cobra-poison into leg.
4.1 P.M. Ligature applied immediately. Permanganate of
potash applied immediately.
4.5 P.M. Twitching.
4.10 „ Dying.
4.13 „ Convulsion.
4.14 „ Dead.
Experiment XX.
April 4th, 1878. — Guinea-pig weighing 1 lb.
3.45' 20'' P.M. Injected f gr. = 4 centigrammes of cobra-
poison under skin of leg. A ligature was applied round the
leg in one minute, and in five minutes permanganate of potash
was rubbed into an incision made over the site of injection.
3.52 P.M. Ligature cut.
3.53 „ Twitching violently ; leg paralysed.
3.55 „
3.57 „ Dying.
3.58 „ Dead — less than 13 minutes.
EXPERIMENTS ON A METHOD OF PHE-
VENTING DEATH FROM SNAKE BITE,
CAPABLE OF COMMON AND EASY PRAC-
TICAL APPLICATION.
By Sir Lauder Brunton, M.D., F.E.S., Sir Joseph Fayrek,
Bart., K.C.S.L, F.R.S., and Leonard Eogers, M.D., B.S.,
etc., Indian Medical Service.

(Eeprinted from the Proceedings of the Royal Society, vol. Ixxiii, 1904.)

Although this paper is a joint one, the authors wish to mention

that each has had a different part in its production. The whole
research may be fairly regarded as the natural outcome of the
work begun in India nearly forty years ago by one of us (Fayrer),
and this is the only ground on which his name can be associated
with this paper. The instrument employed was designed by
another of us (Brunton), and the actual experimental work was
entirely carried out by a third (Eogers).
The first experiments on the use of permanganate of potash
as an antidote to snake-poison was made by one of us (Fayrer)
in 1869, both by the local application of a solution and by
injection into the veins,* on the ground of its being a chemical
antidote. The animals experimented upon were dogs, but the
permanganate of potash did not seem to have any power to
avert the lethal action of the poison. It was shown also by
Wynter Blythf that cobra-venom, when mixed m vitro with
permanganate of potash, becomes innocuous. His results were
confirmed by two of us, who showed that some other substances
had a similar power.J They tried, by the injection of strong
* J. Fayrer, M.D., London, J. and A. Churcliill, The Thanatophidia of India,
1872, p. 95.
t A. Wynter Blytli, M.E.C.S., " The Poison of the Cobra," The Analyst,
February 28th, 1877, p. 204.
X Brunton and Fayrer, *' Note on the Effect of Various Substances in Destroy-

1878,ing vol.
the Activity of Cohrdb-Voi^on" Froceedings of the Royal ISociety,dvaie20t'h,
27, p. 465.
 150 METHOD OF TREVENTING DEATH FROM SNAKE BITE,

solution of permanganate of potash, and also by its local appli-

cation to an incision made over the bite, to destroy the lethal
action of cobra-poison previously injected, but their experiments
were unsuccessful, the permanganate appearing to be unable to
overtake the poison which had got the start of it.
In 1881 Messrs. Couty and Lacerda* made a number of
experiments upon the effect of permanganate of potash on
serpents' venom, and Lacerda found that permanganate of
potash not only destroyed the lethal action of the venom when
mixed with it in vitro, but also preserved life when a 1-per-
cent, solution of permanganate was injected into the tissues
close to the place where the venom had been previously injected,
and also when both venom and antidote were injected directly
into the vein. At the time of presenting his note to the
Academy of Science, in Paris, M. Lacerda was apparently
unaware of the previous experiments by Blyth, Brunton, and
Fayrer. In a later publicationf he discusses their experiments,
but claims for himself to have scientifically demonstrated per-
manganate ofpotash to be a precious antidote to serpent venom,
and to have brought it into common use, and thinks, therefore,
that the priority belongs to him ; but he was apparently
unaware that instructions for its use with the ligature had
many years before been promulgated by Fayrer in India.
In the winter of 1881 a number of experiments were made
by Dr. Vincent Eichards, who found, like the previous experi-
menters, that cobra-poison was completely destroyed by per-
manganate of potash when mixed with it in vit7V, so that
death did not follow the injection of the mixture either
hypodermically ox into a vein. He found also that when cobra-
poison was injected into a dog, and the injection followed either
immediately or after an interval of four minutes by a hypo-
dermic injection into the same part of a solution of permanganate
of potash, no symptoms of cobra-poisoning resulted, but after
the development of symptoms of cobra-poisoning permanganate
of potash failed to have any effect, whether injected locally or
into a vein or both.

*Coutj QjudLskcerdoj, Comptes JRendus, \ol. xcil, 1^. 465.

t Lacerda, Comptes Rendus, vol. xciii, p. 466.
 CAPABLE OF COMMON AND EASY PRACTICAL APPLICATION. 151

These results obtained both by Lacerda* and Eichards seemed

to give good hope that permanganate of potash might be used
to lessen the appalling fatalities from snake bite in India, but
it is evident that the hypodermic injection of a solution can
never be widely employed, because the hypodermic syringe is
expensive, it is liable to get out of order just at the times that
it is wanted, and the solution may become dried or spilt, or
may not be available. It is evident that the first requisite for
any antidote to snake poisoning is that it shall be always at
hand ; second, that it shall be easily applied ; and thirdly, that
it shall be cheap.
About two years ago, one of us (Brunton) was asked, on
behalf of a young officer going out to India, to design an instru-
ment which might be used in case of snake bite. He did so
accordingly, and he has since had a similar one made for him
by Messrs. Arnold and Sons, which seems to combine the three
requisites just noted. It consists of a lancet-shaped blade
about half an inch long, long enough, in fact, to reach the
deepest point of a bite by the largest snake. He has had some
instruments made with a double edge, like an ordinary lancet,
and others with one edge sharp and the other edge blunt, so as
to press in the permanganate. The lancet is set in a wooden
handle about an inch and a half long, which is hollowed at the
other end so as to form a receptacle to hold the permanganate.
Two wooden caps are fitted over the ends of the instrument, one
to keep in the permanganate, and the other to protect the
lancet. Such an instrument, if turned out in large numbers,
could be sold at such a small price as to be within reach of even
the Indian labourer, and might be sold everywhere in the same
way as packets of quinine are at present.

FiO-. 1. — Lancet for use in snake bite, showing the steel blade, the cap which
covers it, the hollow wooden handle for holding crystals of permanganate of
potash, and the cover which retains them.

* Dr. J. B. de Lacerda, Rio de Janeiro, Lombaerts, etc., " O Veneno ophidico

e seus antidotos," 1881, p. 64.
152 METHOD OF PREVENTING DEATH FROM SNAKE BITE,

The plan now proposed is to make a free opening into the site
of the bite, and to rub in crystals of permanganate. For this
purpose the limb should be surrounded by a tight bandage above
the bite, the puncture of the tooth or teeth should be freely cut
into by the lance-shaped blade and the crystals of permanganate
introduced and rubbed round. A few drops of saliva may be
added.
To test the efficacy of the proposed plan, several lethal doses
of venom dissolved in a few drops of water, go as to resemble, as
far as possible, the natural poison, are to be injected into the
limb of an animal, a ligature placed round the limb above the
seat of injection, an incision made, and crystals of permanganate
placed in the wound, moistened and rubbed in.

Experimental Investigation, hy Leonard Rogers.

