The body consists of a soft granular protoplasmic substance, the middle

of which is occupied by a large oval mass called the “nucleus;” while,

at its hinder end, is a “contractile vacuole,” conspicuous by its

regular rhythmic appearances and disappearances. Obviously, although

the _Colpoda_ is not a monad, it differs from one only in subordinate

details. Moreover, under certain conditions, it becomes quiescent,

incloses itself in a delicate case or _cyst_, and then divides into

two, four, or more portions, which are eventually set free and swim

about as active _Colpodæ_.

But this creature is an unmistakable animal, and full-sized _Colpodæ_

may be fed as easily as one feeds chickens. It is only needful to

diffuse very finely ground carmine through the water in which they

live, and, in a very short time, the bodies of the _Colpodæ_ are

stuffed with the deeply-coloured granules of the pigment.

And if this were not sufficient evidence of the animality of _Colpoda_,

there comes the fact that it is even more similar to another well-known

animalcule, _Paramæcium_, than it is to a monad. But _Paramæcium_ is

so huge a creature compared with those hitherto discussed—it reaches

1/120 of an inch or more in length—that there is no difficulty in

making out its organisation in detail; and in proving that it is not

only an animal, but that it is an animal which possesses a somewhat

complicated organisation. For example, the surface layer of its body is

different in structure from the deeper parts. There are two contractile

vacuoles, from each of which radiates a system of vessel-like canals;

and not only is there a conical depression continuous with a tube,

which serve as mouth and gullet, but the food ingested takes a definite

course, and refuse is rejected from a definite region. Nothing is

easier than to feed these animals, and to watch the particles of indigo

or carmine accumulate at the lower end of the gullet. From this they

gradually project, surrounded by a ball of water, which at length

passes with a jerk, oddly simulating a gulp, into the pulpy central

substance of the body, there to circulate up one side and down the

other, until its contents are digested and assimilated. Nevertheless,

this complex animal multiplies by division, as the monad does, and,

like the monad, undergoes conjugation. It stands in the same relation

to _Heteromita_ on the animal side, as _Coleochæte_ does on the plant

side. Start from either, and such an insensible series of gradations

leads to the monad that it is impossible to say at any stage of the

progress—here the line between the animal and the plant must be drawn.

There is reason to think that certain organisms which pass through a

monad stage of existence, such as the _Myxomycetes_, are, at one time

of their lives, dependent upon external sources for their protein

matter, or are animals; and, at another period, manufacture it, or are

plants. And seeing that the whole progress of modern investigation is

in favour of the doctrine of continuity, it is a fair and probable

speculation—though only a speculation—that, as there are some plants

which can manufacture protein out of such apparently intractable

mineral matters as carbonic acid, water, nitrate of ammonia, metallic

and earthy salts; while others need to be supplied with their carbon

and nitrogen in the somewhat less raw form of tartrate of ammonia and

allied compounds; so there may be yet others, as is possibly the case

with the true parasitic plants, which can only manage to put together

materials still better prepared—still more nearly approximated to

protein—until we arrive at such organisms as the _Psorospermiæ_ and the

_Panhistophyton_, which are as much animal as vegetable in structure,

but are animal in their dependence on other organisms for their food.

The singular circumstance observed by Meyer, that the _Torula_ of

yeast, though an indubitable plant, still flourishes most vigorously

when supplied with the complex nitrogenous substance, pepsin; the

probability that the _Peronospora_ is nourished directly by the

protoplasm of the potato-plant; and the wonderful facts which have

recently been brought to light respecting insectivorous plants, all

favour this view; and tend to the conclusion that the difference

between animal and plant is one of degree rather than of kind; and that

the problem whether, in a given case, an organism is an animal or a

plant, may be essentially insoluble.

VIII.

ON CERTAIN ERRORS RESPECTING THE STRUCTURE OF THE HEART

ATTRIBUTED TO ARISTOTLE.

In all the commentaries upon the “Historia Animalium” which I have met

with, Aristotle’s express and repeated statement, that the heart of

man and the largest animals contains only three cavities, is noted as

a remarkable error. Even Cuvier, who had a great advantage over most

of the commentators in his familiarity with the subject of Aristotle’s

description, and whose habitual caution and moderation seem to desert

him when the opportunity of panegyrising the philosopher presents

itself, is betrayed into something like a sneer on this topic.

