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XII (B)

SECOND LETTER

25 October 1792

SOURCES

PRINTED HANDWRITTEN
Phil. Tr. P. 1. 1793, p. 27 

Ant. Coll. T. II, P. 1. p- 141

Cart. Volt. J 16, L 8, N 26

 

REMARKS

TITLE from Phil. Tr.

DATE from Phil. Tr. (§ 8 of Memoir III on Animal Electricity – letter to Aldini – V says this letter was written in August).

______________________

 

J 16 consists in various fragments written in French.

L 8 is a complete Ms. with common additions (§ 52 and part of § 52) and passages from handwritten sources of No. VIII ( C )

In Cart. Volt. No. 26, there is a letter of acknowledgement, thanks and congratulations from Sir

Joseph Banks, the President of the London Royal Society, dated London. 27 November 1792

 

SECOND LETTER

25 October 1792

(24.) Besides, it is clear that my observations, relating to the sciatic nerve and the leg, also apply to the brachial nerve and the arm. They also apply to any other nerve and to the muscles and limbs governed by those nerves.

(25.) These preparations are similar to those described by Mr. GALVANI. They prove that it is advantageous to expose the nerves and even better to dissect them free. This condition, however, is not absolutely necessary, because we obtain the same convulsions and motions of the limbs when we simply uncover the muscles and leave the nerves hidden under the muscles in their natural position, as shown by all my other experiments reported previously. (Section 21, 22, 23).

(26.) After performing these experiments on reptiles, birds and small quadrupeds, I studied larger animals: rabbits, dogs, lambs, bovines. The effects were not only similar to those described previously but even stronger and more durable, because the vital heat persisted for a longer time in those large animals and in their limbs. On the other hand I must mention that in most cold-blooded animals and particularly in frogs, viability persists for several hours in severed limbs. This prolonged viability, which renders the limbs so sensitive to weak electrical stimulations, lasts only a few minutes in warm-blooded animals and usually disappears before the animal heat has vanished.

(27.) After performing successful experiments in large and small animals of all species, be they intact or skinned, beheaded or variously dissected, and also on severed large limbs (most of the time without using the preparation called for by Mr. GALVANI, namely, without exposing the nerves) I decided to go further, that is, to study small limbs, or even single muscles or small fragments of a muscle. The new successes I thus achieved led me to other discoveries, which I shall present shortly, after describing some of my experiments.

(28.) EXPERIMENT E. In a frog, I severed a leg with the thigh attached, or a leg alone, or even one half or one quarter of a leg. By applying the tin foil to a part of the severed limb and the silver lamina to a different part, as usual and on establishing contact between the two metal plates, I always obtained some convulsions and movements. In some cases I separated a single muscle, e.g. the gluteus or the gastrocnemius or even a small fragment of those muscles, no larger than a grain of barley. The same effects were obtained, namely, application of two plates elicited brisk, spasmodic contractions of the entire muscle or fragment.

EXPERIMENT F. I repeated the same experiments on a whole leg, part of it, or a muscle or a muscle fragment, in chickens and other birds ; in slices of the gluteus muscle of a rabbit, lamb etc., and I obtained the same results, as long as the flesh was warm.

(29.) Thus, we elicit strong contractions in the muscles of warm-blooded and cold-blooded animals, and in all resected fragments of the muscles, This is achieved by directly applying the metal plates to the muscle itself, without preparing or exposing the nerves. On the other hand, we previously showed that we can stimulate the muscles by applying the metal plates to two neighboring parts of a nerve (Sections 19 and 20, experiments A and B). I draw the conclusion that it is not necessary to provoke a discharge of electric fluid between the nerve and the muscle or to move the fluid from the inside to the outside of the muscle through the nerve and the conducting circuit, or vice versa, as Mr. GALVANI assumes. I also conclude that our experiments do not suggest any similarity between the muscle and the Leyden flask. Where is the similarity with the flask, when the two metal plates, connected by a conducting wire, are applied, in close proximity to each other, on the surface of the same nerve (Experiments A and B) or on the surface of two similar muscles, or on the same muscle? (Experiments C, D, E, G). We must admit that any effort to claim a similarity with the Leyden flask would be vain.

(30.) EXPERIMENT G. If we apply silver foil to one thigh of a frog and a tin foil to the corresponding region on the other thigh, we elicit contraction of the muscles and the ordinary movements of the legs as soon as we connect the two plates with a conducting arc.

