VII.--From the first appearance of an individualized centre in the

vascular area of the human embryo, that centre (punctum saliens) and the

vessels immediately connected with it, undergo a phaseal metamorphosis,

till such time after birth as they assume their permanent character. In

each stage of metamorphosis, the embryo heart and vessels typify the

normal condition of the organ in one of the lower classes of animals.

The several species of the organ in these classes are parallel to the

various stages of change in the human organ. In its earliest condition,

the human heart presents the form of a simple canal, similar to that of

the lower Invertebrata, the veins being connected with its posterior

end, while from its anterior end a single artery emanates. The canal

next assumes a bent shape, and the vessels of both its ends become

thereby approximated. The canal now being folded upon itself in

heart-shape, next becomes constricted in situations, marking out the

future auricle and ventricle and arterial bulb, which still communicate

with each other. From the artery are given off on either side

symmetrically five branches (branchial arches), which arch laterally

from before, outwards and backwards, and unite in front of the

vertebrae, forming the future descending aorta. In this condition, the

human heart and vessels resemble the Piscean pipe. The next changes

which take place consist in the gradual subdivision, by means of septa,

of the auricle and ventricle respectively into two cavities. On the

separation of the single auricle into two, while the ventricle as yet

remains single, the heart presents that condition which is proper to the

Reptilian class. The interauricular and interventricular septa, by

gradual development from without inwards, at length meet and coalesce,

thereby dividing the two cavities into four--two auricles and two

ventricles--a condition proper to the Avian and Mammalian classes

generally. In the centre of the interauricular septum of the human

heart, an aperture (foramen ovale) is left as being necessary to the

foetal circulation. While the septa are being completed, the arterial

bulb also becomes divided by a partition formed in its interior in such

a manner as to adjust the two resulting arteries, the one in connexion

with the right, the other with the left ventricle. The right ventricular

artery (pulmonary aorta) so formed, has assigned to it the fifth

(posterior) opposite pair of arches, and of these the right one

remaining pervious to the point where it gives off the right pulmonary

branch, becomes obliterated beyond this point to that where it joins the

descending aorta, while the left arch remains pervious during foetal

life, as the ductus arteriosus still communicating with the descending

aorta, and giving off at its middle the left pulmonary branch. The left

ventricular artery (systemic aorta) is formed of the fourth arch of the

left side, while the opposite arch (fourth right) is altogether

obliterated. The third and second arches remain pervious on both sides,

afterwards to become the right and left brachio-cephalic arteries. The

first pair of arches, if not converted into the vertebral arteries, or

the thyroid axes, are altogether metamorphosed. By these changes the

heart and primary arteries assume the character in which they usually

present themselves at birth, and in all probability the primary veins

corresponded in form, number, and distribution with the arterial

vessels, and underwent, at the same time, a similar mode of

metamorphosis. One point in respect to the original symmetrical

character of the primary veins is demonstrable--namely, that in front of

the aortic branches the right and left brachio-cephalic veins, after

joining by a cross branch, descend separately on either side of the

heart, and enter (as two superior venae cavae) the right auricle by

distinct orifices. In some of the lower animals, this double condition

of the superior veins is constant, but in the human species the left

vein below the cross branch (left brachio-cephalic) becomes obliterated,

whilst the right vein (vena cava superior) receives the two

brachio-cephalic veins, and in this condition remains throughout life.

After birth, on the commencement of respiration, the foramen ovale of

the interauricular septum closes, and the ductus arteriosus becomes

impervious. This completes the stages of metamorphosis, and changes the

course of the simple foetal circulation to one of a more complex

order--viz., the systemic-pulmonary characteristic of the normal state

in the adult body.