“Du reste il n’attribue à cet organe que trois cavités,

erreur qui prouve au moins qu’il en avait regardé la

structure.”[36]

To which remark, what follows will, I think, justify the reply, that it

“prouve au moins” that Cuvier had not given ordinary attention, to say

nothing of the careful study which they deserve, to sundry passages in

the first and the third books of the “Historia” which I proceed to lay

before the reader.

For convenience of reference these passages are marked _A_, _B_, _C_,

etc.[37]

Book i. 17.—(_A_) “The heart has three cavities, it lies

above the lung on the division of the windpipe, and has

a fatty and thick membrane where it is united with the

great vein and the aorta. It lies upon the aorta, with

its point down the chest, in all animals that have a

chest. In all, alike in those that have a chest and in

those that have none, the foremost part of it is the

apex. This is often overlooked through the turning upside

down of the dissection. The rounded end of the heart is

uppermost, the pointed end of it is largely fleshy and

thick, and in its cavities there are tendons. In other

animals which have a chest the heart lies in the middle

of the chest; in men, more to the left side, between the

nipples, a little inclined to the left nipple in the

upper part of the chest. The heart is not large, and its

general form is not elongated but rounded, except that

the apex is produced into a point.

(_B_) “It has, as already stated, three cavities, the

largest of them is on the right, the smallest on the

left, the middle-sized one in the middle; they have all,

also the two small ones, passages (τετρημένας) towards

the lung, very evidently as respects one of the cavities.

In the region of the union [with the great vein and the

aorta] the largest cavity is connected with the largest

vein (near which is the mesentery); the middle cavity

with the aorta.

(_C_) “Canals (πόροι) from the heart pass to the lung and

divide in the same fashion as the windpipe does, closely

accompanying those from the windpipe through the whole

lung. The canals from the heart are uppermost.

(_D_) “No canal is common [to the branches of the

windpipe and those of the vein] (οὐδεὶς δ’ ἐστὶ κοινὸς

πόρος) but through those parts of them which are in

contact (τὴν σύναψιν) the air passes in and they [the

πόροι] carry it to the heart.

(_E_) “One of the canals leads to the right cavity, the

other to the left.

(_F_) “Of all the viscera, the heart alone contains blood

[in itself]. The lung contains blood, not in itself but

in the veins, the heart in itself; for in each of the

cavities there is blood; the thinnest is in the middle

cavity.”

Book iii. 3.—(_G_) “Two veins lie in the thorax alongside

the spine, on its inner face; the larger more forwards,

the smaller behind; the larger more to the right, the

smaller, which some call _aorta_ (on account of the

tendinous part of it seen in dead bodies), to the left.

These take their origin from the heart; they pass entire,

preserving the nature of veins, through the other viscera

that they reach; while the heart is rather a part of

them, and more especially of the anterior and larger one,

which is continued into veins above and below, while

between these is the heart.

(_H_) “All hearts contain cavities, but, in those of very

small animals, the largest [cavity] is hardly visible,

those of middling size have another, and the biggest all

three.

(_I_) “The point of the heart is directed forwards, as

was mentioned at first; the largest cavity to the right

and upper side of it, the smallest to the left, and the

middle-sized one between these; both of these are much

smaller than the largest.

(_K_) “They are all connected by passages (συντέτρηνται)

with the lung, but, on account of the smallness of the

canals, this is obscure except in one.

(_L_) “The great vein proceeds from the largest cavity

which lies upwards and to the right; next through the

hollow middle part (διὰ τοῦ κοίλου τοῦ μέσου) it becomes

vein again, this cavity being a part of the vein in which

the blood stagnates.

(_M_) “The aorta [proceeds from] the middle [cavity], but

not in the same way, for it is connected [with the middle

cavity] by a much more narrow tube (σύριγγα).

(_N_) “The [great] vein extends through the heart,

towards the aorta from the heart.

(_O_) “The great vein is membranous like skin, the aorta

narrower than it and very tendinous, and as it extends

towards the head and the lower parts it becomes narrow

and altogether tendinous.

(_P_) “In the first place, a part of the great vein

extends upwards from the heart towards the lung and

the attachment of the aorta, the vein being large and

undivided. It divides into two parts, the one to the

lung, the other to the spine and the lowest vertebra of

the neck.

