Experiments and Observations Relative to the Influence Lately Discovered by M. Galvani and Commonly Called Animal Electricity

Part 3

Chapter 34,002 wordsPublic domain

But the most important, and characteristic difference, which I have yet been able to discover, between this new influence and electricity, consists in their effects upon the contractile power of animals and of plants. The contractions of animals excited by electricity have a tendency to destroy that power upon which contractions depend. But the contractions excited, by the application of the metals, have, in all my experiments, had the directly opposite effect. The more frequently contractions have been, in this way, excited, the longer do they continue excitable: and the longer are the parts, upon which such experiments are made, preserved from putridity. An influence, capable of exciting contractions, without occasioning exhaustion, was a thing I so little expected to find, and so contrary to the character which had been given of this, both by Galvani and by Dr Valli, that I, at first, distrusted my own observation of the fact: but the number of comparative experiments, which I had afterwards occasion to make, though with views different from that of ascertaining the point in question, convinced me that this influence, so far from destroying the contractility of muscles, has a tendency to preserve it. Oxygene is, so far as I know, the only stimulus in nature, whose effects are at all analogous.

When a frog has been long dead, I have been sometimes more than a quarter of an hour without being able to excite a single contraction by the application of the metals: but after this, without at all varying the means employed, contractions have appeared, and have become gradually more and more vigorous.

It is said, (for I have never had an opportunity of making the experiment,) that a stream of electricity passed through a sensitive plant produces an almost immediate collapse of its leaves. But the influence, discovered by Galvani, produced no such effect in the following experiment. Having separated the leg of a frog from its body, I freed its crural nerve from surrounding parts, and with one hand held it supported upon the end of a probe. An assistant placed a piece of silver under its foot, and held the zinc with which it was to be touched. A sensitive plant formed the medium of communication between us. He held the bottom of its stem between his fingers, while I held the top: so that when the silver was touched by the zinc, the influence passed up the plant, and through the whole of its stem. The frog’s leg instantly contracted, and repeated its contractions every time the silver and zinc were in contact: but the leaves of the plant did not collapse; neither did they when any of its branches formed part of the circuit.

I must, however, confess that the plant, upon which this experiment was made, had been kept through the winter. With a young one the result might possibly be different; but such an one I have not yet had it in my power to procure.

The torpedo does not appear at all affected by the influence which itself produces. Animals, in which Galvani’s phenomena are produced, are strongly affected. From this circumstance, and still more from the presence of metals being absolutely requisite to their production, some may be induced to believe, that the influence, which causes them, is something external to animals; and that it arises from the mutual contact of the metals only. I must confess I was, for some time, inclined to entertain this opinion; and its probability appeared to be not a little increased by observing that its effects differed with the metals employed, and were strongest when their surfaces were extended, and applied horizontally to each other. I began, therefore, to suspect that it might be some hitherto undiscovered property of metals; for that it was not an electrical phenomenon, seemed still further proved by the circumstance above related. It has been demonstrated, by the very interesting discoveries of M. Volta, that, ‘wherever the capacity of holding electricity is greater, there the intensity of electricity is less’:--‘and that the capacity of a conductor is increased, when, instead of remaining quite insulated, the conductor is presented to another conductor not insulated; and this increase is more conspicuous, according as the surfaces of those conductors are larger, and come nearer to each other[10].’

When, therefore, a plate of silver, communicating with the leg of a frog, was laid upon glass, and a plate of zinc was lowered horizontally upon it, the capacities of both, for any electricity which they might have contained, must have been so much increased, that no one will suspect the contractions of the frog’s leg, to have been occasioned by any discharge of the electrical fluid from them.

As little are we authorised to suppose, that the contractions were produced in consequence of the metals attracting the electrical fluid from the leg: for, since the leg was insulated, it is impossible that it should have received a new supply of electricity, after having been deprived by the metals of what it naturally possessed; and consequently, after once or twice contracting, no further contractions should have taken place: but this is contrary to the fact.

I have before shewn, that flakes of gold leaf, placed between the metals, were not affected by their approach to each other; and that, besides, a quantity of electricity, sufficient strongly to affect an electrometer, was far too weak to excite contractions in the muscles of a frog.

That this influence, however, whatever it be, is not derived from the metals alone, but that animals at least contribute to its production, as well as indicate its presence, is, I think, rendered highly probable, by what I have already urged, relative to the necessity of a communication between the metals, and the muscles, as well as between the metals and the nerves.

I may likewise observe, that animals appear to have a much more complete controul over its effects, than one would expect them to have over an influence wholely external to them.