VIII.--Such being the phases of metamorphosis of the primary (branchial)

arches which yield the vessels in their normal adult condition, we

obtain in this history an explanation of the signification not only of

such of their anomalies as are on record, but of such also as are

potential in the law of development; a few of them will suffice to

illustrate the meaning of the whole number:--lst, The interventricular

as well as the interauricular septum may be arrested in growth, leaving

an aperture in the centre of each; the former condition is natural to

the human foetus, the latter to the reptilian class, while both would be

abnormal in the human adult. 2nd. The heart may be cleft at its apex in

the situation of the interventricular septum--a condition natural to the

Dugong, A similar cleavage may divide the base of the heart in the

situation of the interauricular septum. 3rd. The partitioning of the

bulbus arteriosus may occur in such a manner as to assign to the two

aortae a relative position, the reverse of that which they normally

occupy--the pulmonary aorta springing from the left ventricle and the

systemic aorta arising from the right, and giving off from its arch the

primary branches in the usual order. [Footnote 1] 4th. As the two aortae

result from a division of the common primary vessel (bulbus arteriosus),

an arrest in the growth of the partition would leave them still as one

vessel, which (supposing the ventricular septum remained also

incomplete) would then arise from a single ventricle. 5th. The ductus

arteriosus may remain pervious, and while co-existing with the proper

aortic arch, two arches would then appear on the left side. 6th. The

systemic normal aortic arch may be obliterated as far up as the

innominate branch, and while the ductus arteriosus remains pervious, and

leading from the pulmonary artery to the descending part of the aortic

arch, this vessel would then present the appearance of a branch

ascending from the left side and giving off the brachio-cephalic

arteries. The right ventricular artery would then, through the medium of

the ductus arteriosus, supply both the lungs and the system. Such a

state of the vessels would require (in order that the circulation of a

mixed blood might be carried on) that the two ventricles freely

communicate. 7th. If the fourth arch of the right side remained pervious

opposite the proper aortic arch, there would exist two aortic arches

placed symmetrically, one on either side of the vertebral column, and,

joining below, would include in their circle the trachea and oesophagus.

8th. If the fifth arch of the right side remained pervious opposite the

open ductus arteriosus, both vessels would present a similar

arrangement, as two symmetrical ducti arteriosi co-existing with

symmetrical aortic arches. 9th. If the vessels appeared co-existing in

the two conditions last mentioned, they would represent four aortic

arches, two on either side of the vertebral column. 10th. If the fourth

right arch, instead of the fourth left (aorta), remained pervious, the

systemic aortic arch would then be turned to the right side of the

vertebral column, and have the trachea and oesophagus on its left. 11th.

When the bulbus arteriosus divides itself into three parts, the two

lateral parts, in becoming connected with the left ventricle, will

represent a double ascending systemic aorta, and having the pulmonary

artery passing between them to the lungs. 12th. When of the two original

superior venae cavae the right one instead of the left suffers

metamorphosis, the vena cava superior will then appear on the left side

of the normal aortic arch. [Footnote 2] Of these malformations, some are

rather frequently met with, others very seldom, and others cannot exist

compatible with life after birth. Those which involve a more or less

imperfect discharge of the blood-aerating functions of the lungs, are in

those degrees more or less fatal, and thus nature aborting as to the

fitness of her creation, cancels it.

[Footnote 1: This physiological truth has, I find, been applied by Dr.

R. Quain to the explanation of a numerous class of malformations

connected with the origins of the great vessels from the heart, and of

their primary branches. See The Lancet, vol. I. 1842.]

[Footnote 2: For an analysis of the occasional peculiarities of these

primary veins in the human subject, see an able and original monograph

in the Philosophical Transactions, Part 1., 1850, entitled, "On the

Development of the Great Anterior Veins in Man and Mammalia." By John

Marshall, F.R.C.S., &c. ]