(_Q_) “The vein which extends to the lung first divides

into two parts for the two halves of it and then extends

alongside each tube (σύριγγα) and each passage (τρῆμα),

the larger beside the larger and the smaller beside the

smaller, so that no part [of the lung] can be found

from which a passage (τρῆμα) and a vein are absent.

The terminations are invisible on account of their

minuteness, but the whole lung appears full of blood. The

canals from the vein lie above the tubes given off from

the windpipe.”

The key to the whole of the foregoing description of the heart lies

in the passages (_G_) and (_L_). They prove that Aristotle, like

Galen, five hundred years afterwards, and like the great majority

of the old Greek anatomists, did not reckon what we call the right

auricle as a constituent of the heart at all, but as a hollow part, or

dilatation, of the “great vein.” Aristotle is careful to state that his

observations were conducted on suffocated animals; and if any one will

lay open the thorax of a dog or a rabbit, which has been killed with

chloroform, in such a manner as to avoid wounding any important vessel,

he will at once see why Aristotle adopted this view.

For, as the subjoined figure (p. 185) shows, the vena cava inferior

(_b_), the right auricle (_R.a._), and the vena cava superior and

innominate vein (_V.I._) distended with blood seem to form one

continuous column, to which the heart is attached as a sort of

appendage. This column is, as Aristotle says, vein above (_a_) and vein

below (_b_), the upper and the lower divisions being connected διὰ

τοῦ κοίλου τοῦ μέσου—or by means of the intervening cavity or chamber

(_R.a._)—which is that which we call the right auricle.

[Illustration:

A dog having been killed by chloroform, enough of the

right wall of the thorax was removed, without any notable

bleeding, to expose the thoracic viscera. A carefully

measured outline sketch of the parts _in situ_ was then

made, and on dissection, twenty-four hours afterwards,

the necessary anatomical details were added. The woodcut

is a faithfully reduced copy of the drawing thus

constructed; and it represents the relations of the heart

and great vessels as Aristotle saw them in a suffocated

animal.

All but the inner lobe of the right lung has been

removed; as well as the right half of the pericardium and

the right walls of the right auricle and ventricle. It

must be remembered that the thin transparent pericardial

membrane appears nothing like so distinct in nature.

_a.b._, Aristotle’s “great vein”; _V.I._, right vena

innominata and vena cava superior; _b_, the inferior

vena cava; _R.a._, the “hollow middle” part of the great

vein or the right auricle; _R.v′_, the prolongation of

the cavity of the right ventricle _R.v_ towards the

pulmonary artery; _tr_, one of the tricuspid valves;

_Pc_, the pericardium; _I.sv_, superior intercostal

vein; _Az_, vena azygos; _P.A._, right pulmonary artery;

_Br_, right bronchus; _L_, inner lobe of the right lung;

_Œ_, œsophagus; _Ao_, descending aorta; _H_, liver, in

section, with hepatic vein, vena portæ, and gall-bladder,

_gb_, separated by the diaphragm, also seen in section,

from the thoracic cavity.]

But when, from the four cavities of the heart recognised by us moderns,

one is excluded, there remain three—which is just what Aristotle says.

The solution of the difficulty is, in fact, as absurdly simple as that

presented by the egg of Columbus; and any error there may be, is not

to be put down to Aristotle, but to that inability to comprehend that

the same fact may be accurately described in different ways, which

is the special characteristic of the commentatorial mind. That the

three cavities mentioned by Aristotle are just those which remain if

the right auricle is omitted, is plain enough from what is said in

(_B_), (_C_), (_E_), (_I_), and (_L_). For, in a suffocated animal,

the “right cavity” which is directly connected with the great vein,

and is obviously the right ventricle, being distended with blood, will

look much larger than the middle cavity, which, since it gives rise

to the aorta, can only be the left ventricle. And this, again, will

appear larger than the thin and collapsed left auricle, which must

be Aristotle’s left cavity, inasmuch as this cavity is said to be

connected by πόροι with the lung. The reason why Aristotle considered

the left auricle to be a part of the heart, while he merged the right

auricle in the great vein, is, obviously, the small relative size of

the venous trunks and their sharper demarcation from the auricle.

