471. The theory of Dr. Crawford is essentially the same, which supposes that venous blood contains a peculiar compound of carbon and hydrogen, termed hydro-carbon, the elements of which unite in the lungs with the oxygen of the air, forming water with the one and carbonic acid with the other. Mr. Cooper, for many years past, has taught the same doctrine in his lectures, without any knowledge of the fact that Crawford had suggested a similar modification of his theory.

472. It is now established that more oxygen disappears than is accounted for by the amount of carbonic acid that is generated. The experiments of Dr. Edwards had already shown this in so decisive a manner that physiologists almost universally admitted it as an ascertained fact. The calculations of Mr. Finlaison, to whom the opinions of physiologists on this point were unknown, have now determined the precise amount of oxygen (444 et seq.), and the probable amount of azote ([459]) absorbed. By many physiologists it is supposed that the oxygen retained by the lungs, as long as it remains in this organ, enters only into a state of loose combination with the blood; that in this state of loose combination, it is carried from the lungs into the general system; and that it is only in the system that the union becomes intimate and complete. According to this view, the lungs are merely the portal by which the substances employed in respiration are received and discharged, the essential changes induced taking place in the system. That it is through the lungs that the oxygen required by the system is received, is an opinion founded on experiments no less exact than decisive; it is in accordance with the most probable theory of the production and distribution of animal heat (chap. ix.); and the preponderance of evidence in its favour is so great that, in the present state of our knowledge, it may be considered as established; but it will appear hereafter that the lungs are by no means passive in the process, and that, physiologically considered, they as truly constitute a gland secreting carbonic acid gas as the liver is a gland secreting bile.

473. Such are the main facts which have been ascertained relative to respiration, as far as this function is performed by the lungs. But the liver is a respiratory organ as well as the lungs. It decarbonizes the blood. It carries on this process to such an extent, that some physiologists are of opinion that the liver is the chief organ by which the decarbonization of the blood is effected. The following considerations show that whatever be the relative amount of its action, the liver powerfully co-operates with the lungs in the performance of a respiratory function.

1. The liver, like the lungs, is a receptacle of venous blood; blood loaded with carbon. The great venous trunk which ramifies through the lungs is the pulmonary artery, containing all the blood which has finished its circuit through the system. The great venous trunk which ramifies through the liver is the vena portæ, containing all the blood which has finished its circuit through the apparatus of digestion. The liver is a secreting organ, distinguished from every other secreting organ by elaborating its peculiar secretion from venous blood. Carbon is abstracted from the venous blood that flows through the lungs in the form of carbonic acid; carbon is abstracted from the venous blood that flows through the liver in the form of bile.

2. All aliment, but more especially vegetable food, contains a large portion of carbon, more it would appear than the lungs can evolve. The excess is secreted from the blood by the liver, in the form of resin, colouring matter, fatty matter, mucus, and the principal constituents of the bile. All these substances contain a large proportion of carbon. After accomplishing certain secondary purposes in the process of digestion, these biliary matters, loaded with carbon, are carried out of the system together with the non-nutrient portion of the aliment. In the decarbonizing process performed by the lungs and the liver, the chief difference would seem, then, to be in the mode in which the carbon that is separated is carried out of the system. In the lungs it is evolved, as has been stated, in union with oxygen in the form of carbonic acid; in the liver, in union with hydrogen in the form of resin and fatty matter.

3. Accordingly, in tracing the organization of the animal body from the commencement of the scale, it is found that among the distinct and special organs that are formed, the liver is one of the very first. It would appear to be constructed as soon as the economy of the animal requires a higher degree of respiration than can be effected by the nearly homogeneous substance of which, very low down in the scale, the body is composed. Invariably through the whole animal series, the magnitude of the liver is in the inverse ratio to that of the lungs. The larger, the more perfectly developed the lungs, the smaller the liver; and conversely, the larger the liver the smaller and the less perfectly developed the lungs. This is so uniform that it may be considered as a law of the animal economy. In the highly organized warm-blooded animal, with its large lungs, divided into numerous lobes, and each lobe composed of minute vesicles respiring only air, the magnitude of the liver compared with that of the body is small. In the less highly organized animal of the same class, with its smaller and less perfectly developed lung, respiring partly air and partly water, the liver increases as the lung diminishes in size. In the reptile with its little vesicular lung, divided into large cells, the liver is proportionally of greater magnitude. In the fish which has no lung, but which respires by the less highly organized gill, and only in the medium of water, the proportionate size of the liver is still greater; but in the molluscous animal, in which the lung or the gill is still less perfectly developed, the bulk of the liver is prodigious.

4. In all animals the quantity of venous blood which is sent to the liver increases, as that transmitted to the lung diminishes. In the higher animal the great venous trunk which ramifies through the liver (the vena portæ) is formed by the veins of the stomach, intestines, spleen, and pancreas, which are the only organs that transmit their blood to the liver. In the reptile, besides all these organs, the hind legs, the pelvis, the tail, the intercostal veins forming the vena azygos and in some orders of this class, even the kidneys also send their blood to the liver; but in the fish, in addition to all the preceding organs, the apparatus of reproduction likewise transmits its blood to the liver. The very formation of the venous system in the different classes of animals seems thus to point to the liver as a compensating and supplementary organ to the lung.

5. The permanent organs in the lower animal are a type of the transitory forms through which the organs of the higher animal pass in the progress of their growth. Thus the liver of the human fœtus is of such a disproportionate size, as to approximate it closely to that of the fish or of the reptile. After the birth of the human embryo, respiration is effected in part by the lung; but before birth the lung is inactive, no air reaches it; it contributes nothing to respiration; the decarbonizing action of the blood is accomplished, not by the lung, but by the liver; hence the prodigious bulk of the fœtal liver and its activity in the secretion of bile, and especially towards the latter months of pregnancy, when all the organs are greatly advanced in size and completeness.

6. Pathology confirms the evidence derived from comparative anatomy and physiology. When the function of the lung is interrupted by disease, the activity of the liver is increased. In inflammation of the lung (pneumonia); in the deposition of adventitious matter in the lung (tubercles), by which the air vesicles are compressed and obliterated, the lung loses the power of decarbonizing the blood in proportion to the extent and severity of the disease with which it is affected. In this case the secretion of bile is increased. In diseases of the heart the liver is enlarged. In the morbus cæruleus ([516]) the liver retains through life its fœtal state of disproportion.

7. In the last place, there is a striking illustration of the respiratory action of the liver, in the vicarious office which it performs for the lung, during the heat of summer in cold, and all the year round in hot climates. In the heat of summer, and more especially in the intense and constant heat of a warm climate, in consequence of the rarefaction of the air, respiration by the lung is less active and efficient than in the winter of the cold climate. During the exposure of the body to this long-continued heat, there is a tendency to the accumulation of carbon in the blood. An actual accumulation is prevented, by an increased activity in the secretion of bile, to which the liver is stimulated by the heat. In order to obtain the material for the formation of this unusual quantity of bile, it abstracts carbon largely from the blood; to this extent it compensates for the diminished efficiency of the lung, and thus removes through the vena portæ that superfluous carbon which would otherwise have been excreted through the pulmonary artery.