Part 2

The other nerves which issue from the brain exhibit no remarkable features. Those which originate in the spinal marrow are mostly derived from the ganglions, and are sometimes interwoven with the muscles, as the woof with the warp in a piece of cloth[99]; those from the three or four first commonly rendering to the muscles of the legs, wings, and other parts of the _trunk_, and those from the remainder to the _abdomen_. After their origin they often divide and subdivide, and terminate in numerous ramifications that connect every part of the body with the _sensorium commune_. A _pair_ of nerves is the most usual number that proceeds from each side of a ganglion[100]; but this is by no means constant, since in the louse, the hive-bee, and several other insects, only a _single_ nerve thus proceeds[101]; and in the larva of _Ephemeræ_, while _two_ pairs issue from the _six first_ ganglions, only a _single_ one is emitted by the _five last_[102]. In the spinal marrow of the rhinoceros-beetle, both larva and imago, the nerves consist of simple filaments which diverge like rays in all directions[103]: the same circumstance distinguishes the cheese-maggot, only some of the nerves appear to branch at the end[104]: in the louse, the last ganglion sends forth posteriorly three pairs of nerves which render to the abdomen[105]. Sometimes, though rarely, nerves originate in the _internodes_ of the spinal marrow. Cuvier indeed has asserted that in invertebrate animals _all_ the nerves spring from the ganglions, and never immediately from the spinal marrow; but Swammerdam, in describing those of the silk-worm, mentions and figures four pairs as proceeding from the four anterior internodes, excluding the first[106]; and at the same time he gives it as his opinion, that all the nerves in insects really originate from the marrow itself, and not from the ganglions, which he asserts are of a different substance, and are inclosed in the marrow for the sake of giving it greater firmness[107]. In this opinion, however, he seems singular[108]. Those remarkable nerves described by Lyonet under the name of _spinal bridle_ (_bride épinière_) also take their origin, not from the ganglions, but from a bifurcation of the spinal marrow. Of these, in the caterpillar of the goat-moth there are _ten_, the first issuing from the bifurcation of the internode between the fourth and fifth ganglions, and the remainder from the succeeding ones. After approaching the succeeding ganglion, these nerves form a pair of branches that diverge nearly at right angles from the bridle, and producing several lesser branches, lose themselves in the sides of the animal[109]. Besides the nerves above mentioned, two generally issue from the posterior part of the last ganglion, diverging in opposite and oblique directions: some of these render to the parts of generation; and in the silk-worm, and probably other species, the innermost pair is perforated for the passage of the _vasa deferentia_[110].

After duly considering this general outline of the nervous system of insects, the question will continually occur to you,--is then what you have called the _brain_ the _sensorium commune_ of these animals, in the same manner as it is in those with warm blood? To this query a negative must be returned. In the latter, the brain is the common centre to which, by means of the nerves and spinal marrow, all the sensations of the animal are conveyed, and in which all its perceptions terminate. The nerves and spinal marrow are merely the _roads_ by which the sensations travel; and if their communication with the brain, by any means be cut off at the neck, the whole trunk of the animal becomes paralytic, evidently proving that the organ by which it feels is the brain. This, however, is so far from being the case in insects, that in them, if the head be cut off, the remainder of the body will continue to give proofs of life and sensation longer than the head: both portions will live after the separation, sometimes for a considerable period; but the largest will survive the longest, and will _move_, _walk_, and occasionally even _fly_, at first almost as actively without the head, as when united to it. Lyonet informs us, that he has seen motion in the body of a wasp _three_ days after it had been separated from the head; and that a caterpillar even _walked_ some days after that operation; and when touched, the headless animal made the same movements as when intire[111]. Dr. Shaw has observed--an observation confirmed in Unzer's _Kleine Schriften_,--that if _Geophilus electricus_ be cut in two, the halves will live and appear vigorous even for a _fortnight_ afterwards; and what is more remarkable, that the _tail_ part always survives the _head_ two or three days[112]. The _sensorium commune_ of insects, therefore, does not, as in the warm-blooded animals, reside in the brain alone, but in the spinal marrow also. It was on this account probably that Linné denied the existence of a _brain_ in insects, regarding it merely as the first ganglion of the spine.

