Books: The Harvard Classics Volume 38
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ON THE MOTION OF THE HEART AND BLOOD IN ANIMALS
CHAPTER I
THE AUTHOR'S MOTIVES FOR WRITING
When I first gave my mind to vivisections, as a means of
discovering the motions and uses of the heart, and sought to
discover these from actual inspection, and not from the writings
of others, I found the task so truly arduous, so full of
difficulties, that I was almost tempted to think, with
Fracastorius, that the motion of the heart was only to be
comprehended by God. For I could neither rightly perceive at
first when the systole and when the diastole took place, nor when
and where dilatation and contraction occurred, by reason of the
rapidity of the motion, which in many animals is accomplished in
the twinkling of an eye, coming and going like a flash of
lightning; so that the systole presented itself to me now from
this point, now from that; the diastole the same; and then
everything was reversed, the motions occurring, as it seemed,
variously and confusedly together. My mind was therefore greatly
unsettled nor did I know what I should myself conclude, nor what
believe from others. I was not surprised that Andreas Laurentius
should have written that the motion of the heart was as
perplexing as the flux and reflux of Euripus had appeared to
Aristotle.
At length, by using greater and daily diligence and
investigation, making frequent inspection of many and various
animals, and collating numerous observations, I thought that I
had attained to the truth, that I should extricate myself and
escape from this labyrinth, and that I had discovered what I so
much desired, both the motion and the use of the heart and
arteries. From that time I have not hesitated to expose my views
upon these subjects, not only in private to my friends, but also
in public, in my anatomical lectures, after the manner of the
Academy of old.
These views as usual, pleased some more, others less; some chid
and calumniated me, and laid it to me as a crime that I had dared
to depart from the precepts and opinions of all anatomists;
others desired further explanations of the novelties, which they
said were both worthy of consideration, and might perchance be
found of signal use. At length, yielding to the requests of my
friends, that all might be made participators in my labors, and
partly moved by the envy of others, who, receiving my views with
uncandid minds and understanding them indifferently, have essayed
to traduce me publicly, I have moved to commit these things to
the press, in order that all may be enabled to form an opinion
both of me and my labours. This step I take all the more
willingly, seeing that Hieronymus Fabricius of Aquapendente,
although he has accurately and learnedly delineated almost every
one of the several parts of animals in a special work, has left
the heart alone untouched. Finally, if any use or benefit to this
department of the republic of letters should accrue from my
labours, it will, perhaps, be allowed that I have not lived idly,
and as the old man in the comedy says:
For never yet hath any one attained
To such perfection, but that time, and place,
And use, have brought addition to his knowledge;
Or made correction, or admonished him,
That he was ignorant of much which he
Had thought he knew; or led him to reject
What he had once esteemed of highest price.
So will it, perchance, be found with reference to the heart at
this time; or others, at least, starting hence, with the way
pointed out to them, advancing under the guidance of a happier
genius, may make occasion to proceed more fortunately, and to
inquire more accurately.
CHAPTER II
ON THE MOTIONS OF THE HEART AS SEEN IN THE DISSECTION OF LIVING
ANIMALS
In the first place, then, when the chest of a living animal is
laid open and the capsule that immediately surrounds the heart is
slit up or removed, the organ is seen now to move, now to be at
rest; there is a time when it moves, and a time when it is
motionless.
These things are more obvious in the colder animals, such as
toads, frogs, serpents, small fishes, crabs, shrimps, snails, and
shell-fish. They also become more distinct in warm-blooded
animals, such as the dog and hog, if they be attentively noted
when the heart begins to flag, to move more slowly, and, as it
were, to die: the movements then become slower and rarer, the
pauses longer, by which it is made much more easy to perceive and
unravel what the motions really are, and how they are performed.
In the pause, as in death, the heart is soft, flaccid, exhausted,
lying, as it were, at rest.
In the motion, and interval in which this is accomplished, three
principal circumstances are to be noted:
1. That the heart is erected, and rises upwards to a point, so
that at this time it strikes against the breast and the pulse is
felt externally.
2. That it is everywhere contracted, but more especially towards
the sides so that it looks narrower, relatively longer, more
drawn together. The heart of an eel taken out of the body of the
animal and placed upon the table or the hand, shows these
particulars; but the same things are manifest in the hearts of
all small fishes and of those colder animals where the organ is
more conical or elongated.