(31.) Is this the way, I wonder, to induce the discharge of two Leyden flasks, by connecting two homologous surfaces? Let us abandon these ideas of flasks and discharges and these far-fetched explanation; let us simply state that here, and in all similar experiments, we have a transport of electric fluid from one side to the other, provided the two sides are properly covered with metal plates. The transport is brought about not by a relative excess of fluid, which is difficult to imagine between two similar parts (of the body), but by the different nature of the plates, which must be made of different metals, as I explained previously (Sections 20 and 21. Experiments B and C) and in subsequent writings.

Actually

(32.) if two muscles, or two sites on the same muscle are similarly "armed", that is, they are covered with plates of the same metal, having similar tempering and hardness, flexibility or stiffness, smoothness or roughness, and the plates are applied in the same way, we can connect them with a conducting arc but will obtain no convulsion or movement.

(33.) I recognize that it is not easy to understand how and why, by applying two different metals to similar parts of an animal, and even to two neighbouring points on the same muscle, we disturb the balance of the electric fluid and cause it to flow continuously from one site to another; and why the flow starts as soon as we connect the two plates with a conducting object and lasts as long as the connection is maintained. Anyhow, whether or not we are able to understand their cause, these facts are proven by the experiments I described and will be confirmed by many others. At the end of the description, I will try to provide some explanations. These are facts that we must add to our present knowledge of electricity, even though they may seem extraordinary and difficult to reconcile with the established laws. What I have discovered is a new, very peculiar law, which does not relate to animal electricity, but to common electricity. In effect, the flow of electric fluid is not short-lived, like a discharge, but continuous, and persists as long as the connection between the two plates is maintained, whether the plates are applied to living or dead animal tissues, or to other non-metallic conducting objects, provided they are good conductors, like water or moist bodies. However, before I proceed to describe new experiments that definitely prove my contentions, I must briefly dwell upon those I have already reported (Section 20-32).

(34.) These experiments apparently suggest that we can elicit strong contractions in any muscle of any animal, provided they are viable, by simply applying two plates made of different metals. Such a conclusion, however, is too general and my experience taught me that some restrictions must be applied, relating to both the classes and genera of animals and to the different muscles of a given animal.

(35.) With regard to the different classes of animals: while it is true that all the quadrupeds, birds, fish, reptilians and amphibians which I tested do exhibit the phenomena I described, it remains true that worms and many insects do not. I vainly tried with earthworms, leeches, slugs, snails, oysters, caterpillars. I was unable to elicit any movements, even by means of small or medium-sized sparks or with discharges of artificial electricity. I proceeded as follows:

EXPERIMENT I. I applied the tin foil and the silver plate to various external and internal parts of slugs, leeches, earthworms, to the best of my ability. I established communication between the metal plates, either by bringing them closer and closer to each other until they touched, or by connecting them through another metal object. however, by performing all these manoeuvres, I never obtained any movement in any part of their body.

EXPERIMENT L. I made flask discharges to pass through their body, both insulated and non-insulated. The discharges were strong enough to elicit a medium size spark and to give me a little shock; no movements or convulsions appeared.

(36.) Shall we conclude that the most imperfect animals, the whole class of worms and many insects don’t have the same sensitivity and irritability, the same electrical mobility, if I may use this expression, which is observed in more perfect animals? I don’t want to draw this general conclusion from my experiments, which I performed, so far, on a small number of worms and insects. On the other hand, I must say that my experiments were successful, without great difficulty, in crayfish, scarabs, grasshoppers, butterflies and flies. It will be useful here to explain how I handle these animals, which are difficult to experiment with, due to their small size or because they are covered with scales.

EXPERIMENT M. After beheading the fly, butterfly, scarab etc., I cut their chest wall, over its entire length, with a penknife or small scissors and I introduce a piece of tin foil into the wound, near the neck (so-called silver paper is very convenient for this purpose). Then I deeply introduce, at a more caudal level, the edge of a silver plate or a small coin. When I push the latter object until it touches the tin foil, the legs start bending and wriggling and the other parts of the body also start shaking. It is amusing to elicit the song of a cicada etc. in this way.