When living and entire frogs are placed upon a plate of zinc, or tin-foil, and a piece of silver, or of gold, is passed over different parts of their legs, and thighs, till it come into contact with the plate; contractions are very seldom produced, and scarcely ever, if the frogs be healthy and upon their guard. But the instant their sciatic nerves are divided, the contractions produced are as free and vigorous, as if the legs had been completely separated from the body. This difference is not owing to the silver coming in contact with the wound, necessarily made in order to divide the nerve; for I have always taken care that it should not, and indeed when it did, no contractions were produced, unless the nerve had been divided.

Taking off the head of an animal, or intercepting, in any way, the influence of its will upon the muscles of the part excited, has precisely the same effect. But the will is not able to controul the effects of electricity, when the electricity is otherwise sufficiently strong to excite muscles to contraction. I have repeatedly found that even by the strongest voluntary contractions of the muscles of my arm, I have not been able altogether to counteract the involuntary ones, produced by electrical sparks, nor have I found that frogs could ever counteract them.

On attending carefully to the state of the muscles of the legs of living frogs, at the instant the metals were applied, I could perceive by the touch, that, in many frogs, though by no means in all, their muscles were perfectly soft and relaxed: a proof that they have other means of counteracting the involuntary contractions, which the metals have a tendency to produce, besides keeping their muscles in a state of permanent and voluntary contraction.

Footnote 1:

Cavallo.

Footnote 2:

On this metal Cronstedt has the following very curious remark: “It seems to become electrical by friction, and then its smaller particles are attracted by the loadstone; which effects are not yet properly investigated.” Zinc is an ingredient of the best amalgam for smearing the rubbers of electrical machines: But I have not been able to render a bar of zinc electrical by friction, nor to find that its smaller particles were in any state attracted by the loadstone, unless they had been scraped off by means of an instrument of iron. But, in this way, the dust of any metal is rendered susceptible of the influence of the loadstone.

Footnote 3:

If further experiments should establish decidedly, that the mutual contact of two different metals is absolutely necessary for the productions of Galvani’s phenomena, may not this circumstance afford an useful test of the purity of the precious metals? For instance, contractions in an animal produced by the contact of a piece of gold or silver, whose purity we wish to ascertain, with a piece of the same metal known to be pure, would then prove incontestably the presence of alloy.

Footnote 4:

In an able lecture, which Dr Monro lately delivered, chiefly upon this subject, he demonstrated the possibility of exciting contractions in the limb of a frog, without either of the metals he employed being in contact with it; or having any other communication with it than through the medium of some moist substance. In varying this experiment, I find, that if a frog be divided in two parts, just above the origin of the sciatic nerves, and put into a bason of water, the hind legs may be thrown into strong contractions, by bringing zinc, or tin-foil, and silver, in contact with each other, at the distance of at least an inch from the divided spine, so long as they are kept nearly in a right line with it. Water, in this case, is the only communication between the metals and the origin of the nerves.

Footnote 5:

The contractions produced seemed to be strong in proportion to the extent of the surfaces of the metals in contact, strongest when a large plate of zinc is laid horizontally upon a large plate of silver or gold. If the zinc be suffered to remain in contact with the silver, for a little time, the contractions of the leg cease. The zinc may then be slid over the silver, till it even touch the leg without renewing the contractions: but, in withdrawing the silver, the leg contracts at the instant the silver parts from it!

Footnote 6:

‘L’eau trop échauffée, ou qui est en éboullition, disperse l’électricité, de manière à en détruire les phenomènes.’

‘L’excès du froid prive l’eau même de la propriété de conduire le fluide en question.’--_Dr Valli, Lettre 9me._

Footnote 7:

It was in this way, indeed, that I have always excited contractions, when I have employed this new mode of influencing animals, as a test of remaining life in any part of them.

They were constantly kept in fresh water, as the situation most natural to them, during the whole of the time they were under experiment; and their skins were suffered to remain as entire as possible, since I found their muscles lost their contractile power, in a few hours, and became rigid when exposed, deprived of their skins, to the action of the water.

Footnote 8:

M. Fontana, in the first volume of his work on Poisons, mentions some facts, which may, to some, appear to give considerable countenance to this explanation. The microscopical eels found in dry and smutty wheat; the seta equina or gordius of Linnaeus; and the wheal polypus, all, when dry, become apparently dead: but again recover motion and life when moistened with water. One of the latter was put, by M. Fontana, upon a bit of glass, and exposed, during a whole summer, to the noon-day sun. It became so dry that it was like a piece of hardened glue. A few drops of water, however, did not fail to restore it to life. Another was, in this way, recovered after a similar exposure of a year and a half. Father Gumillo, a Jesuit, and the Indians of Peru, are quoted by the same author, on the authority of Bonguer, as speaking of ‘a large and venemous snake, which being dead and dried in the open air, or in the smoke of a chimney, has the property of coming again to life, on its being exposed, for some days, to the sun, in a stagnant and corrupted water.’