3. The heart being grasped in the hand, is felt to become harder
during its action. Now this hardness proceeds from tension,
precisely as when the forearm is grasped, its tendons are
perceived to become tense and resilient when the fingers are
moved.
4. It may further be observed in fishes, and the colder blooded
animals, such as frogs, serpents, etc., that the heart, when it
moves, becomes of a paler color, when quiescent of a deeper
blood-red color.
From these particulars it appears evident to me that the motion
of the heart consists in a certain universal tension--both
contraction in the line of its fibres, and constriction in every
sense. It becomes erect, hard, and of diminished size during its
action; the motion is plainly of the same nature as that of the
muscles when they contract in the line of their sinews and
fibres; for the muscles, when in action, acquire vigor and
tenseness, and from soft become hard, prominent, and thickened:
and in the same manner the heart.
We are therefore authorized to conclude that the heart, at the
moment of its action, is at once constricted on all sides,
rendered thicker in its parietes and smaller in its ventricles,
and so made apt to project or expel its charge of blood. This,
indeed, is made sufficiently manifest by the preceding fourth
observation in which we have seen that the heart, by squeezing
out the blood that it contains, becomes paler, and then when it
sinks into repose and the ventricle is filled anew with blood,
that the deeper crimson colour returns. But no one need remain in
doubt of the fact, for if the ventricle be pierced the blood will
be seen to be forcibly projected outwards upon each motion or
pulsation when the heart is tense.
These things, therefore, happen together or at the same instant:
the tension of the heart, the pulse of its apex, which is felt
externally by its striking against the chest, the thickening of
its parietes, and the forcible expulsion of the blood it contains
by the constriction of its ventricles.
Hence the very opposite of the opinions commonly received appears
to be true; inasmuch as it is generally believed that when the
heart strikes the breast and the pulse is felt without, the heart
is dilated in its ventricles and is filled with blood; but the
contrary of this is the fact, and the heart, when it contracts
(and the impulse of the apex is conveyed through the chest wall),
is emptied. Whence the motion which is generally regarded as the
diastole of the heart, is in truth its systole. And in like
manner the intrinsic motion of the heart is not the diastole but
the systole; neither is it in the diastole that the heart grows
firm and tense, but in the systole, for then only, when tense, is
it moved and made vigorous.
Neither is it by any means to be allowed that the heart only
moves in the lines of its straight fibres, although the great
Vesalius giving this notion countenance, quotes a bundle of
osiers bound in a pyramidal heap in illustration; meaning, that
as the apex is approached to the base, so are the sides made to
bulge out in the fashion of arches, the cavities to dilate, the
ventricles to acquire the form of a cupping-glass and so to suck
in the blood. But the true effect of every one of its fibres is
to constringe the heart at the same time they render it tense;
and this rather with the effect of thickening and amplifying the
walls and substance of the organ than enlarging its ventricles.
And, again, as the fibres run from the apex to the base, and draw
the apex towards the base, they do not tend to make the walls of
the heart bulge out in circles, but rather the contrary; inasmuch
as every fibre that is circularly disposed, tends to become
straight when it contracts; and is distended laterally and
thickened, as in the case of muscular fibres in general, when
they contract, that is, when they are shortened longitudinally,
as we see them in the bellies of the muscles of the body at
large. To all this let it be added, that not only are the
ventricles contracted in virtue of the direction and condensation
of their walls, but farther, that those fibres, or bands, styled
nerves by Aristotle, which are so conspicuous in the ventricles
of the larger animals, and contain all the straight fibres (the
parietes of the heart containing only circular ones), when they
contract simultaneously by an admirable adjustment all the
internal surfaces are drawn together as if with cords, and so is
the charge of blood expelled with force.
Neither is it true, as vulgarly believed, that the heart by any
dilatation or motion of its own, has the power of drawing the
blood into the ventricles; for when it acts and becomes tense,
the blood is expelled; when it relaxes and sinks together it
receives the blood in the manner and wise which will by-and-by be
explained.
