Experiments on Animals. Stephen Paget
the blood could not well, because of the interposing valves, be sent by the veins to the limbs, it should be sent by the arteries, and return through the veins, whose valves did not oppose its course that way."
But between this observation, which "invited him to imagine" a theory, and his final proofs of the circulation, lay a host of difficulties; and it is certain, from his own account of his work, that experiments on animals were of the utmost help to him in leading him "out of the labyrinth."
III.—After Harvey
1. The Capillaries
The capillary vessels were not known in Harvey's time: the capillamenta of Cæsalpinus were not the capillaries, but the νευ̂ρα (Greek: neura) of Aristotle. It was believed that the blood, between the smallest arteries and the smallest veins, made its way through "blind porosities" in the tissues, as water percolates through earth or through a sponge. The first account of the capillaries is in two letters (De Pulmonibus, 1661) from Malpighi, professor of medicine at Bologna, to Borelli, professor of mathematics at Pisa. In his first letter, Malpighi writes that he has tried in vain, by injecting the dead body, to discover how the blood passes from the arteries into the veins:—
"This enigma hitherto distracts my mind, though for its solution I have made many and many attempts, all in vain, with air and various coloured fluids. Having injected ink with a syringe into the pulmonary artery, I have again and again seen it escape (become extravasated into the tissues) at several points. The same thing happens with an injection of mercury. These experiments do not give us the natural pathway of the blood."
But, in his second letter, he describes how he has examined, with a microscope of two lenses, the lung and the mesentery of a frog, and has seen the capillaries, and the blood in them:—
"Such is the divarication of these little vessels, coming off from the vein and the artery, that the order in which the vessel ramifies is no longer preserved, but it looks like a network woven from the offshoots of both vessels."
He was able, in a dead frog, to see the capillaries; and then, in a living frog, to see the blood moving in them. But, in spite of this work, it took nearly half a century before Harvey's teaching was believed by all men—Tantum consuetudo apud omnes valet.
2. The Blood-pressure
Harvey had seen the facts of blood-pressure—the great quantity of blood passing through, and the swiftness of its passage—but he had not measured it. Keill's experiments on the blood-pressure (1718) were inexact, and of no value; and the first exact measurements were made by Stephen Hales, who was rector of Farringdon, Hampshire, and minister of Teddington, Middlesex; a Doctor of Divinity, and a Fellow of the Royal Society. His experiments, in their width and diversity, were not surpassed even by those of John Hunter, and were extended far over physiology, vegetable physiology, organic and inorganic chemistry, and physics; they ranged from the invention of a sea-gauge to the study of solvents for the stone, and he seems to have experimented on every force in Nature. The titles of his two volumes of Statical Essays (1726–1733) show the great extent of his non-clerical work:—
Volume I. Statical Essays, containing Vegetable Statics, or an Account of some Statical Experiments on the Sap in Vegetables, being an Essay towards a Natural History of Vegetation; also, a Specimen of an Attempt to Analyse the Air, by a great Variety of Chymio-Statical Experiments.
Volume II. Statical Essays, containing Hæmostatics, or an Account of some Hydraulic and Hydrostatical Experiments made on the Blood and Blood-vessels of Animals; also, an Account of some Experiments on Stones in the Kidneys and Bladder, with an Enquiry into the Nature of those anomalous Concretions.
"We can never want matter for new experiments," he says in his preface. "We are as yet got little further than to the surface of things: we must be content, in this our infant state of knowledge, while we know in part only, to imitate children, who, for want of better skill and abilities, and of more proper materials, amuse themselves with slight buildings. The farther advances we make in the knowledge of Nature, the more probable and the nearer to truth will our conjectures approach: so that succeeding generations, who shall have the benefit and advantage both of their own observations and those of preceding generations, may then make considerable advances, when many shall run to and fro, and knowledge shall be increased."
His account of his plan of measuring the blood-pressure, and of one of many experiments that he made on it, is as follows:—
"Finding but little satisfaction in what had been attempted on this subject by Borellus and others, I endeavoured, about twenty-five years since, by proper experiments, to find what was the real force of the blood in the crural arteries of dogs, and about six years afterwards I repeated the like experiments on two horses, and a fallow doe; but did not then pursue the matter any further, being discouraged by the disagreeableness of anatomical dissections. But having of late years found by experience the advantage of making use of the statical way of investigation, not only in our researches into the nature of vegetables, but also in the chymical analysis of the air, I was induced to hope for some success, if the same method of enquiry were applied to animal bodies. …
"Having laid open the left crural artery (of a mare), I inserted into it a brass pipe whose bore was ⅙ of an inch in diameter; and to that, by means of another brass pipe which was fitly adapted to it, I fixed a glass tube of nearly the same diameter, which was 9 feet in length; then, untying the ligature on the artery, the blood rose in the tube 8 feet 3 inches perpendicular above the level of the left ventricle of the heart, but it did not attain to its full height at once: it rushed up gradually at each pulse 12, 8, 6, 4, 2, and sometimes 1 inch. When it was at its full height, it would rise and fall at and after each pulse 2, 3, or 4 inches, and sometimes it would fall 12 or 14 inches, and have there for a time the same vibrations up and down, at and after each pulse, as it had when it was at its full height, to which it would rise again, after forty or fifty pulses."
3. The Collateral Circulation
After Hales, came John Hunter, who was five years old when the Statical Essays were published. His experiments on the blood were mostly concerned with its properties, not with its course; but one great experiment must be noted here that puts him in line with Harvey, Malpighi, and Hales. He got from it his knowledge of the collateral circulation; he learned how the obstruction of an artery is followed by enlargement of the vessels in its neighbourhood, so that the parts beyond the obstruction do not suffer from want of blood: and the facts of collateral circulation were fresh in his mind when, a few months later, he conceived and performed his operation for aneurysm (December 1785). The "old operation" gave him no help here; and "Anel's operation" was but a single instance, and no sure guide for Hunter, because Anel's patient had a different sort of aneurysm. Hunter knew that the collateral circulation could be trusted to nourish the limb, if the femoral artery were ligatured in "Hunter's canal" for the cure of popliteal aneurysm; and he got this knowledge from the experiment that he had made on one of the deer in Richmond Park, to see the influence of ligature of the carotid artery on the growth of the antler. The following account of this experiment was given by Sir Richard Owen, who had it from Mr. Clift, Hunter's devoted pupil and friend:—
"In the month of July, when the bucks' antlers were half-grown, he caused one of them to be caught and thrown; and, knowing the arterial supply to the hot 'velvet,' as the keepers call it, Hunter cut down upon and tied the external carotid; upon which, laying his hand upon the antler, he found that the pulsations of the arterial channels stopped, and the surface soon grew cold. The buck was released, and Hunter speculated on the result—whether the antler, arrested at mid-growth, would be shed like the full-grown one, or be longer retained. A week or so afterward he drove down again to the park, and caused the buck to be caught and thrown. The wound was healed about the ligature; but on laying his hand on the antler, he found to his surprise that the warmth had returned, and the channels of supply to the velvety formative covering were again pulsating. His first impression was that his operation had been defective. To test this, he had the buck killed and sent to Leicester Square. The arterial system was injected. Hunter found that the external carotid had been duly tied. But certain small branches, coming off on