In order to test in as practical a manner as possible the value
of the suggestion of the two first-named authors of this com-
munication,'the following experiments were carried out at the
Physiological Laboratory of the London University by the
third-named author. In the first place it was necessary to
ascertain if crystals of permanganate destroy the activity of
other venoms besides that of the cobra, for we are not aware that
its action in this direction has been tested against any extensive
series of snake venoms. As the value of the suggested
treatment would evidently be greatly enhanced if the per-
manganate could be shown to act efficiently against every class
of snake venom, a series of experiments were carried out to test
this point. The venoms in solution were mixed with small
quantities of a 10-per-cent solution of pure crystalline per-
manganate ofpotash in 0*9 per cent. NaCl, and after given
times the mixtures were injected into pigeons, several times a
lethal dose of each venom being used, so that if recovery took
place it would be evident that the permanganate had destroyed
the activity of the poisons. The following table (p. lol)
summarises the results of these experiments.
It will be seen that the table includes venoms of each main
subdivision of snakes, namely the two true vipers, the Daboia
Bussellii of India and the Puff Adder of Africa, the Pit Viper,
 CAPABLE OF COMMON AND EASY PRACTICAL APPLICATION. 153

the Crotal'Hs horridus, the Colubrine snake tlie Bungarus

fasciatus, and one of the Hydrophid?e or Sea-snakes, namely, the
Bnhydrina hengalensis. In the case of each, ten or more lethal
doses were neutralised by very small quantities of permanganate
in solution, and in most of them 20 lethal doses were readily thus
rendered harmless. The only failure was in Experiment 7, in
which 32'2 milligrammes of Bungarus fasciatus venom was added
to 25 milligrammes of permanganate of potash in solution, and in
this case by far the greater part of the poison must have been
neutralised, for in previous experiments one-eighteenth part of
the venom per kilogramme, used in Experiment 7, killed a
pigeon in one hour. Further experiments showed that 25 milli-
grammes of the permanganate of potash did entirely neutralise
16*1 milligrammes of Bungarus fasciatus venom. It is evident,
then, that the salt will neutralise about its own weight of this
venom, but that its power in this direction has a definite limit,
as might have been expected. It is clear, then, that this agent
does act on every class of snake venom and renders them inert.
Owing to the limited time available and the small number of
animals for which a license had been obtained, the actual
experiments on the treatment after injection of the venoms
have been so far limited to those of the cobra as a typical
representative of tlie-Golubrine class, and of the Dahoia Russellii
as a common and deadly viper. Rabbits and cats were used in
the investigation, the latter on account of their mixed diet and
firmer tissues resembling more closely the human subject. The
venoms were dissolved in as small a quantity of sterile normal
saline solution (0*9 per cent. NaCl) as possible, so as to
resemble in concentration the natural venom. The portion
of the limb to be operated on was cleaned of hair by scissors
beforehand (as the human subject is free from this obstacle to
treatment). The strong solution of venom was then injected
into the subcutaneous tissues of the cleaned part of a hind limb
a little above the paw, as most snake bites in the human
subject occur on the distal parts of the extremities. After a
given measured time a ligature consisting of a piece of bandage
was tied loosely round the thigh and twisted up tightly by
means of a piece of stick or a pencil so as to temporarily stop
154 METHOD OF PREVENTING DEATH FROM SNAKE BITE,

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 CAPABLE OF COMMON AND EASY PEACTICAL APPLICATION. 155

the circulation through the distal part of the limb in Order to

check further absorption of the poison. An incision was then
made in the long axis of the limb over the seat of injection of
the poison, and the edges dissected up slightly on either side so
as to fully expose the affected tissues and to form a small
pocket, into which the crystals of permanganate were next
placed, and after moistening with a few drops of sterile normal
salt solution (water, or even saliva, would serve in an emer-
gency) they were well rubbed in until the exposed tissues
presented a uniformly blackened appearance. About 3 minutes
were usually occupied by the little operation, on the completion
of which the ligature was released and a dressing and bandage
applied to the wound. The animals were under chloroform
throughout the operation, including the injection of the venom.
The amount of permanganate held by the instrument made for
these experiments was J gramme, this quantity being used in
each of the experiments.
The results of the experiments so far performed may most
conveniently be summarised in the following table, by means of
which they may readily be studied. The actual doses of venoms
injected are given in Column 4, and the dose per kilogramme
weight in Column 5. The time which was allowed to elapse
after the injection of the poison before the application of the
ligature (Column 6) was usually J minute, which it was calcu-
lated would be sufficient to allow a handkerchief, or, in the case
of a native, a strip of a pugari or of the cotton garments
commonly worn by the poorer classes in the tropics, being tied
round the limb and twisted up to form an efficient ligature.
In a few of the later experiments this application of the liga-
ture was delayed for 5 and 10 minutes. In Column 8 the time
is shown which was taken over the operation from the applica-
tion to the release of the ligature, while the ultimate resulc is
shown in Column 9. In most of the control experiments a
ligature was applied round the thigh for about the same time as
in the operations, as it appeared possible that the ligature
might delay somewhat the absorption of the poison, although it
could scarcely affect the ultimate result of its action, owing to
the poison being an essentially cumulative one.
156 METHOD OF PREVENTING DEATH FROM SNAKE BITE,

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 CAPABLE OF COMMON AND EASY PRACTICAL APPLICATION, 157

The first six experiments of Table II were performed on

rabbits, with the result that only prolongation of life was
obtained. Thus, after a dose of 10 milligrammes per kilo-
gramme (Experiment 1), death took place only a little quicker
than after one-tenth of this dose in a control animal (Experi-
ment 5). Again, 5 milligrammes per kilogramme in a treated
animal caused death in 3-J hours (Experiment 2), but 0*5 milli-
p-ramme per kilogramme in a control killed in the same time
(Experiment 6). The rapidity of death in this last animal
shows that 0*5 milligramme per kilogramme is still much above
the minimal lethal dose of cobra- venom for rabbits, so that the
doses used in the treated cases were many times a lethal dose
(about live to fifty times), and were thus mostly proportionally
larger doses than a cobra could eject in the case of a man. The
tissues of a rabbit are also more delicate than those of a cat or
of a man, so that absorption of the poison may be unusually
rapid in rabbits, which are extremely susceptible to snake-
venoms.
Turning next to the results of the experiments on cats, much
more satisfactory results were obtained. Thus, the control
experiments showed that 1 milligramme per kilogramme pro-
duced death in 50 hours, this being the m-inimal lethal dose of
the cobra-venom used in these experiments for cats (Experi-
ment 15). A dose of 5 milligrammes per kilogramme caused
death in 28 hours, the time having probably been prolonged by
the application of a ligature after the injection (Experiment 12).
A dose of 10 milligrammes per kilogramme proved fatal in
3 hours, although a ligature had been applied as in the treated
cases (Experiment 8). On comparing the result of treated
cases with the above control we find only one death occurred in
six experiments. The one fatal result took place after a dose
of 5 milligrammes per kilogramme (Experiment 9), this having
been the first case treated, in which the permanganate was not
as thoroughly rubbed in, and the site of injection was not as
completely exposed as in later experiments, and in this case
death did not take place until over 30 hours. On the other
hand, in Experiment 7 recovery took place after 10 milli-
grammes per kilogramme (10 lethal doses), while in two other
158 ^METHOD OF PREVENTING DEATH FROM SNAKE BITE,

cases recovery took place after five lethal doses had been
injected, in one of which (Experiment 11) 5 minutes were
allowed to elapse before the treatment was carried out, while in
Experiment 13 recovery ensued from lethal doses treated
10 minutes after injection.
The above results are very encouraging, for it appears from
D. D. Cunningham's observations that the average amount of
venom ejected by a full- sized cobra is not more than 10 lethal
doses for a man, while other writers give much smaller amounts.
Further, in many cases the full dose will not actually be
injected into the human tissue for various reasons.
In Table III a similar series of experiments with Daboia
venom are summarised. Here, again, in the case of rabbits,
only very marked prolongation of life was obtained, although
the dose used in Experiment 17 was less than four lethal doses,
so that it is clear that in the case of rabbits the method was
not very successful.
On the other hand, the experiments with cats were as
successful as in those of the cobra series given above ; for
only one of the six cases treated with permanganate died, and
in this instance (Experiment 21) the very large dose of
50 milligrammes per kilogramme was injected, and the treat-
ment was delayed for 5 minutes. This dose is probably
relatively larger than could be injected by any known viper in
the case of a full-grown man. Further, in this case death did
not take place until upwards of 24 hours after the injection,
while in a control experiment with the same dose (Experi-
ment 22) a fatal result occurred in four hours. Further,
with the same large dose recovery took place when treatment
was carried out half a minute after injection. Again, 30 milli-
grammes per kilogramme (three lethal doses) killed a control
cat in 4 J hours, but in three cases treated, ^, 5, and 10 minutes
respectively after injection, all recovered, as did one after
10 milligrammes per kilogramme, although a control with this
last dose died in 30 — 40 hours. In all the experiments of both
series the recovered animals were alive and well five days and
upwards after the injection of the venoms, which is two days
longer than death has ever taken place in any of the control
animals.
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CAPABLE OF COMMON AND EASY PEACTICAL APPLICATION. 159
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160 METHOD OF PREVENTING DEATH FROM SNAKE BITE,