(37.) Thus, it would be wrong to class the insects among those animals that are devoid of the electric properties we are discussing here. At most, if the caterpillars appear to lack these properties, we can say that in the larva stage, before they attain perfect shape through metamorphosis, and before they acquire new organs in the process, they are similar to worms in many respects, including lack of electrical sensitivity.

(38.) Finally, if I may express my thoughts here, only those animals that have well formed limbs, joints and muscles that bring about the motion of the limbs, in particular the flexor and elevator muscles, and the nerves which govern them, react with a real, spasmodic contraction to weak discharges of artificial electricity , or to a weak flow of electric fluid produced by two different metal plates. The spasmodic contractions provoke the motion and violent shaking of the limbs. By contrast, worms and those insects that have no well-developed limbs and joints, have no flexor muscles and exhibit only vermicular movements are not affected by this kind of electricity. This is a quite different animal organization, a different mechanism of motion, that has been discovered and explained for many species. These ideas of mine, which are still somewhat vague, are based on experiments. Further experiments will confirm or correct them.

(39.) With regard to different muscles in the same animal, I can speak with more certainty. It is not correct that any muscle will react with a contraction to the weak currents we use here. An important distinction must be made regarding their function in the animal organization. All muscles are not subject to the rule of will, and capable of producing spontaneous movements. I found that only those muscles that are governed by the will can be stimulated and produce a movement as a result of a weak flow of electric fluid produced by the contact with two different metals. This is not the case for those muscles that are not directly governed by the will, e.g., the muscles of the stomach, bowels, etc, including the heart, which is usually so irritable. The muscles of the diaphragm do react to the current (I predicted this response before making the experiment) because their motion is governed by the will.

EXPERIMENT N. It is surprising that a slice of good muscular flesh, cut out, e.g., from the thigh of a lamb that was slaughtered one half hour or one full hour before; it is surprising, I say, that this piece of muscle, quite cold, insensitive to any mechanical or chemical stimulation, is so strongly affected by the electrical fluid transmitted from one part to another, to the point of being seized by very strong spasmodic contractions; whereas the heart, recently removed from the same animal, when it is still warm and very irritable, is not affected at all by the same treatment, namely, the application of well fitting metal plates, connected by a conducting body. Again, it is surprising that the heart beat, when it is weak and slow, does not become more frequent; or, when the heart is quiescent or dormant, it does not wake up again, as it does after a very weak mechanical or chemical stimulation.

(40.) Thus, the electric fluid, which seems to be the proper stimulus for the muscles governed by the will, is not an effective stimulus for the heart and the other muscles whose animal and vital movements are not controlled by the will. But what would you say if I were to show that the electric fluid is not even the direct, effective cause of the movements of the "voluntary" muscles, that is, the muscles controlled by the will? In these muscles, the electric fluid is only an indirect cause, because it directly affects only the nerves. This is what I learned from many experiments, which forced me to abandon several attractive and much broader concepts. I was attracted by the idea, which was also advanced by Mr. GALVANI, that when the electric fluid hits the muscles with sufficient strength, it acts as a stimulus which arouses their irritability; that all muscular movements result from the flow of electric fluid into the muscles, whether we use the artificial electricity or the natural animal electricity; that the movements that occur naturally in the living animals, or at least the voluntary movements, were brought about by the same cause, namely, the direct action of the electric fluid on the muscles. However, I repeat, I had to abandon, with regret, all these attractive ideas that seemed to provide a wonderful explanation for our observations. Yes, we must assign certain limits to the action of electricity in animals, which should be conceived as being capable of directly exciting the nerves, as I previously stated and will now demonstrate [1]

(41.) To begin with, the fact that electricity acts on nerves, and that the nerves, thus stimulated, excite all the attached muscles, even without the current reaching the muscles, has been proved by Experiments A. and B., and also by an experiment by Mr. GALVANI, which was the first experiment of all, and originated the subsequent experiments, according to his own narration. In these experiments of the Professor from Bologna, and in my own experiments which I just mentioned , it is clear that the current crosses only a portion of the crural nerve and does not flow through any muscle of the leg. Nevertheless the muscles of the leg, which are controlled by that nerve, undergo convulsive contractions.