Cuvier and other modern physiologists, from the ganglionic structure of this organ, are of opinion that it is not the analogue of the _cerebro-spinal_ system of vertebrate animals, but rather of their _great sympathetic_ nerves. Indeed, considering solely the _external_ structure of the nervous system of insects, a great resemblance strikes us between it and these nerves; for besides its general ganglionic structure, there is also in them an _upper_ ganglion in the neck, seemingly corresponding with what we have named the brain of insects, from which the nervous chord dips to the lower part of the neck, where it forms a _second_ ganglion, which appears to correspond with what we have considered as their second ganglion[113]. We may observe, however, that at least in one respect there is even an _external_ resemblance between the brain of insects and that of vertebrate animals:--it most commonly consists, as has been stated, like them, of two lobes, often very distinct; a circumstance which not unfrequently distinguishes the other ganglions[114], and is not borrowed from the ganglions of the great sympathetics. With respect to the internal structure of the ganglions and spinal marrow of insects, we know little to build any theory upon, except that the internal substance of the former is filled with air-vessels; at least so Lyonet, as has been already observed, found in the goat-moth, while only the tunics of the latter are covered by them. Taking the above resemblance to the brain of vertebrates into consideration, there appears ground for thinking that the nervous system of insects, like some of their articulations[115], is of a _mixed_ kind, combining in it both the cerebro-spinal and the ganglionic systems; and this will appear further if we consider its _functions_.

That learned and acute physiologist Dr. Virey, assuming as an hypothesis, that the structure of the system in question is simply ganglionic, and merely analogous to the sympathetic system of vertebrate animals, has built a theory upon the assumption, which appears evidently contradicted by facts. Because, as he conceives after Cuvier, insects are not gifted with a real brain and spinal marrow, he would make it a necessary consequence that they have no degree of _intellect_, no memory, judgement or free will; but are guided in every respect by instinct and spontaneous impulses,--that they are incapable of instruction, and can superadd no acquired habits to those which are instinctive and inbred[116]. This consequence would certainly necessarily follow, was their nervous system perfectly analogous to the sympathetic of warm-blooded animals. But when we come to take into consideration the _functions_ that in insects this system confessedly discharges, we are led to doubt very strongly the correctness of the assumption. Now in these animals the system in question not only renders to the nutritive and reproductive organs, which is the principal function of the great sympathetic nerves in the vertebrates; but by the common organs maintains a connexion with the external world, and acquires ideas of things without, which in them is a function of the cerebral system: from the same centre also issue those powers which at the bidding of the will put the limbs in action, which also belongs to the cerebral system. That insects have memory, and consequently a real brain, has been before largely proved, as also that they have that degree of intellect and judgement which enables them to profit by the notices furnished by their senses[117]. What can be the use of eyes,--of the senses of hearing, smelling, feeling, &c. if they are not instructed by them what to choose and what to avoid? And if they _are_ thus instructed--they must have sufficient intellect to apprehend it, and a portion of free will to enable them to act according to it. With regard to the assertion that they are incapable of instruction, or of acquiring new habits; few or no experiments have been tried with the express purpose of ascertaining this point: but some well-authenticated facts are related, from which it seems to result that insects may be taught some things, and acquire habits not instinctive. They could scarcely be brought from their wild state, and domesticated, as bees have been so universally, and both ants and wasps occasionally[118], without some departure from the habits of their wild state; and the fact of the corsair-bees, that acquire predatory habits before described[119], shows this more evidently: but one of the most remarkable stories to our purpose upon record, is that of M. Pelisson, who, when he was confined in the Bastile, tamed a spider, and taught it to come for food at the sound of an instrument. A manufacturer also in Paris, fed 800 spiders in an apartment, which became so tame that whenever he entered it, which he usually did bringing a dish filled with flies but not always, they immediately came down to him to receive their food[120].

All these circumstances having their due consideration and weight, it seems, I think, most probable, that as insects have their communication with the external world by means of certain organs in connexion with their nervous system, and appear to have some degree of intellect, memory, and free will, all of which in the higher animals are functions of a cerebral system, and at the same time in other respects manifest those which are peculiar to the sympathetic system,--it is most probable, I say, as was above hinted, that in their system _both_ are _united_.