The above results are very encouraging, as the Viperine

poisons are much less powerful, weight for weight, than are
most of the Colubrines and Hydrophidse, so that the amount of
venom ejected by them can seldom, if ever, be more than two
or three times a lethal dose for man.
In the course of the experiments it was observed that, even
when the incision mas made only 30 seconds after the injection
of the poison into the subcutaneous tissues, a distinct blood-
stained effusion is found, which serves as a very useful guide to
the location and limits of the injected poison ; after five or ten
minutes the effusion is more extensive, and in these cases the
incisions were prolonged up the limb for about 2 inches in
order to try and destroy as much of the venom as possible
The fact that as favourable results have been obtained after
five minutes as after half a minute, may very possibly depend
on the effusion noted materially checking the absorption of the
poisons, so that at the end of that time the rate of absorption
may become very much less rapid than during the first few
seconds after its injection. That a very rapid absorption occurs
during the first few seconds after the injection (probably on
account of the action of the poison in preventing clotting of the
blood locally) is certain, for it was showm by Fayrer many years
ago that a dog bitten in the tail by a full-sized cobra died, in
spite of the tail being cut off between the bitten part and the
body a few seconds after the bite. In such cases, however, the
dose received is relatively much larger than could be injected
by a cobra in the case of such a large animal as man, so that in
practice (except in the very rare cases where the poison is
injected directly into a vein) a fatal dose may not enter the
system for some considerable time after the bite. This
probability is supported by the fact that, in the case of
Colubrine poisons at any rate, the minimal lethal dose is the
same whether the venom is given subcutaneously or intra-
venously, yetit takes one or two days to produce death when
injected under the skin, but only 5 — 20 minutes when inserted
into a vein, so that under the former conditions the whole of the
poison does not enter the circulation for a long period. These
facts suggest the hope that the method of treatment here
 CAPABLE OF COMMON AND EASY PRACTICAL APPLICATION. 161

advocated may produce good results, even when it is not put

into operation until considerably longer periods than in any of
the above experiments, especially when only slightly supra-
minimal lethal doses have been received into the tissues.

Conchtsions.
Further experiments will be necessary to ascertain the exact
limits of the value of this form of treatment, and they will be
undertaken immediately by one of us (Eogers) in India, fresh
venoms being tried, as it is possible that they may be more
rapidly absorbed than those which have been dried and
redissolved. We think, however, that the results reported in
this communication are sufficiently promising to make it
advisable to place them on record, with a view to a trial being
given to the method in suitable cases, especially as the crystals
of permanganate of potash are actively antiseptic without
acting as more than a superficial escharotic, so that the treat-
ment has no markedly injurious effect which can be weighed
for an instant against the terrible results of bites by venomous
snakes. The process here recommended has already yielded
experimental results far in advance of anything hitherto
attained.
It is w^orthy of note that the earlier experiments of the first
two authors were stopped nearly 30 years ago by the passing
of the Act for regulating experiments on animals in England,
but for which this logical sequence of their earlier work might
very probably have been made many years ago.

(95) M
 163

INDEX.

Abdominal Cavity, injection into, of Areolar Tissue, effect on, of

Cobra -poison, effects of, 43 Cobra-poison, 43, 123-6
— Vein (Frog), injection into, of Cobra- Crotalus-'^oison, 123-6
poison, effects of, 75 Dahcda-^oi^on, 24
Absorption, Mode of action of Snake- Injection into, of Liquor ammonise,
poison, 18, 105, 106, 160 for Snake- bite, 109
Extreme rapidity of, 19 Armstrong, Dr., on Chemical Characters
— of l*oison in general, 105 of Cobra- poison, 8, 9 & note
Means of arresting, 106 Artificial Respiration in Cobra-poisoning,
Albuminoid body, chief constituent of 83, 84, 85, 87
seq.
Cobra- poison, 9 effect on Cardiac action, 96 et
Alcoholic extracts of
Cobra-poison, action of, 9, 14, 37, effect on Convulsions, 90, 91
67-8, 91 Life prolonging action, 5, 17,
Eattlesnakg-poison, action of, 10 20-2, 107-8
— Precipitates of with Pure Oxygen, unsuccessful,
Cobra-poison, action of, 9, 16 129-30
Or ot a lus -poison, action of, 10 in Curara-poisoning, Life-saving
action of, 45
American Kattlesnake-poison, experiments
with, 114 et seq. in Poison cases (general), 17
Ammonia, useless as Antidote to Snake- Asphyxia, the usual cause of death from
89
Cobra-poison, ::0 Sf note, 55,
poison, 37, 38, 39, 109-10,
Fontana on, 4, 8
Ataxy from Crotalus-poison, 112
Amoeboid movements of Blood Corp ii«cles,
Auricles, as affected by Independent
effect on, of Cobra-poison, Pulsations of Pulmonary
127, 133 Veins
136 and Yena Cava, 135,
Analysis of Cobra-poison by Armstrong,
9
Animals and Birds experimented on with
Snake-poison, and Brunton's Beenaed,Claude, on Rapid Absorption
Permanganate Lancet, 153, of Poison by Blood, 19
157-60, Tables I., II., III., Bernard's method of Ligaturing, 59
154, 156, 159
Bicliat, on commencing point of Somatic
Animals, Poisoned, effect on, of Inspiration Death, 20
of Pure Oxygen, 111. 129- Birds^ .see under Names
30 Black Snake virus, a,ction of, 112
Antidotes to Snake-poisons (see especially Blake, on Rapidity of Absorption of Poison
Permanganate of Potash), by Blood, 19
cause of apparent success Blood, action on, of Cobra-poison, 23, 25,
43, 44, 90
of, 7-8
Chemical, limitation of, 137, 142 — as affected by Cro^«^M*-poisoning
Essentials in, 6 4* note, 19 (a) before Death, 113
Inefficiency of, 3, 4, 8, 17, 110 (b) after Death, 113
Physiological, 137 — as affected by Dahoia-ipoisoning, 18
Anti- vivisection Act, effect of, on efforts note, 25, 44, 84, 113 <f- note,
to provide Antidotes for S( .^ee Cases
Snake-bites, 161 — Changesof Colour in, in Dahoia--po\?>on-
Aortal injection of Cobra-poison, 75 -6 ing, 26, 27, 28
(95) M 2
KM INDEX.

