INTERNAL ANATOMY AND PHYSIOLOGY OF INSECTS, CONTINUED.
CIRCULATION.
We learn from the highest authority, that the blood is the life of the animal[348]: every object of creation, therefore, that is gifted with animal life, we may conclude, in some sense, has blood, which in this large sense may be defined—The fluid that visits and nourishes every part of a living body[349]. But the Great Author of nature has varied the machinery by which this nutritive fluid is formed and distributed, gradually proceeding from the most simple to the most complex structure; in which he seems to have seen it fit to invert the process observable in the systems of sensation and respiration, where the ascent is from the most complex, to the most simple structure. In the lowest members of the animal creation, the blood seems the portion they imbibe of the fluid medium in which they reside, which when chylified, distributes new molecules to all parts of their frame[350]. In others, as in insects, it is formed by the chyle that transpires through the intestinal canal into the general cavity of the body, where it receives oxygen from the air-vessels, and is fitted for nutrition[351]. In these animals it is accompanied by a long dorsal vessel, the first step towards a heart, which alternately contracts and dilates with an irregular systole and diastole, but appears to have no vascular system connected with it, though in their preparatory states it has an extra-vascular circulation which ceases in the perfect insect. Again: in others, as the Tubicoles, Annelida, &c., a real circulation has been discovered; that is to say, a system of veins and arteries, but unaccompanied by a muscular heart[352]. In the Arachnida and Branchiopod Crustacea the long dorsal vessel is also found; but in these it is connected with an arterial and venous system, which receives, distributes, and returns the blood[353]. It has therefore now become a true heart, and there is a regular circulation; and in the Decapod Crustacea the dorsal vessel is contracted into an oval form, and placed nearly in the centre of the trunk[354]. In the great majority of invertebrate animals the blood is white, but in the Annelida, to which Class the common dew-worm belongs, a curious anomaly takes place—for it is red[355]. Thus a gradual ascent is made to the circulating system of the vertebrate and red-blooded animals. In all, however, the blood is the principal instrument of nutrition and accretion; and is on that account properly so denominated, though not connected with a circulating system.
Having given you this general outline of the means by which the blood is distributed in the different Classes of animals, I shall now confine myself to the case of insects and Arachnida, beginning with the former.
I. If you examine attentively the back of any smooth caterpillar with a transparent skin, you will perceive in that part an evident pulsation, as though a fluid were pushed at regular intervals towards the head, along a narrow tube which seems to run the whole length of the body. Accurate dissections have proved that this appearance is real, that there is actually present in the back of most insects, placed immediately under the skin and furnished with numerous air-vessels, a longitudinal vessel[356] originating in the head near the mouth[357], running parallel with the alimentary canal nearly to the anus, containing a fluid which is propelled in regular pulsations of from 20 to 100 per minute, more or less as the weather is colder or warmer[358], causing a sensible alternate systole and diastole from the anal extremity towards the head. In the Cossus these pulses were observed by Lyonet to begin in the eleventh segment, from which they passed from segment to segment, till they arrived at the fourth, where they terminated[359]. This vessel is what Malpighi, who first discovered it, termed a heart, or rather series of hearts[360]; but which Reaumur, who injected it, regarded as a simple artery without striking contractions[361]: but to steer clear of any hypothesis, I shall merely call it the dorsal vessel (Pseudo-cardia). When carefully taken out of the body it is found to be a membranous tube, appearing to be closed at each end[362], in many larvæ of equal diameter every where, but in perfect insects usually widest at the anal extremity[363], and attenuated into a very slender filament towards the head. In some insects, however, as in the larva of the chamæleon-fly (Stratyomis Chamæleon), it is attenuated at both ends, and in the Ephemera is alternately constricted and dilated as Malpighi describes that of the silkworm[364], a dilated portion belonging to each segment[365]. In the Cossus, and probably others, after the third segment, it is furnished with nine pair, the three posterior pair being the largest, of triangular transverse bundles of muscular fibres, which Lyonet denominates its wings[366], the action of which produces its systole and diastole, and their propagation from the tail towards the head[367]. Under the last pair of these wings it is strengthened by a large number of circular muscular fibres[368]. I have stated it as appearing to be closed at each extremity, because Cuvier and most writers have so regarded it, and probably it is so closed in the perfect insect; but from Lyonet's words it should seem that, in the larva of the Cossus, he considered it as open and expanded at its anterior end[369]. He seems also to suspect, that, by means of what he calls the frontal ganglions, a fluid is derived from the dorsal vessel to the spinal marrow. He likewise describes a large nerve as passing through it and becoming recurrent[370]. Carus, as we shall soon see, has also proved that this tube is not closed in larvæ.