IX.--The portal system of veins passing to the liver, and the hepatic

veins passing from this organ to join the inferior vena cava, exhibit in

respect to the median line of the body an example of a-symmetry, since

appearing on the right side, they have no counterparts on the left. As

the law of symmetry seems to prevail universally in the development of

organized beings, forasmuch as every lateral organ or part has its

counterpart, while every central organ is double or complete, in having

two similar sides, then the portal system, as being an exception to this

law, is as a natural note of interrogation questioning the signification

of that fact, and in the following observations, it appears to me, the

answer may be found. Every artery in the body has its companion vein or

veins. The inferior vena cava passes sidelong with the aorta in the

abdomen. Every branch of the aorta which ramifies upon the abdominal

parietes has its accompanying vein returning either to the vena cava or

the vena azygos, and entering either of these vessels at a point on the

same level as that at which itself arises. The renal vessels also have

this arrangement. But all the other veins of the abdominal viscera,

instead of entering the vena cava opposite their corresponding arteries,

unite into a single trunk (vena portae), which enters the liver. The

special purpose of this destination of the portal system is obvious, but

the function of a part gives no explanation of its form or relative

position, whether singular or otherwise. On viewing the vessels in

presence of the general law of symmetrical development, it occurs to me

that the portal and hepatic veins form one continuous system, which

taken in the totality, represents the companion veins of the arteries of

the abdominal viscera. The liver under this interpretation appears as a

gland developed midway upon these veins, and dismembering them into a

mesh of countless capillary vessels, (a condition necessary for all

processes of secretion,) for the special purpose of decarbonizing the

blood. In this great function the liver is an organ correlative or

compensative to the lungs, whose office is similar. The secretion of the

liver (bile) is fluidform; that of the lungs is aeriform. The bile being

necessary to the digestive process, the liver has a duct to convey that

product of its secretion to the intestines. The trachea is as it were

the duct of the lungs. In the liver, then, the portal and hepatic veins

being continuous as veins, the two systems, notwithstanding their

apparent distinctness, caused by the intervention of the hepatic

lobules, may be regarded as the veins corresponding with the arteries of

the coeliac axis, and the two mesenteric. The hepatic artery and the

hepatic veins evidently do not pair in the sense of afferent and

efferent, with respect to the liver, both these vessels having

destinations as different as those of the bronchial artery and the

pulmonary veins in the lungs. The bronchial artery is attended by its

vein proper, while the vein which corresponds to the hepatic artery

joins either the hepatic or portal veins traversing the liver, and in

this position escapes notice.[Footnote]

[Footnote: In instancing these facts, as serving under comparison to

explain how the hepatic vessels constitute no radical exception to the

law of symmetry which presides over the development and distribution of

the vascular system as a whole, I am led to inquire in what respect (if

in any) the liver as an organ forms an exception to this general law

either in shape, in function, or in relative position. While seeing that

every central organ is single and symmetrical by the union of two

absolutely similar sides, and that each lateral pair of organs is double

by the disunion of sides so similar to each other in all respects that

the description of either side serves for the other opposite, it has

long since seemed to me a reasonable inference that, since the liver on

the right has no counterpart as a liver on the left, and that, since the

spleen on the left has no counterpart as a spleen on the right, so these

two organs (the liver and spleen) must themselves correspond to each

other, and as such, express their respective significations. Under the

belief that every exception (even though it be normal) to a general law

or rule, is, like the anomaly itself, alone explicable according to such

law, and expressing a fact not more singular or isolated from other

parallel facts than is one form from another, or from all others

constituting the graduated scale of being, I would, according to the

light of this evidence alone, have no hesitation in stating that the

liver and spleen, as opposites, represent corresponding organs, even

though they appeared at first view more dissimilar than they really are.

In support of this analogy of both organs, which is here, so far as I am

aware, originally enunciated for anatomical science, I record the

following observations:--1st. Between the opposite parts of the same

organic entity (between the opposite leaves of the same plant, for

example), nature manifests no such absolute difference in any case as

exists between the leaf of a plant and of a book. 2ndly. When between

two opposite parts of the same organic form there appears any

differential character, this is simply the result of a modification or

metamorphosis of one of the two perfectly similar originals or

archetypes, but never carried out to such an extreme degree as to

annihilate all trace of their analogy. 3rdly. The liver and the spleen

are opposite parts; and as such, they are associated by arteries which

arise by a single trunk (coeliac axis) from the aorta, and branch right

and left, like indices pointing to the relationship between both these

organs, in the same manner as the two emulgent arteries point to the

opposite renal organs. 4thly. The liver is divided into two lobes, right

and left; the left is less than the right; that quantity which is

wanting to the left lobe is equal to the quantity of a spleen; and if in

idea we add the spleen to the left lobe of the liver, both lobes of this

organ become quantitatively equal, and the whole liver symmetrical;

hence, as the liver plus the spleen represents the whole structural

quantity, so the liver minus the spleen signifies that the two organs

now dissevered still relate to each other as parts of the same whole.