Blood (cont.) Carbolic Acid, action of, on Cobra -poison,

138
— Coagulability of, offecfc on, of Snako- j
on
Guinea-pig, injected with
Colubrine,
poison 18 Cobra-poison, 146
Viperine, 18 ^ note Carbonic anhydride gas, given off by
— Coagulation of decomposing Cobra-poison, 8
in Cobra-poisoning, 25, 114, 117-8, Cardiac action. Arrest of
140, 141 from Cobra-poison, 11 note
in Cro^ato-poisoning, 113, 118 Snake- venom, 94, 104, 112
in DaJwta-poisoninu, 25, dif see Cases — Diseases, Valvular, importance in, of
'— Corpuscular changes, see Erythrocytes \ Independent Pulsation of
— Extravasation of, in Cobra-poisoning, i Pulmonary Veins and Vena
Cava, 136
10, 13, 37, 43, 113, 117-8, !
— Ganglia, action on (suspected)
123-6 of Cobra-poison, 96
in Croifa/w5-poisoning, 113, 117-8,
123-6 ^ of Snake-poison, 7, 112
in Daboia-Tpoisoning, 24 — Pulsations in Frogs, in Cobra- and
Curare-poisoning, 96
in Snake-bite, 113, value of, 160 '.
— Frogs', eflect on, of Cobra-i)oison, 127 | in application to the Heart of
Cobia-poison, 96, 9y, 100,
— Newts', effect (imperceptible) on, of ; Tracings, 101
Cobra-poison, 127
— of Eabbit poisoned by Crotalus-yirus, \ persistence of, after apparent
effect of, on Guinea-pig, j Death, in Cobra-poisoning,
121 I 12, 13, 14, 15, 16, 20, 21,
Blood-changes, due to Snake-poison, 6, 7, j 22, 30, 38, 39, 42, 69, 82, 95
18, 84, 90 I in Baboia-^oimnmg, 27, 28
Blood-poisoning from Cobra-bite, 43 1 Carp, effect on, of Curare injection, 36
Blood-pressure in Cobra-poisoning, 97, i Cats, action on, of
101-3, 104 j — — Chloride of Platinum, alone, 138
Blyth, Wynter, on Cobra-venom mixed m \ injected after Cobra-poison,
vitro with Permanganate of 138, 140
Potash, 149 i — Effects on, of Cobra-poison, 83, 84,
Brain, the, why necessary to Life, 20
Brunton, Sir Lauder on Circulation, 100
86-8
— Lancet designed by, for use in Snake- after injection to Mesentery,
bite, Fig. 1., 151, how em- 123, 125
on Nerves, 70, 86-8, 94-5
et seq. ployed, 152, how tested, 152 on Eespiration, 91
JSungarus cceruleus {see also Krait), 23 Vomiting, 84, 89, 139, 140
— — Bite of, eflect of, on Innocuous of Crotalus-ipoison, 121, 123-5
Snakes, 31, 33, on other — Experiments on, with Permanganate of
Venomous Snakes, 34 Potash and Ligature, in
Destructiveness of, 3 Snake-poison, 153-8, Tables
Effect on, of Cobra bite, 33 IL, 111., 154, 159
Sungarus fasciatus, 23 — Independent Pulsation in, of Pulmon-
Destructiveness of, 3 ary Veins and Vena Cava,
Effect on, of Cobra-bite, 33, 34
Venom, neutralised by Permanga- — Length of 134,158
Life 135-6
after Cobra-poisoning,
nate of Potash, 153
Busk, Prof., on Venom and its principle, Caustic Potash as Anti-venom, 6 note
9 note Cerebellum, effect on, of Cobra-poisoning,
85
Cerebro-spinal Nerve centres. Snake-
Calcutta, G-overnment Commission at, on poisons acting on, 112
Snake-poisons, 112, conclu- Cerebrum, apparently unaffected by
sions of, 113-14 Cobra-poison, 57
Callophis, 23 Chemical Analysis of Cobra-poison,
Armstrong, 9 by
Canino, Prince of, on Echidnine, 9 note
Capillary Circulation, effect on, of Cobra - Chloride of Gold, action of, on Cobra-
poisoning, 14, 97
poison, 137, 143-3
 INDEX. 165
Chloride of Gold, action of (cont.) Cobra-poison, Action and effects of {cont.)
on Gruinea-pig, locally applied, Fresh, 16, 57, 59
147, 148 Mixed with Liquor Ammonise
injected, mixed with Cobra- and Liquor Potassae, ih.
— with Salt Solution, 54, 61
poison, 141-2
injected after Cobra-poison, 143 — Action and Effects of, according to
Mode of Entry
— of Mercury,138action of, on Cobra-poison,
Absorption, 18, 19, 42, 43, 46, 47,
106
on Guinea-pig, injected with Cobra-
poison, 146 — Application, 46, 124, 125, 127
— Platinum, action of, alone on Cat, 138 Bite, 30-1
3 etet seq.,
seq. 10, 18 et seq.,
on Cobra-poison in Guinea-pig,
locally applied, 147 Immersion, 49, 54, 1 28
Injected
140 after Cobra-poi«on, 138, Injection, 10-17, 21-2, 26,
' 28-9, 38-9, 41, 43, 47, 49,
— of Zinc, action of, on Cobra-poison, 50, 52, 53, 55, 59 et seq., 70,
138 I 71, 73 et seq., 117, 119
on Guinea-pigs, injected mixed Swallowing, 41, 42, 43
with Cobra-poison, 144-5 — Action of, on Amoeboid movements of
Ciliai'y action, effect on, of Cobra-poison, Leucocytes (Frog's Blood),
126, 130-3 127, 133
Ciliated Epithelium, effect on, of Cobra- analogy between, and that of
poison, 126, 133 Curara-poison, 4
Circulation, action on, of Cobra-poison, on Animals, Birds, &c.
41, 49, 75-6, 95 et seq. on Cats, 83, 84, 86-8, 89, 91,
Failure of, in relation to Somatic 94-5, 100, 123, 125, 139, 140,
Death, 20, and Death from 142, 144
Cobra- poison, ih. on Cold-blooded Animals, 24
— Capillary, effect on, of Cobra-poison, on Dogs, 10, 11, 69, 70, 89, 93
14, 97 on Fish, 35-6
Coagulability and Coagulation of Blood, on Fowls, 11, 21
see under Blood -on Frogs, 29-30, 41, 42, 46 et seq.,
37 Cobra-poison, effects of,
Coagulation of 54, 59 et seq., 71, 73 et seq.,
99, 100, 128
Cobra {Naja tripudians), the most veno- ^ on Guinea-pigs, 10, 13, 15, 16,
mous and destructive Indian 28-9, 38-9, 47, 50, 55, 82,
snake, 3, 23 89, 99, 117, 139, 142, 144
— Bite of, effect of injected with Metallic Salts,
on Innocuous Snakes, 31, 32, 33 138, 143, 144, 145, 146
on Nocuous Snakes 33-5 when Metallic Salts were in-
— Effect on, of Venom of itself, injected, jected after it, 140, 141,
35
143
of Venom of other Snakes, 34, 35 when Metallic Salts were
— Full-sized, average amount of Venom locally applied after it, 147,
ejected by, 158 148
Cobra-poison, Action and effects of on Rabbits, 10, 12, 14, 21-2, 92,
According to form in which used 97, 99, 103, 119
Alcoholic Extract, 9, 14, 37, 67-8, on Lizards, 30
91 on Pigeons, 26
Precipitate, 9, 16 on Rats, 44, 45
— Coagulated, by boiling, 37, 57, on Snails, 36
Sf see Cases on Snakes, 31 et seq.
Concentrated Solution, 81 — Action and effect of, on Blood (see also
Diluted, 11, 12, 15, 21, 22, 37, 49, 44
• Blood, Erythrocytes, Sfc),
52, 74, 75
Dried, 13, 26, 57, 117-8, Sf see («) in large quantities, 113
Cases, passim {h) in small, and in the lower
Energy of, greater than that animals, 114
of Crotalus, 117-18 on Ciliary action, 126, 133
with Liquor Ammonise, 38, 39 on Circulation, 95 et seq.
 16G INDEX.

Cobra-poisoning (coni.)
Cobra-poison, Action'and effects of (conf.) — Vomiting in, to what due, 89
compared with Crotalus poison, 43,
115-9, 123 Cold-blooded Animals, see under Names
with other Colubrine poison, 23 Colubrine Snakes
with that of Dahoia Ruftsellii, Bite of, effect of, on Blood, 18
24 et seq. Indian representatives of, 23
general, 10, 11, 24 Venom of, action of, similar to that
— ■— on Grt'rinination of Seeds, 40 of Crotaline virus. 111
on Invertebrata, 24, 36 minimum Lethal Dose, rate of
Local, 10, 43 Death from, 160
Means of Preventing, 17, 105 et seq., neutralised by Permanganate of
149 et seq.
Potash, 153-8, Table J.
on Motor Nerves, 47, 48, 58, 65, 69, (IV.), 154, Table IT., 156
70,71 Comb of Fowl, Colour of, as affected by
on Muscles, 46 et seq., 54, 128 Cobra-poisoning, 21, 90
on Nervous System, 55 et seq. Conia, action of, compared with that of
seq.
Physiological, see 1 et seq. ^ 23 et Cobra- poison, 57
Conjunctiva, action on, of Cobra-poison,
on Respiration, 89 et seq. 42, 43
Secondary, 54 Constitution, infl uence or, on action of
on Secreting Nerves, 69 Cobra-poison, 39
on Sensory Nerves, 60, 70, 71, Couty and Lacerda's Experiments on
86-8 Permanganate of Potash as
on Spinal Cord, 20, 56, 57, 65, 72
et seq. antidote
150 to Snake-poison,