Galen, however, perhaps more consistently, regarded the left auricle

also as a mere part of the “arteria venosa.” The canal which leads

from the right cavity of the heart to the lung (or, as Aristotle

puts it (_E_), from the lung to the heart) is, without doubt, the

pulmonary artery. But it may be said that, in this case, Aristotle

contradicts himself, inasmuch as in (_P_) and (_Q_) a vessel, which

is obviously the pulmonary artery, is described as a branch of the

great vein. However, this difficulty also disappears, if we reflect

that, in Aristotle’s way of looking at the matter, the line of

demarcation between the great vein and the heart coincides with the

right auriculo-ventricular aperture; and that, inasmuch as the conical

prolongation of the right ventricle which leads to the pulmonary

artery (_R.v′_ in the Figure), lies close in front of the auricle,

its base may very easily (as the figure shows) be regarded as part of

the general opening of the great vein into the right ventricle. In

fact, it is clear that Aristotle, having failed to notice the valves

of the heart, did not distinguish the part of the right ventricle

from which the pulmonary artery arises (_R.v′_) from the proper trunk

of the artery on the one hand, and from the right auricle (_R.a_) on

the other. Thus the root, as we may call it, of the pulmonary artery

and the right auricle, taken together, are spoken of as the “part of

the great vein which extends upwards” (_P_); and, as the vena azygos

(_Az_) was one branch of this, so the “vein to the lung” was regarded

as another branch of it. But the latter branch, being given off close

to the connection of the great vein with the ventricle, was also

counted as one of the two πόροι by which the “heart” (that is to say

the right ventricle, the left ventricle, and the left auricle of our

nomenclature) communicates with the lung.

The only other difficulty that I observe is connected with (_K_). If

Aristotle intended by this to affirm that the middle cavity (the left

ventricle), like the other two, is directly connected with the lung by

a πόρος, he would be in error. But he has excluded this interpretation

of his words by (_E_), in which the number and relations of the

canals, the existence of which he admits, are distinctly defined. I

can only imagine then, that, so far as this passage applies to the

left ventricle, it merely refers to the indirect communication of that

cavity with the vessels of the lungs, through the left auricle.

On this evidence I submit that there is no escape from the conclusion

that, instead of having committed a gross blunder, Aristotle has given

a description of the heart which, so far as it goes, is remarkably

accurate. He is in error only in regard to the differences which he

imagines to exist between large and small hearts (_H_).

Cuvier (who has been followed by other commentators) ascribes another

error to Aristotle:—

“Aristote suppose que la trachée-artère se prolonge

jusqu’au cœur, et semble croire, en conséquence, que

l’air y pénétre (_l. c._ p. 152).”

Upon what foundation Cuvier rested the first of these two assertions,

I am at a loss to divine. As a matter of fact, it will appear from the

following excerpts that Aristotle gives an account of the structure of

the lungs which is almost as good as that of the heart, and that it

contains nothing about any prolongation of the windpipe to the heart.

“Within the neck lie what is called the œsophagus (so

named on account of its length and its narrowness) and

the windpipe (ἀρτηρία). The position of the windpipe in

all animals that have one, is in front of the œsophagus.

All animals which possess a lung have a windpipe. The

windpipe is of a cartilaginous nature and is exsanguine,

but is surrounded by many little veins....

“It goes downwards towards the middle of the lung, and

then divides for each of the halves of the lung. In all

animals that possess one, the lung is divided into two

parts; but, in those which bring forth their young alive,

the separation is not equally well marked, least of all

in man.

“In oviparous animals, such as birds, and in quadrupeds

which are oviparous, the one half of the lung is widely

separated from the other; so that it appears as if they

had two lungs. And from being single, the windpipe

becomes (divided into) two, which extend to each half of

the lung. It is fastened to the great vein, and to what

is called the aorta. When the windpipe is blown up, the

air passes into the hollow parts of the lung. In these,

are cartilaginous tubes (διαφύσεις) which unite at an

angle; from the tubes passages (τρήματα) traverse the

whole of the lung; they are continually given off, the

smaller from the larger.” (Book i. 16.)

That Aristotle should speak of the lung as a single organ divided into

two halves, and should say that the division is least marked in man,

is puzzling at first; but the statement becomes intelligible, if we

reflect upon the close union of the bronchi, the pulmonary vessels and

the mediastinal walls of the pleuræ, in mammals;[38] and it is quite

true that the lungs are much more obviously distinct from one another

in birds.

Aubert and Wimmer translate the last paragraph of the passage just

cited as follows:—

“Diese haben aber knorpelige Scheidewände, welche unter

spitzen Winkeln zusammentreten, und aus ihnen führen

Oeffnungen durch die ganze Lunge, indem sie sich in immer

kleineren verzweigen.”