The fluid which this vessel contains is very abundant; in the animal it appears colourless and transparent like water, but when collected in drops it becomes more or less yellow, and even orange[371]. Examined under the microscope it appears filled with a prodigious number of transparent globules, of incredible minuteness[372]. When mixed with water, which it does readily, its globules lose all their transparency, and coagulate into small clammy masses. After evaporation it becomes hard, and cracks like gum, as blood does also. This gummy substance is so abundant, that the fluid contained in the dorsal vessel of the caterpillar of the Cossus yields a mass of it of the size of a grey pea[373].
From the situation of this dorsal vessel, which is precisely the same with that of the heart in Arachnida and the Branchiopod Crustacea, and from the systole and diastole which keep its fluid contents in constant motion, who can wonder that the physiologists who first discovered it, reasoning analogically, maintained that it was a true heart? But modern comparative anatomists, and those of the highest name, from the absence of a vascular system for a circulation, have contended that it is not a true heart, but an organ appropriated to other purposes: a third hypothesis, and intermediate between these two, has very recently been promulgated, that the organ in question, namely, is a real heart, and in the preparatory states of insects, the centre of a real circulation, which, in the imago state, ceases with the full development of the wings; but that this circulation is extravascular, or without peculiar vessels analogous to veins and arteries.
I shall now enlarge a little upon each of these hypotheses, beginning with the first or original one.
No one will deny that the argument from analogy is strongly in favour of this: I need not therefore dwell upon it, but proceed to others. Swammerdam, to whose exactness in observing, and scrupulous accuracy, every reader of his immortal work will bear testimony, expressly asserts that he has seen vessels issuing from the dorsal vessel in the silkworm, and even succeeded in injecting them with a coloured fluid[374]. Now it seems extremely improbable that so practised and expert an anatomist should have been deceived, especially upon a point which would naturally excite his most earnest and undivided attention. Without this recorded experiment, perhaps, it might be thought, though this was very unlikely, that he had mistaken bronchiæ for veins and arteries: but how could they have been injected from the supposed heart? Another great physiologist, Reaumur, in the caterpillar of the saw-fly of the rose (Hylotoma Rosæ) observed, besides the dorsal vessel, a ventral one of similar form, in which also was a pulsation, but slower than that of the other. This he supposes may be the principal trunk of the veins[375]. Bonnet thought he discovered a similar vessel in a large caterpillar, but with all his attention could perceive no motion in it[376]. Reaumur also fancied he perceived in the grub of Musca vomitoria, in which he in vain looked for the dorsal vessel, a fleshy part which exhibited alternate pulsations; and when with a pair of scissors he made a lateral incision in the insect, amongst other parts that came out, there was one that had movements of contraction and dilatation for several minutes,—this experiment was repeated with the same result upon several grubs[377]. De Geer, whose love of truth and accuracy no one will call in question, saw the appearance of blood-vessels in the leg of the larva of a Phryganea L. (as Lyonet did in those of a flea[378]); and in the transparent thigh of Ornithomia avicularia he discovered a pulse like that of an artery[379]. Baker, whose only object was to record what he saw, speaks of the current of the blood being remarkably visible in the legs of some small bugs[380]: what he meant by that term is uncertain, but they could not be spiders, which he had just distinguished. This author has likewise seen a green fluid passing through the vessels of the wings of grasshoppers[381]; and M. Chabrier is of opinion that insects possess the power of propelling a fluid into the nervures of their wings and withdrawing it at pleasure, as they are elevated or depressed[382]; but this last fact may be independent of a circulation.