on Stomach and Intestines, 69, 89, Convulsions at Death, see Cases, passim
93-4 — from Cobra-poison, 11, 24, 55
on various Organs of Body, 24, 41 causes, 20, 56, 90, 91
et seq.
— from Crofalus-'poison, 112
when Introduced through Different — from Daiom -poison, 25, 27, 28
Channels, 41 Cord, see Spinal Cord
— Action of, Influence on Craspedocephalus of America and West
of Chloride of Platinum and other Indies, 23
Mineral Salts, 140 et seq. Crenation of Red Blood-corpuscles
of Constitution on, 39 in Cobra-poisoning, 44-6, 127
of Reagents, &c., 36 in Croialus-^ohom.ng, 119, 122
of Various Substances, 137 —Crotalus-,
action ofor Rattlesnake-poison
— Activity of, effect of Various Substances
in Destroying, 137 on Animals, 112-13
— Appearance of, 8 on Cats, 121-2, 123-5
on Frogs, 122
— Chemical constitution of, Armstrong
on, 8, 9 on Guinea-pigs, 115, 116, 118,121
— on Rabbits, 119
— Cumulative nature of, 155
— Death-causing action of, 112 on Areolar Tissue, 123
— Decomposition on Ciliary and Amoeboid Action,
Cases caused by, 55, ^ see 126, 127, et seq.
— Decomposition of, 8, 10 Compared with Cobra-poison, 46,
— Elimination of, 5, 6, 17, 106-7, 108 115-8, 123
— Mode of Action of, 17 et seq., 55 et seq. on Infusoria, 130-3
— Nature of, 1, 23 et seq., Ill on Muscles, 46, 54
— Permanganate of Potash as Antidote to, on Seed-germination, 40
149 et seq. on Vallisneria, 132-3
— Used in Experiments of — Blood of Animals as affected by
Armstrong, 10 {a) before Death, 113
— — Brunton and Fayrer, 10 {h) after Death, 113
Cobra-poisoning — Effects of, according to Mode of Entry
— Artificial Respiration in, effect of, 5, Application, 123, 125
17, 20-2, 83, 84, 85, 87, 90, Injection, 115-122
91, 96, 107-8, 129 — Nature and IllPhysiological action of , . , .
— Symptoms of, 10, 11, 24, 55
 167

INDEX.

I)eath(s) {coni.)
€rotaliis-, or Kattlesiiake-poison (coni.)
— Neutralised by Periiianganiite of — from Cobra-poison, causes of, 4, 5, 7,
Potash, 153, Table I. (il.), 7, 20 cf note, 55, 57, 89
154
— — Convulsions at, 11, 20, 24, and see
Cases
— Used in Experiment«i, Age and Appear-
ance of, 114 — from Colubrine and Crotaline poison-
C>*o^«/«*-poisoning, Symptoms of, 112-3 ing, causes of, 111-2
Crural Vein, Injection into, of Cobra- — from Curara- poisoning, 4
poison, results, 5 — from Snake-bite, in India, 1, 2, 3
Cunningham, D. D., Experiments of, on — — Means of preventing, 105
Average amount of Venom — 18
from Snake-noison, how caused, 4, 5, 7,
ejected
158 by Full-sized Cobra,
— Somatic, causes of, 20
Ciu'are- or Curara-poisou, Absorption and Decomposition of Cobra-poison, changes
Excretion of, 105, 106 due to, 8, 10
— Analogy between its action and that of — due to Snake-poison, Cobra, and C'ro*
Cobra-poison, 4, 56, 69 talus
— Artificial Eespiration in, Life-saving, 4 of Mviscle, 54-5, cf •^^'' Cases
— Elimination of, state of Nervous of Tissues, incipient, 10
System after, 4, 5, 6, 58, 106 within Body, 55
— Warmth to combat, 4
Depression, after Cobra-poisoning, 10, 24,
55
— after Cro/c/Zw-y-poisoning, 112
Destructiveness, Relative, of Venomous
Dahoia Riissellii, see also Russell's Viper Snakes, Indian, &c., 3, 23
Bite of, effect of, on Innocuous
Snakes, 31, 33, on other Diaphragm, observations on, in Cobra-
Venomous Snakes, 34, 35 poisoning, 86, 91
Destructiveness of, 3, 23 Distal parts of Extremities, most liable to
— — Effect on, of Cobra-bite, 35 Snake-bite, 153
Venom, action of, compared Avitli Dogs, Cobra-poisoned, effects on, of Per-
that of Cobra-poison, 24 e^
seq. manganate of Potash, 149,
151, 158
Average amount ejected, 160 — Effects on, of Cobra-poisoning, 10, 11,
Blood-changes from, 18 note^ 25, 93-4, 101-3
44, 84, 113 cf' note Paralysis, progressive, 10, 11, 24,
Death-causing action of, 112 69
Effects of Salivation, 10, 24, G9-70
Dried, 25 '■ ■ Vomiting, 10, 11, 24, 68, 89
Injected, 25 ef seq. — Rapid Death
160 of, after Cobra-bite,
• Mixed with Alcohol, 27
on Cats, 153, 157, 158, Table Dose, Lethal, Minimal,
III., 159 for Cats, 157 of Cobra-poison
for Man, 158
on G-uinea-pigs and Pigeons, for Rabbits, 157
24 et seq.
on Rabbits, 153, 157, 158, — Average ejected by full-sized
Table III., 159 Cobra, 158
Length of Life after administra- of Colubrine -poisons, time taken
tion, with control by Per- to kill by, according to Mode
manganate and Ligature, 158 of Entry, 160
Neutralised by Permanganate of Dahoia and Viperine Venom
Potash, 152, 153, 158, Table for Cats, 158
I., (i.), 154, Table III., 156 for Man, 158, 160
for Rabbits, 158
— Symptoms of Poisoning with, 24
Darwin, Charles, the late, on action of Average, ejected, 160
Cobra-poison on Vegetable Dried Cobra-poison, Activity of, 36
Protoplasm, 127 Appearance of, 8, 9
— Experiments suggested by, 126 Keeping quality of, 8
Death(s) from Cobra-bite, rate of, accord- Drosera, Darwin's experiments on, with
ing to point of entry, 41, 160 Cobra-poison, M127-83
(95)
 1()S 4NDEX.

])rowsiness or lethargy Fluidity

(partial or com))lete) of Blood
— ill Cobra-poisoning, 10, 11, 21, 24, pos- after Death from
sible cause, 55, 57 Cobra-poison, 113, 120
— in CroialtiS'-poisoning, 112 Crofalus-j)oisou, 120, 122
Dahoia-Yio\aon, 113, 114
Fontana, on Caustic potash as Anti-
KccHTMOSis after Cobra-poisoning, 10, venom, 6 uof.e
119, 123-6 — on Inutility of all Methods of treating
Snake-bite, 4, 8
— after Cro/alus-ixnsoning, 113, 116, 119,
121, 123-6 Fontana and Legallois on persistence of
Echidnine, the Prince of Canino on, Bodily Life, 20
9 no/e Fore legs. Paralysis of, 85
Echis carinata, Dcstriictiveness of, 3 Fowls, eiject on, of Cobra-poison, 11, 21,.
E)ihydriiia bem/alensU, Venom of, Neu- 24,90
tralised byPermanganate of Fresh fluid Cobra-poison
Potasli. 153 appearance of, 8
Erythrocytes, Crenation of, after keeping quality of, 8
Cobra-poisoning, 44-6, 127 Freshwater Mussel, Cilia of, as affected by
■ Cro/'friM.v-poisoriing, 119, 122 Cobra-poison, 130-3
Evacuations, Involuntary, after Cobra- Frogs, action on, of Cobra-poison, 29-30,.
poisoning, 11, 13, 15, 16, 24, 41,42
on Blood, 127
38, 51, 89, 139, 140, Sf see
Cases ■ on Ciliated Epithelium from,
— after jDfflioia-poisoning, 27, 28 126, 133
Ewart, Dr., deductions from experiments on G-dstrocnemius muscle of,
of, on Pseudechis porpht/-
riacus and Hoplocephains 128-9 96, 99, 100, 101
on Heart,
eurtuft, 112 Injected into Dorsal Lymph-sac^
Excised Heart (of Frog), action on, 30, 50, 53, 59, 61, 62, 64,
of Cobra-poison, 96, 99, 100 65, 66, 67, 71, 73, 74, 76,
Excretion of Poisons, 105 77, 79
— of Snake-poison, 104-5, 106 et seq. — — — on Motor Nerves, 59 et seq., 70,
Means of quickening, 108
Excretion or Elimination of on Muscles, 46 ei seq., 54, 55,
— Cobra-poison, 5, 6, 17, Question of 71
State of Nervous System on Nerves,
128-9 52
after, 5, 6 — on Sensory Nerves, 59-60, 70,
— Curara-poison, State of Nervous Sys-
tem after, 4, 5 on Spinal Cord, 73 et seq.
Exhaustion, after Cobra-poisoning, 10, 24 71 Cro/(i;7«.?- poison, 122
— action on, of
Extravasation of Blood, see nnder Blood, — action on, of Curare-poisoning, 96
200