CHAPTER III
OF THE MOTIONS OF THE ARTERIES, AS SEEN IN THE DISSECTION OF
LIVING ANIMALS
In connexion with the motions of the heart these things are
further to be observed having reference to the motions and pulses
of the arteries.
1. At the moment the heart contracts, and when the breast is
struck, when in short the organ is in its state of systole, the
arteries are dilated, yield a pulse, and are in the state of
diastole. In like manner, when the right ventricle contracts and
propels its charge of blood, the pulmonary artery is distended at
the same time with the other arteries of the body.
2. When the left ventricle ceases to act, to contract, to
pulsate, the pulse in the arteries also ceases; further, when
this ventricle contracts languidly, the pulse in the arteries is
scarcely perceptible. In like manner, the pulse in the right
ventricle failing, the pulse in the pulmonary artery ceases also.
3. Further, when an artery is divided or punctured, the blood is
seen to be forcibly propelled from the wound the moment the left
ventricle contracts; and, again, when the pulmonary artery is
wounded, the blood will be seen spouting forth with violence at
the instant when the right ventricle contracts.
So also in fishes, if the vessel which leads from the heart to
the gills be divided, at the moment when the heart becomes tense
and contracted, at the same moment does the blood flow with force
from the divided vessel.
In the same way, when we see the blood in arteriotomy projected
now to a greater, now to a less distance, and that the greater
jet corresponds to the diastole of the artery and to the time
when the heart contracts and strikes the ribs, and is in its
state of systole, we understand that the blood is expelled by the
same movement.
From these facts it is manifest, in opposition to commonly
received opinions, that the diastole of the arteries corresponds
with the time of the heart's systole; and that the arteries are
filled and distended by the blood forced into them by the
contraction of the ventricles; the arteries, therefore, are
distended, because they are filled like sacs or bladders, and are
not filled because they expand like bellows. It is in virtue of
one and the same cause, therefore, that all the arteries of the
body pulsate, viz., the contraction of the left ventricle; in the
same way as the pulmonary artery pulsates by the contraction of
the right ventricle.
Finally, that the pulses of the arteries are due to the impulses
of the blood from the left ventricle, may be illustrated by
blowing into a glove, when the whole of the fingers will be found
to become distended at one and the same time, and in their
tension to bear some resemblance to the pulse. For in the ratio
of the tension is the pulse of the heart, fuller, stronger, and
more frequent as that acts more vigorously, still preserving the
rhythm and volume, and order of the heart's contractions. Nor is
it to be expected that because of the motion of the blood, the
time at which the contraction of the heart takes place, and that
at which the pulse in an artery (especially a distant one) is
felt, shall be otherwise than simultaneous: it is here the same
as in blowing up a glove or bladder; for in a plenum (as in a
drum, a long piece of timber, etc.) the stroke and the motion
occur at both extremities at the same time. Aristotle, [Footnote:
De Anim., iii, cap. 9.] too, has said, "the blood of all animals
palpitates within their veins (meaning the arteries), and by the
pulse is sent everywhere simultaneously." And further, [Footnote:
De Respir., cap. 20] "thus do all the veins pulsate together and
by successive strokes, because they all depend upon the heart;
and, as it is always in motion, so are they likewise always
moving together, but by successive movements." It is well to
observe with Galen, in this place, that the old philosophers
called the arteries veins. I happened upon one occasion to have a
particular case under my care, which plainly satisfied me of the
truth: A certain person was affected with a large pulsating
tumour on the right side of the neck, called an aneurism, just at
that part where the artery descends into the axilla, produced by
an erosion of the artery itself, and daily increasing in size;
this tumour was visibly distended as it received the charge of
blood brought to it by the artery, with each stroke of the heart;
the connexion of parts was obvious when the body of the patient
came to be opened after his death. The pulse in the corresponding
arm was small, in consequence of the greater portion of the blood
being diverted into the tumour and so intercepted.
Whence it appears that whenever the motion of the blood through
the arteries is impeded, whether it be by compression or
infarction, or interception, there do the remote divisions of the
arteries beat less forcibly, seeing that the pulse of the
arteries is nothing more than the impulse or shock of the blood
in these vessels.
