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HAEMOSPORIDIA

slough off with the pile in about ten days, and in about ten days more the individual is, as a rule, well enough to return to his work. If, for one reason or another, no operation is to be undertaken, and the piles are troublesome, relief may be afforded by warm sponging and by sitz-baths, the pile being gently dried afterwards by a piece of soft linen, smeared with vaseline, and carefully returned into the bowel. Under surgical advice, cocaine or morphia may be brought in contact with the tender parts, either in the form of lotion, suppository or ointment. In operating upon internal piles it is undesirable to remove all the external piles around the anus, lest the contraction of the circumferential scar should cause permanent narrowing of the orifice. If, as often happens, blood clots in the vein of an external pile, the small, hard, tender swelling may be treated with anodyne fomentations, or it may be rendered insensitive by the ether spray and opened by a small incision, the clot being turned out.  (E. O.*) 


HAEMOSPORIDIA, in zoology, an order of Ectospora, which although comparatively few in number and very inconspicuous in size and appearance, have of late years probably attracted greater attention and been more generally studied than any other Sporozoa; the reason being that they include the organisms well known as malarial parasites. In spite, however, of much and careful recent research—to a certain extent, rather, as a result of it—it remains the case that the Haemosporidia are, in some respects, the group of the Ectospora about which our knowledge is, for the time being, in the most unsatisfactory condition. Such important questions, indeed, as the scope and boundaries of the group, its exact origin and affinities, the rank and interclassification of the forms admittedly included in it, are answered quite differently by different workers. For example, one well-known Sporozoan authority (M. Lühe) has recently united the two groups, Haemosporidia and Haemoflagellates, bodily into one, while others (e.g. Novy and McNeal) deny that there is any connexion whatever between “Cytozoa” and Trypanosomes. Again, the inclusion or exclusion of forms like Piroplasma and Halteridium is also the subject of much discussion. The present writer accepts here the view that the Haemosporidia are derived from Haemoflagellates which have developed a gregariniform (Sporozoan) phase at the expense, largely or entirely, of the flagelliform one. The not inconsiderable differences met with among different types are capable of explanation on the ground that certain forms have advanced farther than others along this particular line of evolution. In other words, it is most probable that the Haemosporidia are to be regarded as comprising various parasites which represent different stages intermediate between, on the one side, a Flagellate, and on the other, a typical chlamydospore-forming Ectosporan parasite. While, however, it is easy enough sharply to separate off all Haemosporidia from other Ectospora, it is a very difficult matter to define their limits on the former side. Two principal criteria which a doubtful haemal parasite might very well be required to satisfy in order to be considered as a Haemosporidian rather than a Haemoflagellate are (a) the occurrence of schizogony during the “corpuscular” phase in the Vertebrate host, and (b) the formation of many germs (“sporozoites”) from the zygote; so long as these conditions were complied with, the present writer, at all events, would not feel he was countenancing any protozoological heresy in allowing for the possibility of a Flagellate (perhaps trypaniform) phase or features being present at some period or other in the life-cycle.[1] To render this article complete, however, one or two well-known parasites, hitherto referred to this order, must also be mentioned, which, judged by the above (arbitrary) standard, are, it may be, on the Haemoflagellate side of the dividing line (e.g. Halteridium, according to Schaudinn).

The chief characters which distinguish the Haemosporidia from other Ectospora are the following. They are invariably blood parasites, and for part or all of the trophic period come into intimate relation with the cellular elements in the blood. There is always an alternation of hosts and of generations, an Invertebrate being the definitive host, in which sexual conjugation is undergone and which is to be regarded as the primary one, a Vertebrate being the intermediate or secondary one. The zygote or sporont is at first capable of movement and known as an ookinete. No resistant spores (chlamydospores) are formed, the ultimate germs or sporozoites always being free in the oocyst and not enclosed by sporocysts.

To Sir E. Ray Lankester is due the honour of discovering the first Haemosporidian, a discovery which did not take place until after most of the other kinds of Sporozoa were known. In 1871 this author described the parasite of the frog, which he later termed Drepanidium ranarum. The next discovery was the great and far-reaching one of Laveran, who in 1883 described all the characteristic phases of the malarial parasite which are met with in human blood. While regarding the organism as the cause of the disease, Laveran did not at once recognize its animal and Sporozoan nature, but considered it rather as a vegetable, and termed it Oscillaria malariae. As in the case of the Trypanosomes, we owe to Danilewsky (1885–1889) the first serious attempts to study the comparative anatomy and life-history of these parasites, from a zoological point of view. Danilewsky first named them Haemosporidia, and distinguished between Haemocytozoa and Leucocytozoa. To the brilliant researches of R. Ross and Grassi in the closing years of the 19th century is due the realization of the essential part played by the gnat or mosquito in the life-cycle and transmission of the parasites; and to MacCallum belongs the credit of first observing the true sexual conjugation, in the case of a Halteridium. Since then, thanks to the labours of Argutinsky and Schaudinn, our knowledge of the malarial parasites has steadily increased. Until quite recently, however, very little was known about the Haemosporidia of cold-blooded Vertebrates; but in 1903 Siegel and Schaudinn demonstrated that the same rôle is performed in their case by a leech or a tick, and since then many new forms have been described.

The Haemosporidia are widely distributed and of very general occurrence among the chief classes of Vertebrates. Among Invertebrates they are apparently limited to bloodsucking insects, ticks and leeches.[2] As already stated, the universal habitat of the parasites in the Vertebrate Occurrence: habitat; effects on host. is the blood; as a result, of course, they are to be met with in the capillaries of practically all the important organs of the body; and it is to be noted that while certain phases (e.g. growing trophozoites, mature gametocytes) are found in the peripheral circulation, others (e.g. schizogonous “rosettes,” young gametocytes) occur in the internal organs, liver, kidneys, &c., where the circulation is sluggish. The relation of the parasites to the blood-cells varies greatly. Most attack, probably exclusively, the red blood corpuscles (haematids); a few, however, select the leucocytes, and are therefore known as Leucocytozoa. In the case of Mammalian and Avian forms (malarial parasites) Schaudinn and Argutinsky have shown that the trophic and schizogonic phases are not really endoglobular but closely attached to the corpuscle, hollowing out a depression or space into which they nestle; the gametocytes, on the other hand, are actually intercellular. Forms parasitic in cold-blooded Vertebrates, on the contrary, are always, so far as is known, endoglobular when in relation with the corpuscles; and the same is apparently the case with the Mammalian parasite, Piroplasma. Although in no instance so far described is the parasite actually intranuclear (as certain Coccidia are), in one or two cases (e.g. Karyolysus of lizards and certain species of Haemogregarina) it reacts markedly upon the nucleus and soon causes its disintegration. While many Haemosporidia (e.g. malarial parasites, with the exception of Halteridium) remain in connexion with the same corpuscle throughout the whole period of growth and schizogony, the new generation of merozoites first being set free from the broken-down cell, others (the Haemogregarines,

  1. Compare, for example, the flagellated granules of certain Coccidia, which point unmistakably to a Flagellate ancestry.
  2. A possible exception is a doubtful species of Haemogregarina, which has been described from the walls of the blood-vessels of an Annelid.