I must bespeak your attention to a circumstance connected with the subject of this letter, which merits particular consideration: I mean the gradual change that takes place in the nervous system when insects undergo their metamorphoses; so that, except in the _Orthoptera_, _Hemiptera_, and _Neuroptera_ Orders, in which no change is undergone, the number of ganglions of the spinal chord is less in the imago than in the larva. There seems an exception indeed to this rule in the case of the rhinoceros-beetle, in the larva of which there is only _one_ ganglion, while in the imago there are _four_[121]. But as this one ganglion occupies the whole spinal marrow, it is really of greater extent than the four of the imago; so that even in this case there is a concentration of the cerebral pulp. In some cases, as in _Dytiscus marginalis_, and _Hydrophilus piceus_[122], the imago has only _one_ ganglion less than the larva, but more generally it loses _four_ or _five_. Dr. Herold has traced the gradual changes that take place in the spinal marrow of the common cabbage-butterfly (_Pontia Brassicæ_), from the time that it has attained its full size to its assumption of the imago. Of these I shall now give you some account.

In the full-grown caterpillar, besides the brain there are _eleven_ ganglions, the chords of the four first internodes being double, and the rest single: from each ganglion proceed two pairs of nerves, one from each side. In this the lobes of the brain form an angle with each other[123]. In two days the double chords mutually recede, so as to diminish the interval between the ganglions, and the single ones have become curved: thus the length of the spinal marrow is shortened about a _fourth_, and the fourth and fifth ganglions have made an approach to each other[124]. On the eighth day, when the insect has assumed the pupa but remains still in the skin of the caterpillar, the flexure of the internodes is much increased; the first ganglion is now united to the brain, and the fourth and fifth have joined each other, though they are still distinct; the spinal marrow has now lost considerably more than a _third_ of its length[125]. On the fourteenth day, the internodes, except the double ones, have become nearly straight again; the fourth and fifth ganglions have coalesced so as to form one, and the sixth and seventh have each lost their pairs of nerves[126]. Shortly after this, these last ganglions have nearly disappeared, and the chords of the three first internodes have again approached each other[127]. The next change exhibited is the absorption of the first ganglion by the brain, the union of the chords of the first internode, which is now straight, the approximation of the second and third ganglions, and the enlargement of the one formed by the union of the fourth and fifth, at the expense perhaps of the sixth and seventh, which have now intirely disappeared, and in their place is a very long internode. These united ganglions retain the pairs of nerves they had when separate[128]. Just before the assumption of the _imago_, the direction of the lobes of the brain becomes horizontal, the second and third ganglions unite, and the internode between the third and fourth is shortened[129]. Lastly, when the animal is become a butterfly, the second and third ganglions have coalesced, and are joined to that formed by the union of the fourth and fifth; a short isthmus or rather constriction, with an orifice, being their only separation: each of these united ganglions send forth laterally four pairs of nerves[130]. In his figure, Dr. Herold has not represented the orifice for the passage of the gullet, but doubtless one exists, which for an animal that imbibes only fluid food is probably very minute. In _Hypogymna dispar_, we learn from Cuvier, this orifice is of that description, and of a triangular shape[131].

It can admit of no reasonable doubt that one of the principal intentions of these changes is to accommodate the nervous system to the altered functions of the animal in its new stage of existence, in which the antennæ, eyes, and other organs of the senses, as well as the limbs and muscles moving them, and the sexual organs, being very different from those of the larva, and if not wholly new, yet expanded from minute germs to their full size, may well demand corresponding changes in the structure of the nervous system by which they are acted upon.

But are these changes also concerned, as Dr. Virey conjectures, in producing that remarkable alteration which usually takes place between the _instincts_ of the larva and imago? In order to answer this question, it will be requisite first to quote the ingenious illustration with which this able physiologist elucidates his ideas on this point. "The more readily," he observes, "to comprehend the action of instinct, let us compare the insect to one of those hand-organs in which a revolving cylinder presents different tunes noted at its surface, and pressing the keys of the pipes of the organ, gives birth to all the tones of a song: if the tune is to be changed, the cylinder must be pulled out or pushed in one or more notches, to present other notes to the keys. In the same manner let us suppose that nature has impressed or engraved certain determinations or notes of action, fixed in a determinate series in the nervous system and the ganglions of the caterpillar, by which alone she lives, she will act according to a certain sequence of operations; and, so to speak, she will sing the air engraven within her. When she undergoes her metamorphosis into a butterfly, her nervous system being, if I may so express myself, pulled out a notch, like the cylinder, will present the notes of another tune, another series of instinctive operations; and the animal will even find itself as perfectly instructed and as capable of employing its new organs, as it was to use the old ones. The relations will be the same; it will always be the play of the instrument[132]."

