Bleeding is very rapidly followed by a large inflow of fluid into
the circulating blood—this fluid being derived from all the tissues,
and especially again from the muscles. Or again, when the
bandage from the limb in the above-cited experiment was
removed, the total capacity of the circulatory system was
thereby suddenly increased, and it was found that the total
volume of blood increased correspondingly, the increased volume
of fluid being drawn from the tissues and especially again from
the muscles. The rapidity with which this movement of fluid
into or out of the blood takes place is very striking. The explanation
usually offered is that the movement is effected by
changes in the capillary pressure due to the alteration in the
volume of blood circulating. While this seems feasible when the
volume of blood is increased, it does not offer a satisfactory
explanation of the rapid movement of fluid from the tissues
when the volume of the blood is decreased. One must therefore
look for yet further factors in this instance.
2. Let us next turn attention to the second of our three main variations, viz. that in which the composition of the blood is altered. It has long been known that the injection of water, or of solutions of soluble bodies such as salts, urea, sugar, &c., leads to a very rapid exchange of water and salts between the blood and the tissues. Thus if a solution less concentrated than the blood be injected, the blood is thereby diluted, but with very great rapidity water leaves the blood and is taken up by the tissues. Again, if a strong sugar or salt solution be injected, the first effect is a big discharge of water from the tissues into the blood and the movement of fluid is effected with great rapidity. In these instances a new physical factor is brought into play, viz. that of osmosis. When a solution of lower osmotic pressure than the blood is injected the osmotic pressure of the blood falls temporarily below that of the tissues, and water is therefore attracted to the tissues. The converse is the case when a solution of osmotic pressure higher than the blood is injected. This at first sight seems to be an all-sufficient explanation of the results recorded, but difficulties arise when we find that the tissues are not equally active in producing the effects. Thus it is found that the muscles and skin act as the chief water depot, while such tissues as the liver, intestines or pancreas take a relatively small share in the exchange. Again, when a strong sodium chloride solution is injected a considerable part of the sodium chloride is soon found to have left the blood, and it has been shown that the chloride depot is not identical with the water depot. The lung, for instance, is found to take up relatively far more of the salt than other tissues. Simultaneously with the passage of the salt into the tissue an exchange of water from the tissue into the blood can be observed, both processes being carried out very rapidly. The result is that the blood very quickly returns to a state in which its osmotic pressure is only slightly raised; the tissue, on the other hand, loses water and gains salt, and its osmotic pressure and specific gravity therefore rises. Again, the tissues do not participate equally in producing the final result, nor is the tissue which gives up the largest amount of water necessarily that which gains the largest amount of salt. The results following the injection of solutions of other bodies of small molecular size, e.g. urea or sugar, are quite analogous to those above described in the case of the non-toxic salt solutions. Hence we see that the rate of exchange of fluid and dissolved substance between a tissue and the blood can be extremely rapid and that the exchange can take place in either direction. We may also conclude that the main cause of the exchange, and possibly the only one, is the osmotic action set up by the solution injected, and that muscle tissue is particularly active in the process.
Seeing that a very considerable amount of water or of dissolved substance can be taken up from the blood into a tissue, the question next arises: Where is this material held, in the tissue cell or in the tissue space? Immediately the water or salt leaves the blood it reaches the tissue space, but unless the process be extreme in amount it probably passes at once into the tissue cell itself and is stored there. If the process is excessive oedema is set up and fluid accumulates in the tissue space.
These, taken quite briefly, are some of the more important conditions under which fluid exchanges take place. They are selected here because of the extent and rapidity of the changes effected.
3. The third factor which may bring about a change in the amount of fluid sent to a tissue is a variation in the capillary pressure. A rise in capillary pressure will, if filtration can occur through the capillary wall, cause an increased exudation of fluid from the blood. Thus the rise in general blood-pressure following the injection of a salt solution could cause an increased filtration into the tissues. Or again, the hydraemia following a salt injection would favour an increased exudation because the blood would be more readily filtrable. We, however, know very little of the effect of changes in capillary pressure upon movement of fluid into the tissue spaces and tissues, most of such observations being confined to a study of their effect upon lymph-flow. We will therefore return to them in this connexion.
4. The remaining factor to be mentioned is a change in the character of the capillary wall. It is well known that many poisons can excite an increased exudation from the blood and the tissue may become oedematous. Of such bodies we may mention cantharidin and the lymphogogues of Class I (see later). A like change is also probably the cause of the oedema of nephritis and of heart disease. It has also been suggested that the capillaries of different organs show varying degrees of permeability, a suggestion to which we will return later.
Lymph Formation.—There are two theories current at the present day offering explanations of the manner in which lymph is formed. The first, which owes its inception to Ludwig, explains lymph formation upon physical grounds. Thus according to this theory the lymphatics are open capillary vessels at their origin in the tissues along which the tissue fluid is driven. The tissue fluid is discharged from the blood by filtration, and therefore its amount varies directly with the capillary pressure. The amount of fluid movement also is further determined by osmotic actions and by the permeability of the capillary wall.
The second theory first actively enunciated by Heidenhain regards lymph formation as a secretory process of the capillary wall, i.e. one in the discharge of which these cells perform work and are not merely passive as in the former theory. As we shall see, it is now probable that neither theory is completely correct.
In considering lymph formation we have to examine both the total amount of lymph formed in the body and the variations in amount leaving each separate organ under different conditions. In most investigations the lymph was collected from the thoracic duct, i.e. it was the lymph returned from all parts of the body with the exception of the right arm and right side of the head and neck. The collection of the lymph from organs is much more difficult to effect, and hence has not, to the present, been so extensively studied. We will consider first variations in the amount of the thoracic duct lymph. Lymph is always flowing along the thoracic duct, and if the body is at rest, it has been shown that this lymph is coming practically entirely from the intestines and liver, chiefly, moreover, from the liver. The variations in the amount flowing under various conditions has been extensively studied. We will discuss them under the following headings: Changes brought about (a) by altered circulatory conditions, (b) by the injection of various substances, and (c) as a result of throwing an organ into activity.
Ligature of the portal vein leads to an increased flow of duct lymph. Ligature of the inferior vena cava above the diaphragm also leads to a large increase in the flow of duct lymph. Ligature of the aorta may result in either an increased or decreased flow of direct lymph. One explanation of these results has been offered from a study of the changes in capillary pressure set up in the main organs involved. Thus, after ligature of the portal vein the capillary pressure in the intestines rises, and it was proved that the increase in thoracic duct lymph came from the intestines. Ligaturing the inferior vena cava causes a big rise in the pressure in the liver capillaries, the intestinal capillary pressure remaining practically unaltered. Here it was proved that the increase in lymph-flow came from the liver and was
