ALBERT

Notes: SYNOPSIS. The photographs illustrate male and female gametes before fertilization, several progressive stages in the entrance of the male gamete into the cytoplasm of the female, cytoplasmic fusion of gametes, loss of extranuclear organelles of male gamete, retention of extranuclear organelles of female gamete, movement of pronucleus of male gamete to that of female, progressive stages in fusion of pronuclei, and the formation of the zygote which possesses the extranuclear organelles of the female gamete. Some abortive attempts at fertilization, resulting from failure of gametes to differentiate, are shown.

Notes: The following types of transfauntions were made: from nymphs is an early stage of their molting period to nymphs 1-50 days before ecdysis and vice versa; from recently molted nymphs to nymphs in all stages of the molting period and vicve versa. In such transfaunated half or more of the protozoa in a donor host were transferred to a recipient host. The recipient hosts wee defaunated with oxygen by subjecting them to 60 psi for 2 hours, twice the minimum time necessary to kill all their protozoa. Transfaunations were made 24 hours after defaunation.The sexual cycles, as a result of these transfaunation experiments, in addition to being prevented altogether in several genera, were modified principally in two ways: they were shortened to less than half the time required under natural conditions; they were lengthened, in some genera, from two days to over two months. Whether shorteing or lengthening occurs depends on the type of host used in the experiment. And the factor in the host that is responsible for the modifications is the ecdysone titer; transfaunation from a lower to higher to a lower titer, in general, serves to lengthen it.

Notes: When exogenous ecdysone is injected into a host (adult or intermolt nymph) which has no ecdysone of its own, gametogenesis is induced in the protozoa of such a host, although the host itself never undergoes ecdysis. The time required to induce this sexual process varies in different genera and with the dosage of hormone administered; but it is never possible, regardless of the amount of hormone given, to induce gametogenesis in all the genera at the same time; some still begin a day or two later than others, but never 40-45 days later as occurs in untreated roaches.Likewise, when exogenous ecdysone is administered to a nymphal host in the molting period, the sexual cycles of its protozoa are greatly accelerated, and the host undergoes ecdy-sis much sooner than would otherwise have been the case. Both sexuality in the protozoa and the molting process in the host are accelerated. Only when large doses of ecdysone are administered are the sexual cycles of the protozoa modified in any way except a decided acceleration. Quite large amounts of ecdysone present special problems for the protozoa, problems they never encounter in nature. The growth and differentiation hormone ecdysone causes the protozoa to grow and differentiate so rapidly that many of them cannot keep pace; as a result, degeneration begins, and death follows. But this over-acceleration effect of ecdysone and death of the protozoa occurs only in those genera which have not undergone gametogenesis when the large amount of hormone is administered; those which have completed gametogenesis are not affected at all. The results of acceleration have been observed in most genera, but have been studied more intensively in Tricho-nympha, where they are usually seen to be considerably more pronounced on the nuclei and chromosomes than on the cytoplasm. As a result, this genus, which invariably in nature undergoes only fertilization, is sometimes made to undergo either autogamy or endomitosis. The pronuclei, for example, differentiate and are ready to (and do) fuse before the cytoplasm of the gametocyte divides to form gametes. In the case of endomitosis, precocious differentiation of chromosomes seems to produce, or at least goes along with, premature degeneration of one centriole, the one that would ordinarily degenerate following fusion of gametes. Thus, nuclear division is made impossible.In some genera of the protozoa, the ecdysone titer must drop greatly before meiosis can begin. If the titer, after ecdysis. when it normally drops precipitously, is kept high with injections, meiosis does not begin; it is retarded for as long as the ecdysone titer is kept high.

Notes: SYNOPSIS. Gigantomonas usually exists in the multinucleate stage. Hence, multiple fission is more common than binary fission. In its flagellate as well as its aflagellate state it is always an amoeboid cell which possesses from one to several large, clear pseudopodia. Fairly often in the multinucleate, as well as the uninucleate, stage no extranuclear organelles are present except large plain centrioles, two of which are always closely associated with each nucleus. During nuclear reproduction four centrioles, two old ones and two new ones, are always present with each nucleus. During all nuclear reproductions, regardless of the number of nuclei present, extranuclear organelles, such as flagella, axostyle, undulating membrane, and costa when present are discarded. If they are renewed, it is by the new centrioles at the same time that the old ones produce new central spindles, two always cooperating in the process. Thus, Gigantomonas, like other genera of the Devescovinidae, the family to which it belongs, never has for each nucleus present more than one set of extranuclear organelles, a characteristic which Devescovinidae and Lophomonadidae have in common. It is the only genus of Devescovinidae without a parabasal body.Owing to the liquidness of the cytoplasm, the central spindle, which becomes very long indeed, often extends beyond the cytoplasm and thus pushes the nucleus fastened to this end of it completely out of the cell. Mainly because of this situation, multinucleate forms with an odd number of nuclei occur often; otherwise the nuclear numbers would be 2, 4, 8, 16, 32, etc., because, no matter how many nuclei are present, they all reproduce simultaneously or nearly so.An unusual situation occurs in which Gigantomonas ingests and digests a small species of Holomastigotoides, while several hundred individuals of the latter become attached to and destroy Gigantomonas.