Faintness, after Cobra-poisoning, 10, 24, GrASTEOCNEMii of Frogs, as affected by

55
Cobra-poison, 128-9
Fayrer, Sir J., author of Thanatophidia General efTects of Cobra- and Crotalus-
of India, 1, 17, 31 et poisons, 112, Experiments
passim showing, 115-22
— Experiments by, in Treatment of Snake- G-ermination of Seeds, action on
bite, 109 of Cobra -poison, 40
— Investigations of, on Loss of Life by \ of Rattlesnake-poison, ih.
Snake-bite in India, 1-4, Gills of Mussel, effect on, of Cobra-poison,
18, 19 131
— Priority of use by, of Permanganate of Glands of Drosera, as affected by Cobra-
Potash in Snake-bite, 150
Femoral vein. Injection into, of Cobra- Gold, .see Chloride
poison,of Gold
127-8
poison, results, 18, 19 Grass-snake {Tropidonotus qtdncunciattis) ,
Fish, effects on, of Cobra-poison, 35-6, of effect on, of Bite of Cobra,
Curare, 36 32
 INDEX. 169

G-reen Whip-snake (Passerifa mycteri- Hind Legs and Hind-quarters, Paralysis

zans), effect on of, in iSnake-poisoning, 10,
'■ of Cobra-bite, 32 24, 27, 28, 29, 55, to what
of Dahoia-h\te, 33 duo, 57, 58
Guinea-pigs, action on, of Ho'ploce'phalun curiiia (Tiger Snake, Aus-
Carbolic acid, injected witli Cobra- tralia), Virus of, Death-
poison, 146 causing action of, 112
Chloride of Gold, HydropJnclce, Destructiveness of, 3
injected mixed with Cobra-poison, -- Venom of, effects of, 23, 112
141-2 • Neutralised bv Permanganate of
injected after Cobra-poison, 143
locally applied after injection of Syjynale, 23 Potash,'l53
Cobra-poison, 147, 149
Chloride of Mercury injected with
Cobra-poison, 146 Immunity to Snake-poison
Nitrate of Silver injected with of Mongoose, 39
of Pig, 40
Cobra-poison, 146
— • — Liquor Potassee Permanganatis in- of Venomous Snakes, 31, 34-5, 105
iected with Cobra-poison, India, Loss of Life in, from Snake-bite,
143, 145
1, 2, 3
Perchloride of Iron injected with — Snakes of. Poison of, 1, 23 et seq., 112
Cobra-poison, 145 Relative Destructiveness of, 3
Permanganate of Potash after Infiltration of Tissues, in Cobra-poisoning,
Cobra - poisoniusr, locally 10, 43
applied, 147, 148 Infusoria, action on, of Cobra-poison,
— effects on, of Cobra-poison, 13, 15, 16,
28-9, 38-9, 43, 55 Injections. See130-3
Intravenal, Subcutaneous,
on Circulation, 99 and into various organs, &c.,
tinder Names
on Muscles, 47-9, 50 et seq., 55
on Spinal Cord, 82, 85 — of Permanganate of Potash
producing Vomiting, 10, 24, 38, 84 Locality for, 150, 152, 153, 160
— effects on, of Time for, after bite, 150
Crotalus-^oisow, 115, 116, 118, 121 Intestines {See also Stomach and In-
of Dahoia-^oi^on, 24, 25, 27 testines), Blood-conditions
of injection of Strychnia and Woo- in, after Crotalus-^oi<iomng^
rara, 88 113
Intestinal injections, action of
of Cobra-vii'us, 117
Haller on Independent Pulsation of of Crotalus-^o\s,on, action of, 116
Pulmonary Veins and Vena Intravenal entry of Colubrine-poisons,
Cava, 136 extreme rapidity of Death
Halijs, 23 from, 160
Heart, Action of, how arrested, in Snake- j of Snake-poison, rapidity of effects,
poisoning, 7, 76, 95, 120 j 18, 19, 20
— Action on, of Cobra-poison, 7, 11, 12, — Injections of Cobra-poison, rapidity
13, 14, 15, 16, 20-2, 38, of Death from, 5, 41
effect of, on
39, 50, 76, 95, 104, 120
Cats, 83, 100
Arrest of. in Systole, 96, 120
Dogs, 70, 94
Tetanic Contractions of, 95, 104, Frogs, 75
112
— Action on, of jD«5oi« -poison, 27, 28 i Rabbits, 22, 99, 103, 119
— Frog's, III situ and excised, action on Intravenal injection of Crotalus--po\sox\,
of Cobra-poison, 30, 42, 96, 99, action of
100 on Cat, 121
of Curare-poison, 96 on Rabbit, 119
Ilering, observations of, on Rapid Absorp- Invertebrata, 24
eft'ect on, of Cobra-poison,
tion of Poison by Blood, 19
Hermann, on Poisons, 105 Iron, see Perchloride of Iron
Hilson, Dr., on partial Paralysis in Man, Irritant character of Snake-poison, 18
in Cobra-poisoning, 56
170 INDEX.

