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LECTURE III. CIRCULATION OF THE BLOOD

Two kinds of motion may be distinguished in the animal economy; the one voluntary, or under the command of the will, which takes place at certain intervals, but may be stopped at pleasure. The other kind of motion is called involuntary, as not depending on the will, but going on constantly, without interruption, both when we sleep and when we wake.

Of the first kind is the motion of the limbs, of which I have already spoken in general terms; the object of which is, to change the situation of the animal, and carry it where the will directs.

Among the involuntary motions, the most remarkable is the circulation of the blood, which I shall proceed to consider in this lecture.

There is one motion, however, which claims a middle place between the voluntary and involuntary; I mean respiration. This action is so far under the command of the will, that it may be suspended, increased, or diminished in strength and frequency: but we can only suspend it for a very short time; and it goes on regularly during sleep, and in general, even when we are awake, without the intervention of the will; its continuation being always necessary, as we have already seen, to support life.

The motion of the fluids in the living body is regulated by very different laws, from those which govern the motion of ordinary fluids, that depend upon their gravity and fluidity: these last have a general centre of gravitation to which they incessantly tend. Their motion is from above downwards, when not prevented by any obstacles; and when they meet with obstruction, they either stop till the obstacle is removed, or escape where they find the least resistance. When they have reached the lowest situations, they remain at rest, unless acted upon by some internal impulse, which again puts them in motion.

But the motion of the fluids in an animal body, is less uniform, constant, and regular; it takes place upwards as well as downwards, and overcomes numerous obstacles; it carries the blood from the interior parts of the body to the surface, and from the surface back again to the internal parts; it forces it from the left side of the body to the right, and with such rapidity that not a particle of the fluid remains an instant in the same place.

The principal organ concerned in the circulation of the blood, is the heart; which is a hollow muscle, of a conical figure, with two cavities, called ventricles; this organ is situated in the thorax or chest; its apex or point is inclined downwards and to the left side, where it is received in a cavity of the left lobe of the lungs.

At the basis of the heart on each side are situated two cavities, called auricles, to receive the blood; and these contracting, force the blood into the ventricles, which are two cavities in the heart, separated from each other by a strong muscular partition. The cavity which is situated on the right side of the heart, is called the right ventricle, and that on the left the left ventricle. From the right ventricle of the heart issues a large artery, called the pulmonary artery, which goes to the lungs, and is there divided and subdivided into a vast number of branches, the extremities of which are too small to be visible. These ultimate ramifications unite again into larger branches; these again into branches still larger, and so continually, till at last they form four tubes, called the pulmonary veins, which are inserted into the left auricle of the heart,

From the left ventricle of the heart there issues another large artery, called the aorta, which, in its passage, sends off branches to the heart, arms, legs, head, and every other part of the body. These branches, in the course of their progress, are divided and subdivided into innumerable minute ramifications, the last of which are invisible. These small ramifications unite again into branches continually larger and larger, till they form two great tubes, called the venae cavae; which large veins are inserted into the right auricle of the heart; where a vein, termed the coronary vein of the heart, which returns the blood from the heart itself, also terminates.

From what has been said, it will be evident, that strictly speaking, there are only two arteries and seven veins in the body; one pulmonary artery, which carries the blood from the right ventricle of the heart to the lungs, and four pulmonary veins, which bring it back again; then the aorta or large artery, which carries the blood from the left ventricle of the heart to all parts of the body; the two venae cavae, and the coronary vein of the heart, which bring it back again.

At the beginning of both arteries, where they leave the heart, are placed valves, which allow the blood to flow freely from the heart into the arteries, but which prevent its return to the heart. There are likewise valves between the auricles and ventricles, which permit the blood to flow from the former into the latter, but prevent its return into the auricles. The veins are likewise furnished with valves, which allow the blood to flow from their minute branches along the larger toward the heart, but prevent its returning to these minute branches.

The blood being brought back from all parts of the body into the right auricle of the heart, distends this cavity, and thus causes it to contract; this auricle, by contracting, forces the blood into the right ventricle; this muscular cavity being distended and irritated by the blood, contracts, and propels the blood through the pulmonary artery into the lungs: from hence it is brought back by the pulmonary veins, to the left auricle of the heart, by whose contraction it is forced into the left ventricle. The contraction of this ventricle propels the blood, with great force, into the aorta, through the innumerable ramifications of which, it is carried to every part of the body, and brought back by veins, which accompany these arterial ramifications, and form the venae cavae, which conduct the blood into the right auricle of the heart, from whence it is again sent into the right ventricle, which sends it through the pulmonary artery, to the lungs; the pulmonary veins bring it back again to the heart, from whence it is propelled through the aorta, to all parts of the body: thus running a perpetual round, called the circulation of the blood.