CHAPTER IV
OF THE MOTION OF THE HEART AND ITS AURICLES, AS SEEN IN THE
BODIES OF LIVING ANIMALS
Besides the motions already spoken of, we have still to consider
those that appertain to the auricles.
Caspar Bauhin and John Riolan, [Footnote: i Bauhin, lib. ii. cap.
II. Riolan. lib. viii, cap. I.] most learned men and skilful
anatomists, inform us that from their observations, that if we
carefully watch the movements of the heart in the vivisection of
an animal, we shall perceive four motions distinct in time and in
place, two of which are proper to the auricles, two to the
ventricles. With all deference to such authority I say that there
are four motions distinct in point of place, but not of time; for
the two auricles move together, and so also do the two
ventricles, in such wise that though the places be four, the
times are only two. And this occurs in the following manner:
There are, as it were, two motions going on together: one of the
auricles, another of the ventricles; these by no means taking
place simultaneously, but the motion of the auricles preceding,
that of the heart following; the motion appearing to begin from
the auricles and to extend to the ventricles. When all things are
becoming languid, and the heart is dying, as also in fishes and
the colder blooded animals there is a short pause between these
two motions, so that the heart aroused, as it were, appears to
respond to the motion, now more quickly, now more tardily; and at
length, when near to death, it ceases to respond by its proper
motion, but seems, as it were, to nod the head, and is so
slightly moved that it appears rather to give signs of motion to
the pulsating auricles than actually to move. The heart,
therefore, ceases to pulsate sooner than the auricles, so that
the auricles have been said to outlive it, the left ventricle
ceasing to pulsate first of all; then its auricle, next the right
ventricle; and, finally, all the other parts being at rest and
dead, as Galen long since observed, the right auricle still
continues to beat; life, therefore, appears to linger longest in
the right auricle. Whilst the heart is gradually dying, it is
sometimes seen to reply, after two or three contractions of the
auricles, roused as it were to action, and making a single
pulsation, slowly, unwillingly, and with an effort.
But this especially is to be noted, that after the heart has
ceased to beat, the auricles however still contracting, a finger
placed upon the ventricles perceives the several pulsations of
the auricles, precisely in the same way and for the same reason,
as we have said, that the pulses of the ventricles are felt in
the arteries, to wit, the distension produced by the jet of
blood. And if at this time, the auricles alone pulsating, the
point of the heart be cut off with a pair of scissors, you will
perceive the blood flowing out upon each contraction of the
auricles. Whence it is manifest that the blood enters the
ventricles, not by any attraction or dilatation of the heart, but
by being thrown into them by the pulses of the auricles.
And here I would observe, that whenever I speak of pulsations as
occurring in the auricles or ventricles, I mean contractions:
first the auricles contract, and then and subsequently the heart
itself contracts. When the auricles contract they are seen to
become whiter, especially where they contain but little blood;
but they are filled as magazines or reservoirs of the blood,
which is tending spontaneously and, by its motion in the veins,
under pressure towards the centre; the whiteness indicated is
most conspicuous towards the extremities or edges of the auricles
at the time of their contractions.
In fishes and frogs, and other animals which have hearts with but
a single ventricle, and for an auricle have a kind of bladder
much distended with blood, at the base of the organ, you may very
plainly perceive this bladder contracting first, and the
contraction of the heart or ventricle following afterwards.
But I think it right to describe what I have observed of an
opposite character: the heart of an eel, of several fishes, and
even of some (of the higher) animals taken out of the body,
pulsates without auricles; nay, if it be cut in pieces the
several parts may still be seen contracting and relaxing; so that
in these creatures the body of the heart may be seen pulsating
and palpitating, after the cessation of all motion in the
auricle. But is not this perchance peculiar to animals more
tenacious of life, whose radical moisture is more glutinous, or
fat and sluggish, and less readily soluble? The same faculty
indeed appears in the flesh of eels, which even when skinned and
embowelled, and cut into pieces, are still seen to move.
Experimenting with a pigeon upon one occasion, after the heart
had wholly ceased to pulsate, and the auricles too had become
motionless, I kept my finger wetted with saliva and warm for a
short time upon the heart, and observed that under the influence
of this fomentation it recovered new strength and life, so that
both ventricles and auricles pulsated, contracting and relaxing
alternately, recalled as it were from death to life.
