increased activity entails an increased rate of supply of oxygen. This latter has been brought about in the animal kingdom in two ways, first by an increase in the concentration of the haemoglobin of the blood effected by an increase both in the number of corpuscles and in the amount of haemoglobin contained in each, and secondly by an increase in the rate at which the blood has been made to pass through the tissues. In the lower vertebrates the blood pressure is low and the haemoglobin content of the blood is low, consequently both rate of blood-flow and oxygen-content are low. In contrast with this, in higher vertebrates the blood pressure is high and the haemoglobin content of the blood is high, consequently both rate of blood-flow and oxygen-content are high. We must associate with this important step in evolution the means employed for the more rapid absorption of oxygen and for its increased rate of discharge to the tissues, the most important features of which are a diminution in the size of the corpuscle and the attainment of its peculiar shape, both resulting in the production of a relatively enormous corpuscular surface in a unit volume of blood.
Variations are also found in the white corpuscles as well as in the red, but these differences are not so striking and lie chiefly in unimportant details of structure of individual cells. Enormous variations are to be found in different species of mammals, but the cells generally conform to the types of secreting cells or phagocytes.
The platelets also differ in the different species. In the frog, for instance, many are spindle-shaped and contain a nucleus-like structure. Birds’ blood is stated to contain no platelets. The variations in number of these bodies have not been satisfactorily ascertained on account of the difficulties involved in any attempt to preserve them and to render them visible under the microscope.
Differences are also found in the chemical composition of the plasma. The chief variation is in the amount of protein present, which attains its maximum concentration in birds and mammals, while in reptiles, amphibia and fishes it is much less. The bloods of the latter two classes are much more watery than that of the mammal. Moreover, it has been proved that there are specific differences in the chemical nature of the various proteins present even between different varieties of mammals. Thus the ratio of the globulin fraction to the albumin fraction may vary considerably, and again, one or other of the proteins may be quite specific for the animal from which it is derived.
Clotting.—If a sample of blood be withdrawn from an animal, within a short time it undergoes a series of changes and becomes converted into a stiff jelly. It is said to clot. If the process is watched it is seen to start first from the surfaces where it is in contact with any foreign body; thence it extends through the blood until the whole mass sets solid. A short time elapses before this process commences—a time dependent upon two chief conditions, viz. the temperature at which the blood is kept and the extent of foreign surface with which it is brought into contact. Thus in a mammal the blood clots most quickly at a temperature a little above body temperature, while if the blood be cooled quickly the clotting is considerably delayed and in the case of some animals altogether prevented. For example, human blood kept at body temperature clots in three minutes, while if allowed to cool to room temperature the first sign of clotting may not make its appearance until eight minutes after its removal from the body. The process of clotting is also considerably accelerated by making the blood flow in a thin stream over a wide surface. The full completion of the process occupies some time if the blood be kept quiet, but ultimately the whole mass of the blood becomes converted into a solid. At this stage the containing vessel may be inverted without any drop of fluid escaping. A short time after this stage has been reached drops of a yellow fluid appear upon the surface and, increasing in size and number, run together to form a layer of fluid separated from the clot. This fluid is termed serum; its appearance is due to the contraction of the clot, which thus squeezes out the fluid from between its solid constituents. Contraction continues for about twenty-four hours, at the end of which time a large quantity (one-third or more of the total volume) of serum may have been separated. The clot contracts uniformly, thus preserving throughout the same general shape as that of the vessel in which the blood has been collected. Finally the clot swims freely in the serum which it has expressed.
The cause of the clot formation has been found to be the precipitation of a solid from the liquid plasma of the blood. This solid is in the form of very minute threads and hence is termed fibrin. The threads traverse the mass of blood in every possible direction, interlacing and thus confining in their meshes all the solid elements of the blood. Soon after their deposition they begin to contract, and as the meshwork they form is very minute they carry with them all the corpuscles of the blood. These with the fibrin form the shrunken clot.
If the rate at which blood clots be retarded either by cooling or by some other process the corpuscles may have time to settle, partially or completely, in which case distinct layers may form. The lowermost of these contains chiefly the red corpuscles, the second layer may be grey owing to the high percentage of leucocytes present, while a third, marked by opalescence only, may be very rich in platelets. Above these a clear layer of fluid may be found. This is plasma. The formation of these layers depends solely upon the rate of sedimentation of these elements, the rate depending partly upon differences in specific gravity, and partly upon the tendency the corpuscles have to run into clumps. Horse’s blood offers one of the best instances of the clumping of red corpuscles, and in this animal sedimentation of the red corpuscles is most rapid.
If now such a sedimented blood is allowed to clot the process is found to start in the middle two layers, i.e. in those containing the white corpuscles and platelets. From these layers it spreads through the rest of the liquid, being most retarded, however, in the red corpuscle layer, and particularly so if the sedimentation has been very complete. Not only does the clotting process start from the layers containing the leucocytes and platelets, but in them it also proceeds more quickly. These observations clearly indicate that the clotting process is initiated by some change starting from these elements.
The object of the clotting of the blood is quite clear. It is to prevent, as far as possible, any loss of blood when there is an injury to an animal’s vessels. The shed blood becomes converted into a solid, and this, extending into the interior of the ruptured vessel, forms a plug and thus arrests the bleeding. It is found that clotting is especially accelerated whenever the blood touches a foreign tissue, for instance, the outer layers of a torn blood-vessel wall, muscle tissue, &c., i.e. in exactly those conditions in which rapid clotting becomes of the greatest importance. Yet another very pregnant fact in connexion with clotting is that if an animal be bled rapidly and the blood collected in successive samples it is found that those collected last clot most quickly. Hence the more excessive the haemorrhage in any case, the greater becomes the onset of the natural cure for the bleeding, viz. clotting.
When we begin to inquire into the nature of clotting we have to determine in the first place whence the fibrin is derived. It has long been known that two chemical substances at least are requisite for its production. Thus certain fluids are known, e.g. some samples of hydrocele or pericardial fluid, which will not clot spontaneously, but will clot rapidly when a small quantity of serum or of an old blood-clot is added to it. The constituent substance which is present in the first-named fluids is known as fibrinogen, and that present in the serum or the clot is known as fibrin-ferment or thrombin.
Fibrinogen is present in living blood dissolved in the plasma; it is also present in such fluids as hydrocele or pericardial effusions, which, though capable of clotting, do not clot spontaneously. Thrombin, on the other hand, does not exist in living blood, but only makes its appearance there after blood is shed. It is not yet certain what is the nature of the final reaction between fibrinogen and thrombin. The possibilities are, that thrombin may act—(1) by acting upon fibrinogen, which it in some way converts into fibrin, (2) by uniting with fibrinogen to form fibrin, or (3) by yielding part of itself to the fibrinogen which thus