Thus then we see, that the circulation consists of two circles or stages, one through the lungs, which may be called the pulmonary, or lesser circle, and the other through all parts of the body, which may be termed the aortal, or greater circle.

That the blood circulates in this manner, is evident, from the valves placed at the origin of the arteries, and in the large branches of the veins, which prevent the return of the blood to the heart, in any other manner than that I have described. This is likewise evident, in the common operation of blood letting: when the arm is tied, the vein swells below the ligature, instead of above, and we do not make the opening above the ligature, or on the side next the heart. If the vein were opened above the ligature, it would not bleed. For it only swells next the hand, which shows that the blood does not flow into the vein downwards from the heart, but upwards from the hand.

If the ligature be too tight, the blood will not flow through the opening in the vein. The reason of this, is, that the artery is compressed, in this case, as well as the vein; and as the veins derive their blood from the arteries, it follows that if the blood's motion be obstructed in the latter, none can flow from them into the former: when we wish to open an artery, the orifice must be made above the ligature.

Another proof of the circulation being performed in this manner, is derived from microscopic observations, on the transparent parts of animals, in which the blood can be seen to move towards the extremities, along the arteries, and return by the veins.

The blood, however, does not flow out of the heart into the arteries in a continued stream, but by jets, or pulses; when the ventricles are filled with blood from the auricles, this blood stimulates them, and thereby causes them to contract; by such contraction, they force the blood, which they contain, into the arteries; this contraction is called the systole of the heart. As soon as they have finished their contraction, they relax, till they are again filled with blood from the auricles, and this state of relaxation of the heart, is called the diastole.

This causes the pulsation or beating of the heart. The arteries must, of course, have a similar pulsation, the blood being driven into them only by starts; and accordingly we find it in the artery of the wrist; this beating we call the pulse; the like may also be observed in the arteries of the temples, and other parts of the body. The veins, however, have no pulsation, for the blood flowing on, in an uninterrupted course, from smaller tubes to wider, its pulse becomes entirely destroyed.

The different cavities of the heart do not contract at the same time; but the two auricles contract together, the ventricles being at that time in a state of relaxation; these ventricles then contract together, while the auricles become relaxed.

Both the arteries and veins may be compared to a tree, whose trunk is divided into large branches; these are subdivided into smaller, the smaller again into others still smaller; and we may observe, likewise, that the sum of the capacities of the branches, which arise from any trunk, is always greater than the capacity of the trunk.

The minutest branches of the arteries, being reflected, become veins, or else they enter veins that are already formed, by anastomosis, as it is called; the small veins continually receiving others, become, like a river, gradually larger, till they form the venae cavae, which conduct the blood to the heart.

Anatomical injections prove, that the last branches of the arteries terminate in the beginning of veins; but it is the opinion of many celebrated physiologists, that the arteries carry the blood to the different parts of the body to nourish them, and that the veins commence by open mouths, which absorb or suck up what is superfluous, and return it back to the heart.

From what has been said, it must be evident that there is a considerable resemblance between the circulation of the blood in the animal body, and the circulation of the aqueous fluid on the surface of the globe. In the latter case the water is raised from the ocean, by the heat of the sun, and poured down upon the dry land, in minute drops, for the nourishment and economy of its different parts. What is superfluous is collected into little rills; these meeting with others, form brooks; the union of which produce rivers, that conduct the water to its original source, from which it is again circulated.

In the same manner, the blood is sent by the heart to different parts of the body, for the nourishment and economy of its different parts; what is superfluous is brought back by veins, which, continually uniting, form those large trunks, which convey the vital fluid to the heart.

The blood does not circulate, however, in the manner which I have mentioned, in all parts of the body; for that which is carried by arteries to the viscera, serving for digestion, such as the stomach, bowels, mesentery, omentum, and spleen, is collected by small veins which unite into a large trunk called the vena portarum; this vein enters the liver, and is subdivided in it like an artery, distributing through the liver a great quantity of blood, from which the bile is secreted: and, having served this purpose, the blood is collected by small veins; these unite and form the hepatic vein, which pours the blood into the vena cava, to be conducted to the heart.

The reason of this deviation, is probably, to diminish the velocity of the blood in the liver, for the secretion of the bile; which could not have been effected by means of an artery.