But I cannot think that by διαφύσεις and τρήματα, in this passage,

Aristotle meant either “partitions” or openings in the ordinary

sense of the latter word. For, in Book iii. Cap. 3, in describing

the distribution of the “vein which goes to the lung” (the pulmonary

artery), he says that it

“extends alongside each tube (σύριγγα) and each passage

(τρῆμα), the larger beside the larger, and the smaller

beside the smaller; so that no part (of the lung) can be

found from which a passage (τρῆμα) and a vein are absent.”

Moreover, in Book i. 17, he says—

“Canals (πόροι) from the heart pass to the lung and

divide in the same fashion as the windpipe does, closely

accompanying those from the windpipe through the whole

lung.”

And again in Book i. 17—

“It (the lung) is entirely spongy, and alongside of each

tube (σύριγγα) run canals (πόροι) from the great vein.”

On comparing the last three statements with the facts of the case, it

is plain that by σύριγγες, or tubes, Aristotle means the bronchi and so

many of their larger divisions as obviously contain cartilages; and

that by διαφύσεις χονδρώδεις he denotes the same things; and, if this

be so, then the τρήματα must be the smaller bronchial canals, in which

the cartilages disappear.

This view of the structure of the lung is perfectly correct so far

as it extends; and, bearing it in mind, we shall be in a position to

understand what Aristotle thought about the passage of air from the

lungs into the heart. In every part of the lung, he says, in effect,

there is an air tube which is derived from the trachea, and other tubes

which are derived from the πόροι which connect the lung with the heart

(_suprà_, _C_). Their applied walls constitute the thin “synapses” (τὴν

σύναψιν) through which the air passes out of the air tubes into the

πόροι, or blood-vessels, by transudation or diffusion; for there is no

community between the cavities of the air tubes and cavities of the

canals; that is to say, no opening from one into the other (_suprà_,

_D_).

On the words “κοινὸς πόρος” Aubert and Wimmer remark (_l. c._ p. 239),

“Da A. die Ansicht hat die Lungenluft würde dem Herzen zugeführt,

so postulirt er statt vieler kleiner Verbindungen einen grossen

Verbindungsgang zwischen Lunge und Herz.”

But does Aristotle make this assumption? The only evidence so far as

I know in favour of the affirmative answer to this question is the

following passage:—

“Συνήρτεται δὲ καὶ ἡ καρδία τῇ ἀρτηριᾷ πιμελώδεσι

καὶ χονδρώδεσι καὶ ἰνώδεσι δεσμοῖς· ᾗ δὲ συνήρτεται,

κοῖλόν ἐστιν. φυσωμένης δὲ τῆς ἀρτηρίας μὲν ἐνίοις ἐν οὐ

κατάδηλον ποιεῖ, ἐν δὲ τοῖς μείζοσι τῶν ζῴων δῆλον ὅτι

εἰσέρχεται τὸ πνεῦμα εἰς αὐτὴν” (i. cap. 16).

“The heart and the windpipe are connected by fatty and

cartilaginous and fibrous bands; where they are connected

it is hollow. Blowing into the windpipe does not show

clearly in some animals, but in the larger animals it is

clear that the air goes into it.”

Aubert and Wimmer give a somewhat different rendering of this passage:—

“Auch das Herz hängt mit der Luftröhre durch fettreiche,

knorpelige und faserige Bänder zusammen; und da, wo sie

zusammenhängen, ist eine Höhlung. Beim Aufblasen der

Lunge wird es bei manchen Thieren nicht wahrnehmbar, bei

den grösseren aber ist es offenbar, dass die Luft in das

Herz gelangt.”

The sense here turns upon the signification which is to be ascribed to

εἰς αὐτὴν. But if these words refer to the heart, then Aristotle has

distinctly pointed out the road which the air, in his opinion, takes,

namely, through the “synapses” (_D_); and there is no reason that I

can discover to believe that he “postulated” any other and more direct

communication.

With respect to the meaning of κοῖλόν ἐστιν, Aubert and Wimmer observe:—

“Dies scheint wohl die kurze Lungenvene zu sein.

Schneider bezieht dies auf die Vorkammern, allein diese

werden unten als Höhlen des Herzens beschrieben.”

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