5thly. The liver, as being three-fourths of the whole, possesses the

duct which emanates at the centre of all glandular bodies. The spleen,

as being one-fourth of the whole, is devoid of the duct. The liver

having the duct, is functional as a gland, while the spleen having no

duct, cannot serve any such function. If, in thus indicating the

function which the spleen does not possess, there appears no proof

positive of the function which it does, perhaps the truth is, that as

being the ductless portion of the whole original hepatic quantity, it

exists as a thing degenerate and functionless, for it seems that the

animal economy suffers no loss of function when deprived of it. 6thly.

In early foetal life, the left lobe of the liver touches the spleen on

the left side; but in the process of abdominal development, the two

organs become separated from each other right and left. 7thly. In

animals devoid of the spleen, the liver appears of a symmetrical shape,

both its lobes being equal; for that quantity which in other animals has

become splenic, is in the former still hepatic. 8thly. In cases of

transposition of both organs, it is the right lobe of the liver--that

nearest the spleen, now on the right side--which is the smaller of the

two lobes, proving that whichever lobe be in this condition, the spleen,

as being opposite to it, represents the minus hepatic quantity. From

these, among other facts, I infer that the spleen is the representative

of the liver on the left side, and that as such, its signification being

manifest, there exists no exception to the law of animal symmetry. "Tam

miram uniformitatem in planetarum systemate, necessario fatendum est

intelligentia et concilio fuisse effectam. Idemque dici possit de

uniformitate illa quae est in corporibus animalium. Habent videlicet

animalia pleraque omnia, bina latera, dextrum et sinistrum, forma

consimili: et in lateribus illis, a posteriore quidem corporis sui

parte, pedes binos; ab anteriori autem parte, binos armos, vel pedes,

vel alas, humeris affixos: interque humeros collum, in spinam excurrens,

cui affixum est caput; in eoque capite binas aures, binos oculos, nasum,

os et linguam; similiter posita omnia, in omnibus fere animalibus."

--Newton, Optices, sive de reflex, &c. p. 411.]

X.--The heart, though being itself the recipient, the prime mover, and

the dispenser of the blood, does not depend either for its growth,

vitality, or stimulus to action, upon the blood under these uses, but

upon the blood circulating through vessels which are derived from its

main systemic artery, and disposed in capillary ramifications through

its substance, in the manner of the nutrient vessels of all other organs.

The two coronary arteries of the heart arise from the systemic aorta

immediately outside the semilunar valves, situated in the root of this

vessel, and in passing right and left along the auriculo-ventricular

furrows, they send off some branches for the supply of the organ itself,

and others by which both vessels anastomose freely around its base and

apex. The vasa cordis form an anastomotic circulation altogether

isolated from the vessels of the other thoracic organs, and also from

those distributed to the thoracic parietes. The coronary arteries are

accompanied by veins which open by distinct orifices (foramina Thebesii)

into the right auricle. Like the heart itself, its main vessels do not

depend for their support upon the blood conveyed by them, but upon that

circulated by the small arteries (vasa vasorum) derived either from the

vessel upon which they are distributed, or from some others in the

neighbourhood. These little arteries are attended by veins of a

corresponding size (venules) which enter the venae comites, thus

carrying out the general order of vascular distribution to the minutest

particular. Besides the larger nerves which accompany the main vessels,

there are delicate filaments of the cerebro-spinal and sympathetic

system distributed to their coats, for the purpose, as it is supposed,

of governing their "contractile movements." The vasa vasorum form an

anastomosis as well upon the inner surface of the sheath as upon the

artery contained in this part; and hence in the operation for tying the

vessel, the rule should be to disturb its connexions as little as

possible, otherwise its vitality, which depends upon these minute

branches, will, by their rupture, be destroyed in the situation of the

ligature, where it is most needed.