(42.) Moreover, I maintain that even when the electric current (here, I mean weak artificial discharges, or those currents that are generated by applying two plates of different metals) hits and penetrates the muscles that respond by movement, it does so by stimulating their nerves, not by direct excitation of the muscles themselves. This is shown by my Experiments C. and D. (Sect. 21 and 23), where the tin foil and the silver plate are directly applied to the muscles of the animal, whether whole or dismembered. Here, the muscles that show the most violent contractions are not those covered by the two metal plates, but those that are controlled by a main nerve trunk, which happens to be close to one of the plates. Thus, in a frog, when we apply the tin foil on the lumbar region, where the crural nerves are located at moderate depth, the leg muscles are seized by strong convulsions, more than any other muscle, including those that touch the other plate, that is, the silver plate. I already showed this behaviour in quadrupeds, dogs, lambs etc., in relation to the sciatic nerve (experiment D) and I want to add that the leg does not fail to be shaken when the nerve is not too deeply hidden under the skin and other tissues, provided we apply one of the plates to the proper location. This happens even when the other plate is not located near the gluteus muscle or any other leg muscle, if it is not too far away. This is the reason:

EXPERIMENT O.: After beheading a lizard and exposing its dorsal muscles by removing the skin, I apply a piece of tin foil to the truncated upper end of the torso, in such a way that the foil goes a little beyond the truncated area and reaches the shoulders; then, I place a silver coin over the middle of the spine. Finally, I slide the coin until it touches the foil. Immediately, the legs move, the tail bends tortuously and the entire body bends and jumps from right to left and vice versa. Isn't this due to the upper part of the spinal cord, the main origin of the nerves, being irritated?

(43.) By means of a similar operation [2]  we can obtain approximately the same result in mice, birds etc. However, in these animals we must remove not only the skin and other integuments, but also some flesh, because their back is more fleshy and the main spinal nerves and the spinal cord are hidden by this flesh and by the bones of the spine. It is easy to understand that the flow of electric fluid, produced by the two plates, can penetrate those parts of the body that are covered by the two plates, but only to some extent. In particular, it cannot reach the spinal cord, and the main branches of the nerves, which go into the limbs, if the bones, flesh and other integuments are very thick. Thus, we can easily understand why in large animals (dogs, lambs etc.) we do not succeed in eliciting the movement of all limbs by applying the plates to the back, even after stripping the flesh off. The large branches of the nerves remain too deeply hidden and buried in the tissues. Only a few branches or ramifications lie just under the plates and these branches reach only the adjacent, superficial tissues. As a consequence, we generally see only some shallow palpitations and contractions in a few muscles. If, occasionally, an entire limb is set in motion, this happens because the nerve that goes into that limb and controls its movement is not deeply buried, but is covered only by a thin layer of tissue, that separates it from the metal plates. This was the case in Experiments D and following trials (Section 23 etc.), where we obtained large movements in a dog's or lamb's leg by just applying one of the plates near the sciatic nerve; the thinner the layer of tissue surrounding the nerve, the stronger the movements of the leg.

(44.) Thus, to elicit full movements of the limbs in large animals, and not only superficial contractions and palpitations in a few muscles, it is necessary to know the position and direction of the nerves and to remove the common integuments, the fat, etc., and also to reduce the thickness of the tissues that cover and surround the nerves, before applying the metal plate. It may even be impossible to elicit those movements and convulsions in all the limbs simultaneously; whereas this is easily obtained in small animals, just by removing the skin and part of the other integuments, as previously shown (Sect. 42, Experiments O and P). This is not even necessary in frogs, where we can leave the skin in place because the skin, which is very thin and moist, does not prevent the electric current from reaching the main nerves and the spinal cord.

(45.) Thus, to elicit movements of the limbs, we must take into consideration both the direction of the main nerves and the position of the plates in relation to the muscles. Those muscles that are located in between the plates, or close to one of the plates, are more likely to produce spasmodic contractions, and are often the only ones that produce those contractions. This can happen, for instance, when the plates are not positioned near a large nerve trunk, or when the nerves are deeply located and surrounded by thick layers of tissue.