Notes: SYNOPSIS. The gametes of Trichonympha grandis fuse quite differently from those of all the other species of Trichonympha in Cryptocercus; fusion is partial and temporary instead of complete and permanent as in other species. In this species, as in the others, fusion brings about the disintegration of all the extranuclear organelles of the male gamete, but none of those of the female; these persist to become the organelles of the zygote. Gametic union accomplishes two things: loss of extranuclear organelles of male and fusion of pronuclei. Unlike that of the other species of Trichonympha, the cytoplasm of the male gamete of T. grandis contributes slightly, if at all, to the formation of the zygote.

Notes: SYNOPSIS. The role of eleven different types of achromatic figures in chromosomal movement of Barbulanympha is analyzed. When only one pole is present, no chromosomes are ever connected with it, and hence they do not move. The chromosomes go through their usual life cycle including pairing, but remain in the parent nucleus, which, of course, does not divide. When two poles are present with only one pole near the nucleus, the poles, which are the distal ends of the elongate centrioles, do not cooperate in the formation of a central spindle and the chromosomal behavior is just as if there were only one pole—no movement. The same is true when more than two poles are present with only one near the nucleus.Unless a central spindle is present, movement of chromosomes never occurs. However, when many central spindles are present, sister chromosomes may separate and move to poles which are not directly connected by a central spindle. In other words, sisters may separate without moving along a central spindle.In binucleate cells with one central spindle the chromosomes of one nucleus move to the poles, but those of the other do not. Movement always occurs in the nucleus that has its nuclear membrane depressed by the central spindle. When two or more central spindles are present, the chromosomes of both nuclei may move to the poles.

Notes: SYNOPSIS. Under the influence of ecdysone a uninucleate asexual cell of Paranotila lata gen. et sp. nov. is transformed into a gametocyte in which, as a rule, 8 successive nuclear divisions occur, thus producing 8 male and 8 female pronuclei. No cytokinesis occurs during these divisions. The process should be regarded as a gametogenesis in which only nuclei participate. These gametic pronuclei fuse, male with female, and produce 8 fusion or zygotic nuclei. The cell may now be regarded as a zygote, which soon undergoes plasmotomy, and eventually produces 8 uninucleate, diploid asexual cells. This is the usual course of events. A few exceptions or irregularities have been noted. Throughout gametogenesis, Paranotila is quite indifferent regarding the production of new flagella and axostyles; sometimes they are produced, sometimes they are not.

Notes: SUMMARY. Two centrioles, an old one and a new one, are always present in the resting cell. From prophase onward two new ones and two old ones are present. Beginning with the resting stage, five types of centriole life cycles are described and compared with one another: In type 1, both centrioles are elongate; in type 2, the old one is long and the new one, which is short, elongates in prophase; in type 3, both are short, both elongate in prophase, and both, except for their anterior tips, degenerate in late telophase; in type 4, both are long but in prophase their distal ends become free of the rest of the centrioles, these ends migrate to center or posterior end of cell, where, after they produce the achromatic figure and it completes its function in nuclear division, they degenerate; in type 5, both are short and neither elongates at any stage of its life cycle.New centrioles are produced by the anterior ends of old ones. In their first generation, centrioles produce only extranuclear organelles (flagella, parabasals, axostyles, etc.); in their second and later generations, they produce only the achromatic figure (gametogenesis in Trichonympha and reorganization in Barbulanympha and Rhynchonympha are exceptions to this rule).The distal ends of centrioles in some types of cycles are surrounded by centrosomes; in others they are not. In one type of centriole life cycle a small central spindle is present in the resting cell in two genera; in the other types this is not the case.

Notes: SUMMARY. The role of centrioles in achromatic figure production is considered when the number present varies from 1–8. Each centriole after it becomes elongate produces astral rays from its distal end. Some of these rays remain free; some, by joining centromeres, become chromosomal fibers; and some, by joining and growing along those produced by one or more other centrioles, produce the central spindle portion of the achromatic figures. Thus, one centriole may function cooperatively with one to several others in the production of central spindles. But at least two centrioles must be present, and in the proper spacial relation to each other, to form a central spindle; one by itself can form only free astral rays, no central spindle or chromosomal fibers.The flagellated areas (to which the centrioles are anchored anteriorly) play an important role in determining the position of the distal ends of the centrioles with respect to one another, and the position of these ends, in turn, in a large measure, determines the types of achromatic figures produced, particularly the number of central spindles.

Notes: SUMMARY. Oxygen concentrations of 70–80 per cent of an atmosphere destroy all chromosomes of the flagellate Trichonympha provided the oxygen treatment is carried out during the early stages of gametogenesis at which time the chromosomes are in the process of duplicating themselves. This treatment does no damage to the cytoplasm and its organelles. Following the loss of chromosomes, the centrioles function in the production of the achromatic figure, the flagella, and‘parabasal bodies. Then the cytoplasm divides, thus producing two anucleate gametes which make some progress in the cytoplasmic differentiations characteristic of normal male and female gametes of Trichonympha.It is also possible, with somewhat higher concentrations of oxygen, with temperatures slightly above the freezing point and a longer period of treatment, to destroy the chromosomes of resting asexual nuclei in several genera of the flagellates that live in the roach Cryptocercus. So far as one can determine by observing organisms so treated, their cytoplasm and organelles are not injured.