Jugular rein, injection into Man, Pamlysis56 in, of Peripheral Nerves,

of Cobra-poison, results, 5, 18,
19, 20 uofe, 41, 99, 100, 103, Mantle of Mussel, effect on, of Cobra-
119
^ of C>'0^a//fs-poison, 119, 121 Means of Preventing
poison, ]30-lEffects of Poison,
(a) generally, 17, 105
{b) of Cobra-poison, ih.y 106
Kidneys, Cobra-venom excreted by, 104 Medulla, effect on
Krait (.vee also Bu)i<jartis caruleiis), De- of Cobra-poison, consequences of.
structiveness of, 3 90, 91
of C/*o/fi5Ze<.9-poison, 120
of Snake-poisons, 112
Lacerda, on Permanganate of Potash as Mercury, see Chloride of
Antidote to Snake-poison, | Mesentery, the. Blood-conditions in, after
150 I Crotalus -■^oiiomng, 113
Lancet, for use in Snake-bite, designed by i — Exposed, or withdrawn
Brunton, Fig. 1., 151, how | of Cat, action on, of
used, 152, liow tested, 152 | Cobra-poison, 123, 124, 125
ef seq. Crotalus-^oi'so\\, 123, 124, 125
Legallois, f/ee Fontana and Legallois Methylconia, action of, compared with
Lethargy, after Cobra-poisoning, 10, 11, that of Cobra-venom, 67
21, 24, 55, 57 Mitchell, Dr. Weir, Crotalv.s-])oison sup-
— ■ after Crotalus-x>o\mm\\g, 112 plied by, 114
Leucocytes, ofMovements
Fi'og's Blood, — Observations of, on effects of Crotalus-
of, as Amoeboid
affected
by Cobra-poison, 127, 133 - — on Effects poison, 113
of Rattlesnake-poison on
Life, how maintained, 20 Muscles, 46, 54
Ligatures, Bernard's method of, 59 — on Snake-bite without Poison-ejection,
— in Snake-bite, 17, 106, 150, 152 8
— as used in Kogers* experiments, 153, Mode of Action55 ofet,Cobra-poison,
seq. 17 et seq.,
155,
159 Tables II., IIL, 156,
Mode of Entry of Colubrine Poisons in
Liouville, observations of, on Curare- relation
160 to rapidity of Death,
poisoning, 56
Liquor Ammonise, effect of, on Cobra- Mongoose {Herpestes gnseiis), supposed
poison, 4, 8, 37, 38, 39, Immunity of, to Snake-
109-10
poison, 39
Motor Impressions, effect on, of Cobra-
Liquor Potassa?, effect of, on Cobra-poison,
37, 138, 145 poison, 72, 76-7, 86-8
Liquor Potassoe Permanganatis, action of, Motor Nerves, Paralysis of, in Cobra-
on Guinea-pigs and llabbits poisoning, 58, 65, 69, 70
injected mixed with Cobra- effect of, on Respiration, 90-1
poison, 143, 145 Motor Nerves, Peripheral ends of
Lizards, action on, of Cobra-poison, 30 Paralysis of, from Cobra-poison, 20,
Local effects of Cobra- and Crotalus- 43, 56, 59 et seq.
poison, 113, experiments cause of Death in Curara-poison-
ing, 4
showing, 123-6 condition in, 81
Ludwig and Brunton, observations of, on
Eabbit's Ear, 97 condition of, after Elimination
Lymph-sac, Dorsal, of Frog, effects of of Curara, 4, 5, 6, 58
injection into Miiller, Johannes, on Contractions of the
of Cobra-poison, effects of, Pulmonary Veins and Vena
30, 50, 53, 59, 61, 62, 64, Cava, 136
65, 66, 67, 71, 73, 74, 76, Muscles, effect on, of Cobra-poison, 20,
77,79 43, 46 et seq., 54, 92, 128
of Crotalus.))oi^on, 122 possible, of iSnake-poison, 4, 5
— of Locomotion, effect on, of Cobra-
poison, 11, 24, 55, 56, 85
Mammaey Grlands, Cobra- venom excreted Muscular action, effect on, of Curare, in
Fish, 36
by, 104
 INDEX. 171
Muscular {cont.) Paralysis (cont.)
— System as affected by Cro^a^tts-poison, — after Sniike-poisoning, 18, 31, 32, 33,
112 and see Cases
55, 112
Mussel, Freshwater, Mantle of, effect on, Paramcecium, 131
effect on, of Cobra-poison,
of Cobra-poison, 130-1
Pedler, on Perchloride of Platinum as
destructive of activity of
Naja tripudians, see Cobra Cobra-poison, 137
Peltopelor, 23
]S"ausea, after 112
Cobra-poisoning, 10, 24, 70,
Perchloride of Iron, action of, on Cobra-
— after Colubrine -poisoning, 112 poison, 138
— after Cro^a^MS-poisoning, 112 on Guinea-pig, injected with
Nerve Centres, Cerebro-spinal, Pnralysis Cobra-poison, 145
of, as cause of Death from — of Platinum, destructive of activity of
Snake-bite, 4, 5, 7, 18, 55
Cobra-poison, Pedler's dis-
Nerve -periphery, possible affection of, in covery, 137 ; mode of action
Snake-poisoning, 4, 5 of, 137
Nerves {see also under Names), effect on, Peripheral Distribution of Motor Nerves,
of Cobra-poison, 52 Paralvsis of, see under Motor
Nerves
Nervous System, action on, of Snake-
poison, 55 et seq. Permanganate of Potash, see also Liquor
Rudimentary, in Infusoria, a query, Potassee Permanganatis
132 note action of, on Cobra-poison before
Neurotic character of Snake-poison, 18 entrance to Body, 138
Newts' blood, effect on, imperceptible, of on Cobra-poison in G-uinea-pigs,
Cobra-poison, 127 locally applied, 147, 148
seq.
Nitrate of Silver, action of as antidote to Snake-poison, various
on Cobra-poison, 138 experiments with, 149 et
on Gruinea-pig, injected with
Cobra-poison, 146 Antiseptic qualities of, 161
Escharotic action, superficial, 161
150
• Fayrer's precedence in use of, 149,
OphiocepJialus marulins, effect on, of
Cobra-bite, 35 Lacerda's use of, 150
Ophiophagus elaps, 23, Destructiveness Time occupied by application of,
of, 3 in Snake-bite, 155
Oxygen, Pure, effect of Respiration of, by Peritoneal Injections of
Poisoned
30 Animals, 111, 129- Cobra-poison, effects, 20, 41, 82,
83-4, 85, 86-8, 91, 117-8
Crotalus'-poison, 116, 118
Daboia-'poison, 25
Paeai-tsis after Cobra-poisoning, 11, 12, Phrenic Nerve, Paralysis of, in Cobra-
13, 21, 22, 26, 29, 38, 39, poisoning, 69, 91, 95
42,55 Pig, supposed25 Immunity of, to Snake-
of Bitten spot, 10, 57
of Fish, 35 poison, 40
Pigeons, effect on, of Daboia--poison, 24,
in Lizards, 30-1
of Medulla, effect of, on Respira- — Experiments on, with Cobra-poison and
tion, 90, 91 Permanganate of Potash,
of Motor Nerves, 58, 65, 69, 70, 71, 152, Table I., 154
effect on Respiration, 90-1 Pit-vipers of America and West Indies,
of Muscles, 47 et seq. virulence of, 23
of Phrenic Nerve, 69, 91, 95 — Venom of. Neutralised by Per-
Progressive, 10, 24, 55 manganate of Potash, 152
of Respiratory apparatus, cause of Platinum, see Perchloride of
Death, 4 Poison of Naja triptidians and other
of Secreting Nerves, 70, 108 Indian Venomous Snakes,
of Spinal Cord, 20, 56, 57, 65, 72 Nature and Physiological
— after Curara-povsoning, 4 Action of the. Part I., 1,
Fart II., 23
— after Daboia-poisoning, 27, 28
 172 INDEX.

Poison-glands of Snakes, 105 Eepfile?, Experiments on, with Snake.