The force which impels the blood, is, first, the contraction of the heart, which propels the blood into the arteries with great velocity; but this is not the only force concerned in keeping up the circulation; this is evident, from the diminished heat, and weakened pulse, in a paralytic limb, which ought not to take place, if the blood were propelled merely by the action of the heart.

The arteries are possessed of an elastic and muscular power, by means of which they contract when they are distended or stimulated. It is however by the muscular power alone, that they assist in propelling the blood; for the elasticity of their coats can serve no other purpose than preserving the mean diameter of the vessel. If we suppose the arteries to be dilated by the blood, poured into them by the heart, they will, by their contraction, as elastic tubes, undoubtedly propel the blood: but supposing them to be perfectly elastic, the force of the heart will be just as much diminished in dilating them as the force of the blood is increased by their contraction. We are not however acquainted with any substance perfectly elastic, or which restores itself with a force equal to that with which it was distended: hence the elastic power of the arteries will subtract from, instead of adding to, the power of the heart. It is evident, therefore, that it must be by the muscular power of the arteries, which causes them to contract like the heart, that they propel the blood.

That such is the case, appears from the muscular structure of the arteries observed by anatomists; as also from the effects of mechanical irritation of their coats, which causes them to contract; this is likewise evident from the inflammation produced by the application of stimulating substances to particular parts; for instance, cantharides and mustard. It appears likewise, from the secretion in some parts being preternaturally increased, while the motion of the general mass of the blood continues unaltered.

The contraction of the arteries always propels the blood towards the extreme parts of the body: this must necessarily happen, because the valves at the origin of the arteries prevent its return to the heart, it must therefore move in the direction in which it finds least resistance.

If it were not for this muscular power of the arteries, the force of the heart would not alone be able to propel the blood to the extreme parts of the body, and overcome the different kinds of resistance it has to encounter. Among the causes that lessen the velocity of the blood, may be mentioned the increasing area of the artery; for it was before observed, that the sum of the cavities of the branches from any trunk exceeded the cavity of the trunk: and from the principles of hydrostatics, the velocities of fluids, propelled by the same force, in tubes of different diameters, are inversely as the squares of the diameters, so that in a tube of double the diameter, the velocity will only be one fourth; in one of the triple, only one ninth: and since the arteries may be looked upon as conical, it is evident that the velocity of the blood must be diminished from this cause.

The curvilinear course of the arteries likewise gives considerable resistance; for at every bending the blood loses part of its momentum against the sides; and this loss is evidently proportioned to the magnitude of the angle, at which the branch goes off. Convolutions are frequently made, in order to diminish the force of the blood in particular organs; this is especially the case with the carotid artery before it enters the brain.

The angles which the ramifications of the arteries make, are greater or more obtuse nearer the heart, and more acute as the distance increases; by which means the velocity of the blood is rendered more equal in different parts.

The anastomosing or union of different branches of arteries, likewise retards the velocity of the blood, the particles of which, from different vessels, impinging, disturb each other's motion, and produce a compound force, in which there is always a loss of velocity: and it is evident, from the composition of forces, that this loss must be proportioned to the obliquity of the angle at which the vessels unite.

The adhesion of the blood to the sides of the vessels, likewise causes a loss of velocity in the minuter branches, which may be owing to a chemical affinity: the viscidity or imperfect fluidity of the blood is another retarding cause. All these causes united, would render it impossible for the heart to propel the blood with the velocity with which it moves in the very minute branches of the arteries, if these arteries were not endowed with a living muscular power like the heart, by which they contract and propel their contents.

In the veins, the motion of the blood is occasioned partly by the vis a tergo, and partly by the contraction of the neighbouring muscles, which press upon the veins; and these veins being furnished with valves, the return of the blood towards the arteries is prevented; it must therefore move towards the heart.

That the contraction of the muscles of the body tends very much to promote the circulation of the blood, is evident, from the increase of the circulation from exercise, and likewise from the languid motion of the blood in sedentary persons, and those given to indolence. Hence we may account for the different diseases to which such persons are subject, and know how to apply the proper remedies. Hence likewise, we see the reason why rest is so absolutely necessary in acute and inflammatory diseases, where the momentum of the blood is already too great.

It has been doubted by anatomists, whether the veins were possessed with muscular power; but this seems now to be confirmed. Haller found the vena cava near the heart to contract on the application of stimulants, though he could see no muscular fibres; these, however, have been discovered by succeeding anatomists.

The magnitude of the veins is always greater than that of the corresponding arteries; hence the velocity of the blood must be less in the veins; and hence likewise we may account for their want of pulsation; for the action of the heart upon the arteries is at first very great; but as we recede from the heart, this effect becomes less perceptible; the arterial tube increases both in size and muscularity, in proportion to its distance from the source of circulation. The powers of the heart are spent in overcoming the different resistances which I have noticed, before the blood enters the veins; hence the blood will flow uniformly in these last.