This illustration is doubtless at the first glance very striking and plausible: but a closer examination will, I think, show, that, as in so many other instances in metaphysical reasoning, when fanciful analogies are substituted for a rigid adherence to stubborn facts, it is satisfactory only on a superficial view, and will not stand the test of investigation; and as this is a question intimately connected with what I have advanced on the subject of instinct in a former letter, I must be permitted to go somewhat into detail in considering it.

To prove his position, Dr. Virey ought at least to be able to show that, whenever a change takes place in the instincts of insects in their different states of larva and imago, a corresponding change takes place in the external structure of the nervous chord. But what are the facts? In three whole orders, viz. _Orthoptera Hemiptera_, and _Neuroptera_, as mentioned above[133], the structure of the nervous chord is _not_ changed; and yet we know that many tribes of these orders acquire instincts in their imago state altogether different from those which directed them in their state of larvæ. A perfect _Locust_, for instance, acquires the new instincts of using its wings; of undertaking those distant migrations of which so many remarkable instances were laid before you in a former letter[134]; and, if a female, of depositing its eggs in an appropriate situation. But if such striking changes in the instinct of these tribes can be effected without any perceptible alteration in the structure of the nervous chord, it is contrary to the received rules of philosophical induction to refer to this alteration the changes in the instincts of other tribes where it is found. Is it not far more probable that this alteration has in fact no connexion with the changes of instinct, but is solely concerned with those remarkable changes in the organs of sense and motion, which occur in the larva and imago states of the orders in which it is observed? In a common caterpillar, the form of the body, the legs, the eyes, and other organs of the senses, all strikingly differ from those of the imago; whereas, with the exception of the acquisition of new wings, a perfect locust differs little from its larva: so that we may reasonably expect a corresponding change, such as we find it, in the structure of the nervous chord of the lepidopterous insect, not called for in that of the neuropterous species, in which accordingly it does not take place.

This reasoning, in opposition to Dr. Virey's theory, that the changes of instinct depend on the altered structure of the nervous system, becomes greatly strengthened when we advert to the higher classes of animals, which surely in any investigation of the nature of instinct ought to be closely kept in view; for the faculty, though often less perfect in them than in insects, is still of the _same kind_, and may consequently be expected to follow the same general laws. In a young swallow, for example, all its instincts are not developed at once any more than in an insect. The instinct which leads it to migrate does not appear for some months after its birth, and that of building a nest still later. But we have not the slightest ground for believing that these new instincts are preceded by any change in the structure of the great sympathetic nerve, or of any other portion of the nervous system: and the same may be said as to the sexual instincts developed in quadrupeds some years subsequent to their birth. If, then, these remarkable changes in the instinct of the higher classes of animals can take place independently of any visible change in the nerves, what substantial reason can be assigned why they may not also in the class of insects?

On the whole, I think you will agree with me, that there is nothing in Dr. Virey's hypothesis which should lead me to alter the opinion I have already so strongly expressed in a former letter[135], as to the insufficiency of the mechanical theories of instinct hitherto promulgated, adequately to explain _all_ the phenomena; and unless they do this they are evidently of small value. Such theories as I have there adverted to may often seem to be supported by a few insulated facts, but with others, far more numerous, they are utterly at variance; and, to omit many other instances, I am strongly inclined to doubt the possibility of satisfactorily explaining the _variety_ of instincts exercised by a bee[136], or the _extraordinary_ development of new ones in particular circumstances only[137], on any merely mechanical grounds.

And after all, even suppose it could be demonstratively shown that _every_ instinct is as clearly dependent on secondary causes, as I have formerly admitted that _some_ doubtless seem to be, yet what would this teach us as to the essential nature of instinct? We have advanced indeed a step; but still, as I have before observed in referring to the theories of Brown and Tucker, we have only placed the world upon the tortoise, and instinct, as to its _essence_, which is what we want to detect, is as mysterious as ever: just as, though we can clearly prove that the mind is acted upon by the senses, yet this throws no light upon the essential nature of the mind, which we are forced to admit is inscrutable, as if to teach us humility, and prevent our vainly fancying, that though allowed to discover some of the arcana of nature, we shall ever be able to penetrate into her inmost sanctuaries.