XI.--The branches of the systemic aorta form frequent anastomoses with

each other in all parts of the body. This anastomosis occurs chiefly

amongst the branches of the main arteries proper to either side. Those

branches of the opposite vessels which join at the median line are

generally of very small size. There are but few instances in which a

large blood vessel crosses the central line from its own side to the

other. Anastomosis at the median line between opposite vessels happens

either by a fusion of their sides lying parallel, as for example (and

the only one) that of the two vertebral arteries on the basilar process

of the occipital bone; or else by a direct end-to-end union, of which

the lateral pair of cerebral arteries, forming the circle of Willis, and

the two labial arteries, forming the coronary, are examples. The

branches of the main arteries of one side form numerous anastomoses in

the muscles and in the cellular and adipose tissue generally. Other

special branches derived from the parent vessel above and below the

several joints ramify and anastomose so very freely over the surfaces of

these parts, and seem to pass in reference to them out of their direct

course, that to effect this mode of distribution appears to be no less

immediate a design than to support the structures of which the joints

are composed.

XII.--The innominate artery. When this vessel is tied, the free direct

circulation through the principal arteries of the right arm, and the

right side of the neck, head, and brain, becomes arrested; and the

degree of strength of the recurrent circulation depends solely upon the

amount of anastomosing points between the following arteries of the

opposite sides. The small terminal branches of the two occipital, the

two auricular, the two superficial temporal, and the two frontal,

inosculate with each other upon the sides, and over the vertex of the

head; the two vertebral, and the branches of the internal carotid, at

the base and over the surface of the brain; the two facial with each

other, and with the frontal above and mental below, at the median line

of the face; the two internal maxillary by their palatine, pharyngeal,

meningeal, and various other branches upon the surface of the parts to

which they are distributed; and lastly, the two superior thyroid

arteries inosculate around the larynx and in the thyroid body. By these

anastomoses, it will be seen that the circulation is restored to the

branches of the common carotid almost solely. In regard to the

subclavian artery, the circulation would be carried on through the

anastomosing branches of the two inferior thyroid in the thyroid body;

of the two vertebral, in the cranium and upon the cervical vertebrae; of

the two internal mammary, with each other behind the sternum, and with

the thoracic branches of the axillary and the superior intercostal

laterally; lastly, through the anastomosis of the ascending cervical

with the descending branch of the occipital, and with the small lateral

offsets of the vertebral.

XIII.--The common carotid arteries, Of these two vessels, the left one

arising, in general, from the arch of the aorta, is longer than the

right one by the measure of the innominate artery from which the right

arises. When either of the common carotids is tied, the circulation will

be maintained through the anastomosing branches of the opposite vessels

as above specified. When the vertebral or the inferior thyroid branch

arises from the middle of the common carotid, this vessel will have an

additional source of supply if the ligature be applied to it below the

origin of such branch. In the absence of the innominate artery, the

right as well as the left carotid will be found to spring directly from

the aortic arch.

XIV.--The subclavian arteries. When a ligature is applied to the inner

third of this vessel within its primary branches, the collateral

circulation is carried on by the anastomoses of the arteries above

mentioned; but if the vertebral or the inferior thyroid arises either

from the aorta or the common carotid, the sources of arterial supply in

respect to the arm will, of course, be less numerous. When the outer

portion of the subclavian is tied between the scalenus and the clavicle,

while the branches arise from its inner part in their usual position and

number, the collateral circulation in reference to the arm is maintained

by the following anastomosing branches:--viz., those of the

superficialis colli, and the supra and posterior scapular, with those of

the acromial thoracic; the subscapular, and the anterior and posterior

circumflex around the shoulder-joint, and over the dorsal surface of

the scapula; and those of the internal mammary and superior intercostal,

with those of the thoracic arteries arising from the axillary. Whatever

be the variety as to their mode or place of origin, the branches

emanating from the subclavian artery are constant as to their

destination. The length of the inner portion of the right subclavian

will vary according to the place at which it arises, whether from the

innominate artery, from the ascending, or from the descending part of

the aortic arch.

XV.--The axillary artery. As this vessel gives off throughout its whole

length, numerous branches which inosculate principally with the

scapular, mammary, and superior intercostal branches of the subclavian,

it will be evident that, in tying it above its own branches, the

anastomotic circulation will with much greater freedom be maintained in

respect to the arm, than if the ligature be applied below those

branches. Hence, therefore, when the axillary artery is affected with

aneurism, thereby rendering it unsafe to apply a ligature to this

vessel, it becomes not only pathologically, but anatomically, the more

prudent measure to tie the subclavian immediately above the clavicle.

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