The blood is subject in the veins to retarding causes, similar to those which operate in the arteries, but perhaps not in an equal degree; for the flexures are less frequent in the veins than in the arteries. As the capacity of the arterial tube increases with its distance from the heart, the velocity, from this cause, as has already been observed, is continually diminished; but a contrary effect takes place in the veins; for the different branches uniting, form trunks, whose capacities are smaller than the sums of the capacities of the branches, hence the velocity of the blood in the veins will increase as it approaches the heart.

Another retarding cause may be mentioned, namely, gravity, which acts more on the venous than the arterial system. The effects of gravity on the veins may be exemplified, by a ring being pulled off the finger with ease when the hand is elevated; also by the swellings of the feet that occur in relaxed habits, which swellings increase towards night, and subside in the morning, after the body has been in a horizontal posture for some hours.

In weak persons, the frequency of the pulse is increased by an erect posture, which may probably depend on gravity; as we know, from the observations of Macdonald and others, that an erect posture will make a difference of 15 or 20 beats in a minute. The experiments alluded to, were made by gently raising a person fastened to a board, where there being no muscular exertion, respiration would not be increased; so that the whole effect was probably owing to gravity accelerating the column of arterial blood.

The inverted posture produces a still more remarkable effect in accelerating the pulse, than the erect, for it sometimes causes it to beat 10 or 12 times more in the former case than in the latter.

While we are on this subject, it may not be improper to take notice of the effects of swinging on the circulation, which have been found by Dr. Carmichael Smyth, and others, to diminish the strength and velocity to such a degree, as to bring on fainting. These effects have never been satisfactorily accounted for; but they would seem to admit of an easy explanation on mechanical principles: they are undoubtedly owing, at least in a great measure, to the centrifugal force acquired by the blood.

By a centrifugal force, I mean, the tendency which revolving bodies have to fly off from the centre, which arises from their tendency to move in a straight line, agreeably to the laws of motion. Hence a tumbler of water may be whirled in a circle vertically without spilling it; the centrifugal force pushing the water against the bottom of the tumbler. In the same manner when the human body is made to revolve vertically in the arch of a circle, this centrifugal force will propel the blood from the head and heart towards he extremities; hence the circulation of the blood will be weakened, and the energy of the brain diminished. The contrary, however, will take place on a horizontal swing, as I have frequently observed, both on myself and others; for the centrifugal force in this case will propel the blood from the extremities towards the head.

It has been already observed, that the pulsations of the artery which we feel at the wrist, are occasioned by its alternate dilatations and contractions, which vary according to the strength and regularity of the circulation, which is liable to be affected by the smallest changes in the state of health. Hence physicians make use of the pulse as a criterion whereby to judge of the health of the body. And we may observe that there are few more certain characteristics of the state of the body than the pulse; yet the conclusions that have been drawn from it have often been erroneous; and this has arisen from trusting to observation without the aid of reason.

That we may better understand the phenomena of the pulse, I shall lay down the following postulata. 1st. It is now generally believed, that every part of the arterial system is endowed with irritability, or a power of contracting on the application of a stimulus, and that the blood acting on this contractibility, if the term may be allowed, causes contraction; and that the alternate relaxation and contraction gives the phenomenon pulsation. 2d. The greater the action of the stimulus of the blood, the greater will be the contraction, that is, the nearer will the sides of the artery approach towards the axis. 3d. That the velocity with which a muscular fibre, in a state of debility, contracts, is at least equal to that with which a fibre in a state of strength contracts, is a fact generally allowed by physiologists.

We shall afterwards see, that a deficient action of stimulus on the vessels may arise, either directly from diminishing the quantity of blood contained in them, or indirectly, from the application of too great a stimulant power, which has diminished the capability of contracting inherent in the vessels.

From these postulata, it will be evident, that the greater the action of the arteries, that is, the more powerful their contraction, the longer will be the intervals between the pulsations.

For the velocity being at least equal in debility and in strength, the times between the pulsations will be proportioned to the approach of the sides of the artery towards its axis: but the approach of the sides towards the axis is greater when the arteries are in a state of vigour than when debilitated; consequently the intervals between the pulsations will be greater when the arteries are in a state of vigour than when debilitated.

Hence it is evident, that a frequency of pulse must generally indicate a diminished action or debility; while a moderate slowness indicates a vigorous or just action.