(46.) These observations and Experiments E, F (Sect 28) where a single muscle or even a fragment of muscle, treated in the usual way, does produce strong contractions , might suggest that the electric fluid elicits the movements by directly irritating the muscle fibres, without any nervous intervention. Thus, the role of the nerves would be neither primary nor strictly necessary, as I claim. However, this argument has no strength, so long as we don't demonstrate that those muscles and fragments do not contain nerve fibres. If, on the contrary, such fibres are present (and nervous branches must be present in every fragment that responds to the stimuli, even, I would say, in every muscle fibre), then I can affirm that it is the thin nerve filaments, which are distributed throughout the muscle, that are directly affected by the electric fluid when it flows through the muscular substance. The fluid acts on the sensitive nerves, and no further, and the nerves act on the muscles. Thus, I can maintain that, most likely, the electric fluid has no direct effect on muscular contractions, except insofar as it stimulates the nerves. Briefly, that the electric fluid is not the direct cause of the muscular contraction. This statement, which seems highly plausible in view of the findings that I described in the previous pages, will be verified, in the most obvious way, by a number of experiments which I performed on the tongue. These experiments led me to new discoveries, which are both interesting and curious.

(47.) Having succeeded in eliciting tonic convulsions and strong movements in the muscles and limbs of both small and large animals, without exposing any nerve, just by applying the plates made of different metals to the exposed muscles, I wondered whether we could not obtain similar results in humans. I realized that this could be achieved in amputated limbs. But how about intact, living human beings? This would have required removing the skin, performing deep incisions, cutting out part of the flesh in the regions where we planned to apply the metal plates (as I showed to be necessary in large animals). Fortunately, it come to my mind that the tongue offers to us an exposed muscle, not covered by the thick integuments that cover the rest of the body, a muscle that is very mobile, and controlled by the will. Here is, I thought, a muscle that offers all the necessary conditions for eliciting strong movements by means of the metal plates. With these considerations in mind, I performed the following experiment on my own tongue.

(48.) EXPERIMENT Q. After covering the tip of the tongue and part of its upper surface, over a few twelfths of an inch, with tin foil (so-called "silver paper" is most suitable for this purpose) I applied the convex part of a silver spoon further inside on the flat part of the tongue. By tilting the spoon I brought its tail into contact with the tin foil. I expected to see my tongue quiver and, to this end, I performed the experiment in front of a mirror. The expected movements did not materialize. Instead, I experienced an unexpected sensation, a strong, sour taste on the tip of the tongue.

(49.) At first, I was surprised by this event. However, after some reflection, I realized that the nerves that reach the tip of the tongue have the role of conveying taste sensations not of controlling motion in that flexible organ. Thus, it was natural that the irritation caused by the electric fluid should elicit a sensation of taste and nothing else. To produce movements in the tongue, it would be necessary to apply the metal plates near its root, where the motor nerves are located. I then verified this hypothesis with another experiment.

(50.) EXPERIMENT R. In a recently slaughtered lamb I cut the tongue out, near its root, and I applied tin foil near the cut end and the silver spoon on its surface. I then proceeded to establish a contact between the two metals, as usual. I had the satisfaction of seeing the whole tongue twitch vigorously, lift its tip, twist and bend on both sides every time and as long as the contact was maintained.

(51.) I repeated this experiment with the tongue of a calf, which I placed on a silver dish, after applying the tin foil to its root. The silver dish played the role of the second plate. The result was the same. I repeated the experiment by using the tongue of other small animals, like mice, chickens, rabbits, etc., and I obtained the same effect almost every time. I am saying "almost" because sometimes the effect failed in small animals. This may be due to the tin foil not being properly applied to the right place, where the motor nerves enter the tongue, or to the tongue being already cold. As I already pointed out (Sect. 26), viability is rapidly lost in warm-blooded animals, particularly in the tongue.

I am, etc.

October 25, 1792

A. VOLTA

Revised by John Coggan, Oxford University

 

Notes

[1]   J 16 here reads: "Thus, we must assign narrow limits to the action and influence of animal electricity. We would have loved to share GALVANI's idea, that electricity is the direct and effective cause of all muscular movements. Unfortunately, we cannot maintain these ideas because my experiments show that those muscles that are not controlled by the will do not respond to the action of electricity . Therefore, their movement must be due to a totally different cause. In effect, a fully viable heart, which responds with strong and frequent contractions to all sort of mechanical and chemical stimuli, is not affected (by electrical stimuli) ( translator’s addition) and maintains its rhythm without change, or remains still if it had already lost its rhythmical activity. [Editors’note]

[2] in LS we read: "preparation" [Editors’ note]

 

 

       
   

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