But it would almost require the credulity of an Indian to credit the testimony of the Jesuit.

Footnote 9:

Since what I had before written upon this subject went to the press, I have been informed by a friend, that Dr Lind of Windsor has found, that contractions may be excited in a frog by touching it with iron alone. In a frog very recently killed, I have myself, sometimes, excited contractions, by touching its nerves with iron and steel in conjunction. But I can by no means consider this as a satisfactory proof, that contractions may be excited by the contact of one metal alone; since I have never been able to excite contractions with a piece of iron, of the same quality throughout, applied to a frog which had been so long dead as to leave no suspicion that the contractions were occasioned by mechanical irritation. In Dr Valli’s experiment, with scissars of bad steel, upon a frog recently killed, these circumstances do not appear to have been sufficiently attended to.

Footnote 10:

Phil. Trans. vol. 72. part i. Appen.

SECTION II.

_Has Magnetism any concern in the Phenomena discovered by Galvani?_

In answer to this question I have little to say, as the experiments which it suggested, and which I had an opportunity of making, have been but few.

I have repeatedly excited contractions, both with the natural and the artificial loadstone, but I could never observe any difference between them, and such as were excited by unmagnetised iron, or an ore containing an equal quantity of iron with the natural loadstone.

When the separated leg of a frog was laid upon a plate of iron, and a loadstone was brought in contact both with its nerve and the plate, no contraction was excited. I have often brought frogs, in every state of preparation, as nearly as possible to a very sensible magnetic needle, but no variation in its direction was in any case produced by the contractions of the frogs excited by the metals.

SECTION III.

_What are the relations which subsist between the influence discovered by Galvani, and the muscles, the nervous, and the vascular systems, of animals?_

In proposing to myself a question of this very extensive nature, it will hardly be imputed to me, that I ever entertained, for a moment, the idle expectation of being able completely to solve it. It is prefixed to the following experiments as the most commodious general head under which I could arrange, not only what I had further to say, upon the influence discovered by Galvani, but likewise upon the several physiological subjects, in the examination of which this influence was employed merely as a test.

OF THE MUSCLES.

As I am acquainted with no criterion by which we can assure ourselves of the complete separation of muscular fibres from nerves, without rendering them objects too minute for accurate experiment; it can never be in our power, so far as I am able to judge, to satisfy ourselves, if this new influence can act immediately upon the muscular fibre. A doubt must always remain, whether nerve has not been present; and from this doubt will arise another still more difficult to solve, whether the influence produced or excited by the metals have passed through the nerve to the muscles? or if it have merely acted as a stimulus to the nerve, serving to rouse that unknown energy, by which nerves are known in certain circumstances to excite muscles to contraction.

The following experiments, made upon animals considered by anatomists, in general, as destitute of nerves, may to some appear decisive of this question, but to myself, I confess, they are by no means so. In by far the greater number of animals, we are precluded from the possibility of discovering nerves by their minuteness; yet the actions of these animals, not merely excited by mechanical irritation, but so obviously directed to the attainment of an end, oblige us to infer their existence even where our senses, aided by the best glasses, do not enable us to detect them.

Having laid some earth worms upon a plate of zinc, I tried to excite contractions in them, by passing a rod of silver over different parts of their length, till it came in contact with the plate; but for a long time without producing any effect. Application of the metals to a part recently divided seemed to produce as little effect. At length, I perceived one of them dart itself forwards, whenever the silver was passed under its belly near to a part which had been divided and rejoined. On repeating the experiment again, and with more care, I found, (as in the frog,) that when the animal was perfectly lively, and upon its guard, no contraction could in this way be excited; but that when a part had been rendered more sensible by previous disease, recent irritation, &c. or when the worm was taken unawares by hanging it over a probe, and lowering both upon the plate at the same instant; a sudden and involuntary motion seemed to dart through a great part of the worm’s length from the part touched towards the head; a direction contrary to that in which it takes place in other animals. I never could produce the same effect upon leeches. On varying the experiment, a most whimsical, but satisfactory phenomenon presented itself. I had laid a leech upon a crown piece of silver, placed in the middle of a large plate of zinc. The animal moved its mouth over the surface of the silver without expressing the least uneasiness; but having stretched beyond it and touched the zinc plate with its mouth, it instantly recoiled, as if in the most acute pain, and continued thus alternately touching and recoiling from the zinc, till it had the appearance of being quite fatigued. When placed wholly upon the zinc, it seemed perfectly at its ease; but, when at any time its mouth came in contact with the silver lying upon the zinc, the same expression of pain was exhibited as before.