Hence likewise the opinion of increased action, which has been supposed to take place in fevers, because a frequent pulse was observed, must be false, because the frequency arises from a directly opposite state, and indicates a diminished action of the vascular system.

In a sound and adult man the frequency of the pulse is about seventy beats in a minute; and in an infant, within the first five or six months, the pulse is seldom less than one hundred and twenty, and diminishes in frequency as the child grows older. But though seventy beats in the minute may be taken as a general standard; yet in persons of irritable constitutions the frequency is greater than this, and many, who are in the prime of life, have the pulse only between fifty and sixty.

It is generally observed, that the pulse is slower in the morning, that it increases in frequency till noon, after dinner it again becomes slow, and in the evening its frequency returns, which increases till midnight.

These phenomena may be rationally explained on the principles just laid down. When we rise in the morning, the contractibility being abundant, the stimulus of the blood produces a greater effect, the pulse becomes slow, and the contractions strong; it becomes more frequent, however, till dinner time, from a diminished contractibility; after dinner, from the addition of the stimulus of food and chyle, it again decreases in frequency, and becomes slow till the evening, when its frequency returns, because the contractibility becomes exhausted: and this frequency continues till the vital power have been recruited by sleep.

By the same principles it is easy to explain the quickness of the pulse in infancy, its gradual decrease till maturity, its slowness and strength during the meridian of life, and the return of its frequency during the decline.

Having now described the phenomena of the circulation, it will be proper to examine the changes produced by this function on the blood; and, in the first place, it may be observed, that the blood which returns by the vena cava to the heart, is of a dark colour inclining to purple; while that which passes from the left ventricle into the arteries, is of a bright vermilion hue. The blood which is found in the pulmonary artery has the same dark purple colour with that in the vena cava, while that in the pulmonary vein resembles the aortal blood in its brightness. Hence it would appear, that the blood, during its passage through the lungs, has its colour changed from a dark purple to a bright vermilion, in which state it is brought by the pulmonary vein to the left auricle of the heart; this auricle, contracting, expels the blood into the corresponding ventricle, by whose action, and that of the arteries, it is distributed to all parts of the body. When it returns, however, by the veins, it is found to have lost its fine bright colour. It would appear, therefore, that the blood obtains its red colour during its passage through the lungs, and becomes deprived of it during its circulation through the rest of the body.

That the blood contains iron, may be proved by various experiments: if a quantity of blood be exposed to a red heat in a crucible, the greatest part will be volatilised and burnt; but a quantity of brown ashes will be left behind, which will be attracted by the magnet. If diluted sulphuric acid be poured on these ashes, a considerable portion of them will dissolve; if into this solution we drop tincture of galls, a black precipitate will take place, or if we use prussiate of potash, a precipitate of prussian blue will be formed. These facts prove, beyond doubt, that a quantity of iron exists in the blood.

I shall not now particularly inquire how it comes there; it may partly be taken into the blood along with the vegetable and animal food, which is received into the stomach; for the greatest part of the animal and vegetable substances, which we receive as food, contain a greater or less quantity of iron. Or it may be partly formed by the animal powers, as would appear from the following circumstance. The analysis of an egg, before incubation, affords not the least vestige of iron, but as soon as the chick exists, though it has been perfectly shut up from all external communication, if the egg be burnt, the ashes will be attracted by the magnet.

But, however we may suppose the blood to obtain its iron, it certainly does contain it; if the coagulable lymph and serum of the blood be carefully freed from the red particles, by repeated washing, the strictest analysis will not discover in either of them a particle of iron, while the red globules thus separated will be found to contain a considerable quantity of this metal.

That the red colour of the blood depends upon iron, appears likewise from the experiments of Menghini, which show, that the blood of persons who have been taking chalybeate medicines for some time, is much more florid that it is naturally; the same is agreeable to my own observation. A late analysis, by Fourcroy, has likewise proved, that the red colour of the blood resides in the iron; but, though the red colour of the blood may reside in the iron which it contains, we shall find that this colour is likewise connected with oxidation.

If the dark coloured blood, drawn from the veins, be put under a vessel containing oxygen gas, its surface will immediately become florid, while the bulk of the gas will be diminished. Mr. Hewson enclosed a portion of a vein between two ligatures, and injected into it a quantity of oxygen gas; the blood, which was before dark coloured, instantly assumed the hue of arterial blood. Thuvenal put a quantity of arterial blood under the receiver of an air pump; on exhausting the air it became of the dark colour of venous blood; on readmitting the air, it became again florid. He put it under a receiver filled with oxygen gas, and found the florid colour much increased.