Postmortem conditions in Animals after ! poisons, see Frogs, .Lizards,
Cro/fa 'tt.?-poi8oning, 113-14
Potash, Permanganate of, see Perman- i Eespiration {see also Artificial Eespira-
ganate of Potash, Sc Liquor [ j and "N"ewt8
tion), effect on, of
Potassa? Permanganatis I Cobra-poison, 4, 10, 12-17, 18, 20-2,
Preventing Death from Snake- bite, Expe- 24, 55, 56, 86-8, 89 e/, ye?.
riments on a Method of, 149 I of Crolahts-])o\Bor\, 112
Preventive Treatment in Snake-bite, when I of Curara-poi>oning, 4
solely of use, 7 I of Da/^c/m-poison, 25
Protoplasm {see also Vegetable do.), —
— Failure of,
Cobra- virus as Poison to, in cause
above, ofq.v. Death
132
in Somatic Death, 20
rsenihcMs porphyriacvs (Black Snake), — Function of the Brain in regard to, 20
Virus of,"of.
action 112 Death-causing — Hurried, after Cobra-poisoning, 10, 24,
26, 90
Ptyaline, in relation to the principle of causes of, 90, 92, 93
Cobra-venom, 9 note Ehythm of Independent Pulsation of
Puff Adder, Venom of, Neutralised by Pulmonary Vein, changes
Permanganate of Potash, i in, 134, 135, 136
152
Eichards, Dr. Vincent, Cobra-virus
Pulmonary Veins, and Vena Cava, Inde-
supplied by, 8
pendent Pulsation of, 134,
importance of, 13fi — Experiments by, on Cobra-virus, 8, 96
on Excretion of Snake-poison, 104
Pulsation, Independent, of the Pulmonary
Veins and Vena Cava, 134 on Permanganate of Potash as Anti-
dote to Cobra-poison, 150
on Treatment thereof by Liq. Am-
Eabbit, action on, of Cobra-poison, 10, I monise, 109
— on Physical changes produced by
12, 14, 21-2, 24, 43, 119
Snake-poisoning on the
on Circulation, 97, 99, 103
on Ee&piration, 92 ,. 113-4 by, on use of
II — and others, Blood,Experiments
of Crotalus-^o\s,on, injected, 119 Artificial Eespiration in
of Liquor Pot. Permang. injected Snake-bite, 107-8
with Cobra-poison, 143, 145
Rigor mortis in Cobra-poisoning, 49, 51,
— Blood of one poif=oned by Crotalus-
virus, effect of, on Gruinea-
I 84
Eogers, Leonard, Experimental Investiga-
pig, 121 tion by, into practical value
•— Cerebral operation
tion, 93 on, effect on Eespira-
of Fayrer's use of Perman-
— Experiments on, with Permanganate of ganate of Potash, and of
Potash and Ligature in Brunton's Lancet for speedy
Snake-poison, methods of, introduction thereof, 152
153 et sea., results, 157, — Animals and Birds experimented on,
158, Tables L, II., IIL, 154,156,
159 Tables II., IlL, 156, 159
Eat- Snake {Ftyas mucosa), effect on, of — Methods emj)loyed, 153 et seq.
bite of B^mgarus cceruleus, — Venoms dealt with, 152-3, Tables L,
33, of Cobra, 32, 33 II., III., 154, 156, 159
Eats, Blood of, effect on, of Cobra-poison, preparation of, 152, 153
44-6 Eussell, on Convulsions in Cobra-poison-
Eattlesnake, see Crotalus ing, 20
Eeagents, &c., e^eci of, on action of Eussell's Viper, see also Daboia Eiissellii
Cobra-poison, 36
Eeflex action, effect on, of Cobra-poison,
79-82, 95 Saliva, with Crystals of Permanganate of
— Centres, in Cobra-poisoning, 58, 72, Potash,
155 in Snake-bite, 152,
73
Eegion generally Salivary Grlands, probable Excretion by, of
153 attacked in Snake-bite, Snake-poison, 104
 INDEX. 17a
Salivation of Dogs, after Cobra-j)oisoning, Snakes, see Venomous Snakes, Sf under
Names
10, 2J., 60-70, cause, 70
Saiiguinolent Effusions as effect of Cobra- — Effect on, of Serpent-venom, 31 et seq.
and Croi^a^H* poison, 113, — Poison-glands of, 105
1)5, 123-6 Solution of Permanganate of Potash, for
Schiff, Observations
97 of, on Kabbit's Ear, Snake-bite
— first used by Fayrer, 149
— on Vomiting of Gruinea-pigs, 89 — • strength u*ed by Lacerda, 150
Secreting Xerves, Action on, of Cobra- strength used in tests on Pigeons,
poison, 69, 108 152, and Table I., 154
Seeds, G-ermination of. Action on, of Cobra- Somatic Death, causes of, 20
poison and of Rattlesnake- Somnolence,' after Cobra-poisoning, 55
poison, 40 Sphincters, relaxation of, in Cobra-poison-
Senac, Hypothesis of, on Contraction of
ing, 11, 24
Vena Cava and Pulmonary Spinal cord, effect oh, of Cobra-poison, 20,
Veins, 136 56, 57, 58, 65, 72 et seq.
Sensory Impressions, two kinds, effect on,
— effect on ("probable), of Croz'aZ^s-poison,
of Cobra-poison, 59-60, 72, 113
86-8 Stimulation followed by Exhaustion,
Sensory Nerves, Action on, of Cobra-poison, effect of Cobra-poison on
60, 70, 86-8 — Ciliary action,
— Muscle, 128 126, 130-3
Septic character of Snake-poison, 18
Serpent- venom {see also Snake-poison), Stimulation, not so followed, effect of the
Effect of, on Snakes, 31 et
seq. same on Vegetable Proto-
Stomach and Intestines,
plasm, 127-8 action on, of
Setschenow's Inhibitory Centres, Observa-
tions on, in Cobra-poisoning, Cobra-poison, 69, 89, 93-4
79 et seq. Strychnia and Woorara, mixture of, effect
Silver, see Nitrate of Silver of, compared with that of
Size of Animal in relation to quantity and Cobra-poison, 88
quality of Snake-poiscJn, 11, Subcutaneous Injection of
note Blood of Rabbit poisoned by
Snails, effect on, of Cobra-poison, 36 Crotalns-^o\s>or\. into Guinea-
Snake-bite, Experiments on an Easy
Method of Preventing Death pig, effect,
Cobra-poison, action121and effects
from, 149 et seq. on Cats, 83-4, 91
on Cobras, 33
— Loss of Life from, in India, 1, 2, 3 on Dog, 11
— Treatment of, proposed, 110
on Fowls, 21, 90
Snake-poison, action of, on Nervous
System, 55 on Frogs, 29, 41, 81
— Animals resisting, 39-40. 28
on Guinea-pigs, 13, 15, 16, 17,
— Antidotes to, 3, 4, 6 ^ note, 7, 8, 17,
19, 110, 137 et seq., 143, on Rabbits, 12, 14, 21, 41, 43,
149 et seq. 92,97
— Effects of, conditions governing, 18 of CrotaUis-^Q\:s>oxi, action of, on
on Snakes, 31 et seq. Guinea-pigs, 115
— Excretion of, 104-5, 106 et seq. of Curare, effect of, on Carp, 36
— Irritant, Neurotic and Septic character of Z)a5oia-poison, effects of
■ — ^ on Guinea-pigs, 27, 28
of, 18
on Pigeons, 25
— Mode of action of, 4, 5, 7, 18 et seq.
— Nature of, problems of, 6 of Permanganate of Potash, where
— Permanganate of Potash as antidote to, most effective, 150, 152,
first used by Fayrer, 149, 153, 160
other investigations into,
of Strychnia and "Woorara, effects
149-50 et seq. of, on Guinea-pig, 88
— Similaritv of mode of causing Death Swallowing of Cobra-virus, effects of,
in, 111-2
Snake-poisoning, Symptoms of, 10, 24, Svmptoms of 41-3
Cobra-poisoning, 10, 11, 24,
55, 112 55
Snake-stones, uselessuess of, 110 — of C/*o^aZM6-poisoning, 112-3
174 INDEX.

Tetanic arrest of Cardiac action induced Venomous Snakes, Bites of {see also Snake-
bj Cobra-, Daboia-, and bite), effect of, on them-
ffi/draphis--poison, 95, 104, selves, 31, 33-5, 105
112 Means of105Preventing Death from,
Thanatophidia of India, The, by Sir J.
Fayrer, 1, 17, 31, 39, 40, 42, Easy, of the same, 149 et seq.
106, 109, 110 note Indian, Poison of. Nature, etc., of,
Thoracic cavity. Injection into, of Cobra-
poison, effect of, 41, 50 1,23 Destructiveness of, 3
Relative
Tiger-snake of Australia {Hoplocephalus Venoms, see Snake-poisons
curins), Virus of, action of, Ventricles, Pulsation of, and of Pulmon-
112 ary Veins while Auricles
Tissues {see also Areolar Tissue), effect on, motionless, 135
of Cobra-poisoning, 10 Viperidse,
— Indian, African,
23 virulence of, 23
Tourniquet, improvised, in Snake-bite,
153 Viperine poison, local action of, 48
Transfusion of Blood, possible means of ■ Neutralised by Permanganate of
preventing Death through Potash, 152, 153, 158, 160,
Snake-bite, 109 Tables I., II., III., 154, 156,
159
Tree-snakes {Dendrophis picta), effect on,
of Cobra-bite, 32 Physiological effects of, 112, 113, 114
Trimeresuri, Low fatality from Bites of, 3 Viperine Snakes {see also Crotalus, Dahoia,
seq.
etc.), Blood-changes due to
Bite of, 18 Sf note
Unconscioitsness after Cobra-poisoning, Viscera, after Croi^afes-poisoning, 113 et
24
Voisin, Observations of, on Curare -
poisoning, 56
Yagus(i), division of, effect of Vomiting, after Cobra-poisoning
on Gruinea-pigs, 89 of Cats, 84, 89, 139
on Rabbit, 93 of Dogs, 10, 11, 24, 70, 89
— Inhibitory branches of, action on, of of auinea-pigs, 10, 24, 38, 89
Cobra- and Curare -poison- — after C rotalus-^o\%onva.g, 112
ing, 97 by Cat, 121
— Peripheral ends of, effect on, of Cobra- by Frog, 129 Sf note
poison, 69
Vallisneria, effect on, of Cobra-poison,
132-3 Wabm-bloodbd Animals, see under
Valvular disease of the Heart, value in, of Names
Independent Pulsation of Warmth, applications of, in cases of
Pulmonary Veins and Vena Curara-poisoning, 4
Cava, 136 Woorara, see Strychnia and Woorara
Vegetable Protoplasm, action on, of Snake-
(Cobra-) poison, Darwin on,
127-8 Xenurelaps bungaroides, 23
Vena Cava, Pulmonary Veins and, Inde-
pendent Pulsation of, 134,
importance of, 136 Zinc, see Chloride of Zinc

Haerison and Sons, Printers in Ordicary to His Majesty, St. Martin's Lane,
 3P Fayrer, (Sir) Joseph
941 On the poison of venomous
F3 snakes

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