With the earth worm, this experiment succeeded still more decisively. The animal sprang from the zinc in writhing convulsions; if, when the worm stretched itself forwards, one of its folds lit upon the zinc, it expressed little uneasiness in comparison of what it shewed when the point of its head touched the zinc.

These extraordinary effects were, however, considerably different from those produced by the metals upon the limbs of frogs, and other animals. They had not so much the appearance of involuntary, instantaneous convulsions, as long continued expressions of pain and disgust; such as are produced by applying zinc and silver to the tongue of a child.

A strong presumptive proof, in my humble opinion, that these animals are endowed with a most exquisite organ of sense, and, consequently, that they are not, as has been supposed, destitute of a nervous system.

Doubtful, therefore, if this influence can ever act upon the muscular fibre, except through the medium of nerves, I shall reserve what I have to say upon particular muscles, till I have related some facts relative to the nerves.

OF THE NERVES.

It appears from every experiment, which has been made in prosecution of Galvani’s discovery, that the nerves are very essentially concerned, in all the phenomena which it exhibits. It becomes, therefore, an object of inquiry, highly interesting, to ascertain if all the nerves of the body are equally subjects of this new influence, or if its effects are confined to those appropriated to muscles of voluntary motion. With this view, I surrounded with tin-foil the parvagum and intercostal nerves of several cows and sheep, while the auricles of their hearts were still contracting, and placed one end of a bent silver rod, at one time upon the heart itself, at another upon adjacent muscles, and sometimes upon the nerves; but all without producing the slightest perceptible variations, in the contractions of the heart, or a renewal of them when they had ceased.

I likewise included the caroted artery in the tin-foil; and, at another time, inserted the foil in longitudinal incisions made in the nerves, that it might be more immediately in contact with their substance; but still no contractions followed. I had as little success when I made similar experiments upon a dog, cats, rabbits, fowls, and frogs; yet, in all these animals, I could in general excite vigorous contractions, by arming the nerves of parts obedient to the will: I say in general, for in rabbits I have sometimes failed altogether; especially when they have been drowned in very cold water. Soon after making these experiments, I perceived from one of Dr Valli’s letters, published in the Journal de Physique, that he had made a similar one upon the heart of a dog, and with the same result. The heart, through the medium of its nerves, is not excitable, therefore, by the same means which are found efficacious in exciting other muscles to contraction. I confess I had not expected this result. It has been asserted indeed, by many physiologists of the first name[11], that the heart can in nowise be affected by the application of a stimulus to its nerves, or to the brain; but many considerations excited my doubts upon this subject, and some experiments which I made at this place, more than a year ago, tended to confirm me in an opposite opinion. That both the frequency, and the strength of the heart’s contractions are affected by passions of the mind, is a fact known to every one; but what is much more to the purpose, since we know so little either of mind or of its mode of influencing the body, we know that many derangements of the brain, such as apoplexy, hydrocephalus, phrenitis, &c. together with all kinds of mechanical injuries, (and what are these, but so many stimuli irritating the brain, and consequently the nerves sent to the heart?) affect the motions of the heart most materially and obviously. The contractions of the heart, so long as the brain remains entire, may be affected by a thousand different substances thrown into the stomach; but it appears from the experiment of Mr Kite, that this is by no means the case, when the functions of the brain are suspended by hanging, or drowning[12]. Dr Whytt’s experiment on this subject is one of the most decisive with which I am acquainted. He found, that opium operates much more slowly in destroying the heart’s motion in frogs, deprived of their brain and spinal marrow, than it does when these animals ate entire. Several of my own experiments, though not made expressly with this view, gave the same result with those of Dr Whytt. M. Fontana tells us, he has discovered the heart of the wheel polypus to be a voluntary muscle. It was probably this discovery which led him to try the effects of his will upon his own heart. For the success of his experiment, we have the testimony of his friend Dr Gerardi, Professor of Anatomy in the University of Parma, who, in a very learned little Dissertation on the Origin of the Intercostal Nerve, published in the Journal de Physique for September last, makes the following short mention of it; ‘Je ne dois point oublier de vous dire que M. Fontana a la faculté d’accélérer, ou de retarder à volonté son pouls, sans aucune contraction sensible des muscles.’