From what has been brought forward no doubt can now remain that the question has been decided quite in Ehrlich's favour.
But what then are the physiological functions of the spleen, since that organ is unnecessary for the persistence of life? Doubtless its chief duty is the taking up of the greater part of the decaying fragments of red and white blood corpuscles in the blood-stream, so that this valuable material is not quite lost for the organism. Thus Ponfick has found that after destruction of the red corpuscles the spleen takes up a portion of their "shadows," and for this reason calls the splenic tumour a spodogenous splenic tumour (σποδος, ruins). Ehrlich has made a corresponding observation for the products of dissolution of the white blood corpuscles, and has proved that the splenic tumour which occurs in many infectious diseases and in phosphorus poisoning is to a large extent caused by the parenchyma of the spleen taking up the remains of the neutrophil protoplasm.
The question of the relation of the spleen to the fresh formation of red blood corpuscles is a problem of comparative anatomy. Observations on this point made on one kind of animal can certainly not claim validity for other kinds. In lower vertebrates, as in fishes, frogs, tortoises, and also in birds, the blood-forming activity of the spleen is pronounced and of great importance. In mammalia on the other hand, in some cases this function cannot be demonstrated, and in others only to a very small degree. In the spleen of normal mice nucleated red blood corpuscles are seen in relatively large numbers; in the rabbit they are less numerous and often only to be found with difficulty. In the dog they only make their appearance after anæmia from loss of blood, normally they are absent. In the human spleen nucleated red blood corpuscles are not to be found normally or in cases of severe anæmia, but exclusively in leukæmic diseases. U. Gabbi in his recently published work on the hæmolytic function of the spleen, also emphasises the difference between the various animal species. In guinea-pigs he found that the spleen acts largely as a scavenger of the red blood corpuscles; in rabbits very slightly. Consequently after removal of the spleen in guinea-pigs the number of red blood corpuscles rose 377,000 in the cubic millimetre, and the amount of hæmoglobin 8.2%. After splenectomy in rabbits the increase in these values is absent.
Shortly summarising our analysis of the facts before us, we must say that the importance of the spleen for the production of the white blood corpuscles can in no respect be considerable, and that if these cells really are produced by it, they must be free from granulations. The spleen therefore stands functionally in closer connection with the lymphatic gland system than with the bone-marrow. The spleen has not the least connection with ordinary leucocytosis[14].
(β) The Lymphatic Glands.
As it is impossible experimentally to prevent the lymphatic glands as a whole from contributing to the formation of the blood, we are dependent almost entirely on clinical and anatomical researches for an elucidation of their function.
Since Virchow's definition of the lymphocyte it has been admitted that the lymphocytes of the blood, both the small and larger kinds, are identical with those of the lymphatic glands and the rest of the lymphatic system. This is proved by the complete agreement in general morphological character, in staining properties of the protoplasm and nucleus, and from the absence of granulation.
Abundant clinical experience testifies that the lymphocytes of the blood really do arise from the lymphatic system. Ehrlich had previously observed that when extensive portions of the lymphatic glandular system are put out of action by new growths and similar causes, the number of the lymphocytes may be considerably diminished. These observations have since that time been confirmed by various authors. For example, Reinbach describes several cases of malignant tumour, particularly sarcoma, in which the percentage of lymphocytes, which normally amounts to about 25%, was very considerably lowered; in one case of lymphosarcoma of the neck they only made up 0.6% of the total number. These conditions are quite easily and naturally explained by the exclusion of the lymphatic glands. It is difficult for the advocates of the view that the lymphocytes are the early stages of all white blood corpuscles to reconcile it with these facts. According to their scheme the low number of lymphocytes is to be explained in such cases by their unusually rapid transformation to the polynuclear elements—the old forms; or to appropriate the expression of Uskoff, by a too rapid ageing of the lymphocytes.
Further evidence for the origin of the lymphocytes of the blood from the lymphatic glands is to be obtained from those cases in which we find an increase of the lymphocytes in the blood. These "lymphocytoses" occur, in comparison with other leucocytoses, relatively seldom. Under certain conditions in which a hyperplasia of the lymphatic glandular apparatus makes its appearance, we often see at first an increase of the lymphocytes in the blood. Ehrlich and Karewski in some unpublished work have investigated together a large number of typical cases of lymphoma malignum, and were able constantly to observe a lymphocytosis, which in some cases was of high degree and bore almost a leukæmic character.
Relying on these facts Ehrlich and Wassermann (Dermatolog. Zeitschr. Vol. i., 1894) made the diagnosis in vivo of malignant lymphoma in a rare skin disease, chiefly from the absolute increase of the lymphocytes alone, although no swelling of the glands was palpable. The post-mortem shewed that the chief condition was a swelling of the retro-peritoneal lymph glands to lumps as large as a fist.