Besides this, however, I have occasionally observed, after the
heart and even its right auricle had ceased pulsating,--when it
was in articulo mortis in short,--that an obscure motion, an
undulation or palpitation, remained in the blood itself, which
was contained in the right auricle, this being apparent so long
as it was imbued with heat and spirit. And, indeed, a
circumstance of the same kind is extremely manifest in the course
of the generation of animals, as may be seen in the course of the
first seven days of the incubation of the chick: A drop of blood
makes its appearance which palpitates, as Aristotle had already
observed; from this, when the growth is further advanced and the
chick is fashioned, the auricles of the heart are formed, which
pulsating henceforth give constant signs of life. When at length,
and after the lapse of a few days, the outline of the body begins
to be distinguished, then is the ventricular part of the heart
also produced, but it continues for a time white and apparently
bloodless, like the rest of the animal; neither does it pulsate
or give signs of motion. I have seen a similar condition of the
heart in the human foetus about the beginning of the third month,
the heart then being whitish and bloodless, although its auricles
contained a considerable quantity of purple blood. In the same
way in the egg, when the chick was formed and had increased in
size, the heart too increased and acquired ventricles, which then
began to receive and to transmit blood.
And this leads me to remark that he who inquires very
particularly into this matter will not conclude that the heart,
as a whole, is the primum vivens, ultimum moriens,--the first
part to live, the last to die,--but rather its auricles, or the
part which corresponds to the auricles in serpents, fishes, etc.,
which both lives before the heart and dies after it.
Nay, has not the blood itself or spirit an obscure palpitation
inherent in it, which it has even appeared to me to retain after
death? and it seems very questionable whether or not we are to
say that life begins with the palpitation or beating of the
heart. The seminal fluid of all animals--the prolific spirit, as
Aristotle observed, leaves their body with a bound and like a
living thing; and nature in death, as Aristotle [Footnote: De
Motu Animal., cap. 8.] further remarks, retracing her steps,
reverts to where she had set out, and returns at the end of her
course to the goal whence she had started. As animal generation
proceeds from that which is not animal, entity from nonentity,
so, by a retrograde course, entity, by corruption, is resolved
into nonentity, whence that in animals, which was last created,
fails first and that which was first, fails last.
I have also observed that almost all animals have truly a heart,
not the larger creatures only, and those that have red blood, but
the smaller, and pale-blooded ones also, such as slugs, snails,
scallops, shrimps, crabs, crayfish, and many others; nay, even in
wasps, hornets, and flies, I have, with the aid of a magnifying
glass, and at the upper part of what is called the tail, both
seen the heart pulsating myself, and shown it to many others.
But in the pale-blooded tribes the heart pulsates sluggishly and
deliberately, contracting slowly as in animals that are moribund,
a fact that may readily be seen in the snail, whose heart will be
found at the bottom of that orifice in the right side of the body
which is seen to be opened and shut in the course of respiration,
and whence saliva is discharged, the incision being made in the
upper aspect of the body, near the part which corresponds to the
liver.
This, however, is to be observed: that in winter and the colder
season, exsanguine animals, such as the snail, show no pulsation;
they seem rather to live after the manner of vegetables, or of
those other productions which are therefore designated plant-
animals.
It is also to be noted that all animals which have a heart have
also auricles, or something analogous to auricles; and, further,
that whenever the heart has a double ventricle, there are always
two auricles present, but not otherwise. If you turn to the
production of the chick in ovo, however, you will find at first
no more a vesicle or auricle, or pulsating drop of blood; it is
only by and by, when the development has made some progress, that
the heart is fashioned; even so in certain animals not destined
to attain to the highest perfection in their organization, such
as bees, wasps, snails, shrimps, crayfish, etc., we only find a
certain pulsating vesicle, like a sort of red or white
palpitating point, as the beginning or principle of their life.
We have a small shrimp in these countries, which is taken in the
Thames and in the sea, the whole of whose body is transparent;
this creature, placed in a little water, has frequently afforded
myself and particular friends an opportunity of observing the
motions of the heart with the greatest distinctness, the external
parts of the body presenting no obstacle to our view, but the
heart being perceived as though it had been seen through a
window.
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