Uma nova modalidade de sexo-determinação no grilo sul-americano eneoptera surinamensis

The male of Eneoptera surinamensis (Orthoptera-Eneopteridae) is provided with 9 chromosomes, that is, with 3 pairs of autosomes and 3 sex chromosomes. Spermatogonia. - The autosomes of the spermatogonia are of the same size and U-shaped. One of the sex chromosomes approximately equalling the autosomes in size is telocentric, while the other two are much larger and V-shaped. One of the latter is smaller than the other. The sex chromosomes as showed in Figs. 1 and 2 are designated by X, Yl and Y2, X being the larger V, Yl the smaller one and Y2 the rod-shaped. Primary spermatocytes. - Before the growth period of the spermatocytes all the three sex chromosomes are visible in a state of strong heteropycnosis. X is remarkable in this stage in having two long arms well separated by a wide commissural segment. (Figs. 4, 5 and 6). During the growth period Y2 disappears, while X and Yl remain in a condensed form until metaphase. These may be separated from one another or united in the most varied and irregular manner. (Fig. 7 to 12). In the latter case the segments in contact seem to be always different so that we cannot recognize any homology of parts in the sense os genetics. At diplotene Y2 reappears together with the autosomal tetrads. X and Yl may again be seen as separate or united elements. (Figs. 13 and 14). At later diakinesis and metaphase the three sex chromosomes are always independent from each other, Y2 being typically rod-shaped, X and Yl V-shaped, X being a little larger than Yl. (Fig. 15 to 18). At metaphase the three condensed tetrads go to the equatorial plane, while the sex chromosomes occupy any position at both sides of this plane. In almost all figures which could be perfectly analysed X appeared at one side of the autosomal plate an Yl together with Y2 far apart at the other side. (Figs. 16 and 18). Only a few exception have been found. (Figs. 17 and 19). At anaphase X goes in precession to one pole, Yl and Y2 to the other (Figs. 20 and 21). As it is suggested by the few figures in which a localization of the sex chromosomes different from the normal has been observed, the possibility of other types of segregation of these elements cannot be entirely precluded. But, if this does happen, the resulting gametes should be inviable or give inviable zygotes. Early in anaphase autosomes and sex chromosomes divide longitudinally, being maintained united only by the kinetochore. (Figs. 20 and 21). At metaphase the three sex chromosomes seem to show no special repulsion against each other, X being found in the proximity of Yl or Y2 indifferently. At anaphase, however, the evidences in hand point to a stronger repulsion between X on the one side and both Ys on the other, so that in spite of the mutual repulsion of the latter they finish by going to the same pole. Secondary spermatocytes. - At telophase of the primary spermatocytes all the chromosomes enter into distension without disappearing of view. A nuclear membrane is formed around the chromosomes. All the chromosomes excepting Y2 which has two arms, are four-branched. (Fig. 22). Soon the chromosomes enter again into contraction giving rise to the secondary metaphase plate. Secondary spermatocytes provided as expected with four and five chromosomes are abundantly found. (Figs. 23 and 24). In the former all chromosomes are X-shaped while in the latter there is one which is V-shaped. This is the rod- shaped Y2. In the anaphase of the spermatocytes with four chromosomes all the chromosomes are V-shaped, one of them (X) being much larger than the others. In those with five there is one rod-shaped chromosome (Y2). (Fig. 25), Spermatids. Two classes of spermatids are produced, one with X and other with Yl and Y2. All the autosomes as well as Y2 soon enter into solution, X remaining visible for long time in one class and Yl in the other. (Figs. 26 and 27). Since both are very alike at this stage, one cannot distinguish the two classes of spermatids. Somatic chromosomes in the famale. - In the follicular cells of the ovary 8 chromosomes were found, two of which are much larger than the rest. (Figs. 29 and 30). These are considered as being sex chromosomes. CONCLUSION: Eneoptera surinamensis has a new type of sex-determining mechanism, the male being X Yl Y2 and the female XX. The sex chromosomes segregate without entering into contact at metaphase or forming group. After a review of the other known cases of complex sex chromosome mechanism the author held that Eneoptera is the unique representative of a true determinate segregation of sex chromosomes. Y2 behaving as sex chromosome and as autosome is considered as representing an intermediary state of the evolution of the sex chromosomes.

Provas adicionais da dicentricidade dos cromossômios dos Hemípteros

In order to test Piza's conclusions regarding the dicentricity of Hemipteran chromosomes, two species of bugs of the family Coreidae, namely, Anasa sp. and Leptoglossus stigma (Herbst), are studied in the present paper. a) Anasa sp. - The male of this species has 21 chromosomes, that is, 20 pairs of autosomes and a single sex chromosome. The latter divides equationally in the first division of the spermatocytes and passes undivided to one cell in the second division. In this it moves with its longer axis parallelly to the spindle axis and shows fibrillar connections with both poles. Special attention was paid to the behavior of the chromosomes in the anaphase of the spermatogonia. As it was previously stated (Piza 1946 and 1946a) with regard to other species, the chromosomes are here attached to the spindle by both ends and begin to move toward the poles strongly curved to them. No intercalary fibers could be detected although their existente may not be denied by theoretical reasons developed in another paper (Piza 1946). Mitoses in somatic tissues of the embryo were equally studied. Careful examination of anaphase chromosomes in a great number of cells showed that the chromosomes behave exactly as in the spermatogonia, being equally attached to the spindle by the extremities alone and moving with their ends looking to the pole. A weak median constriction sometimes replaced by a slightly clearer space was observed in prometaphase and even in metaphase chromosomes of the spermatogonia as well as the somatic cells, having already been referred to in the case of Diactor bilineatus. (Piza 1945). Hemipteran chromosomes being considered as iso-chromosomes originated by a longitudinal spliting of the monocentric chromosomes resulting from the second division of the spermatocytes, the median aspect just mentioned may be regarded as the point of union of the separated halves. (See origin of dicentricity in Piza 1946). b) Leptoglossus stigma - This species has spermatogonia provided with 20 pairs of autosomes and one sex chromosome whose behavior differs in nothing from what was stated in regard of the preceding species. In the primary spermatocytes nothing meriting special mention was observed. Orientation, connection with the poles and movements of the sex chromosome in the secondary spermatocytes confirm the views already developed.

Nota prévia sôbre a meiose de Corizus (Liorhyssus) hyalinus (Fabr.) (Hemiptera-Corizidae)

The main facts presented in this paper may be summarized as follows: 1) Corizus (Liorhyssus) hyalinus (Fabr.) has primary spermatocytes provided with 6 autosomal tetrads, one pair of microchromosomes and one sex chromosome. 2) The two microchromosomes present in this species sometimes appear at the primary metaphase as an unequal pair of minute elements. In the secondary spermatocytes the unique microchromosome present may be in the limit of visibility or entirely invisible. This invisibility may be partly due to a loss of colourability. 3) The sex chromosome divides transversely in the first division of the spermatocyte, passing undivided to one pole in the second one. In the latter it becomes fusiform in the beginning of anaphase revealing in this manner its dicentricity. In late anaphase it finishes by passing to one pole leaving in the other pole one of its kinetochores sometimes accompanied by a chromosomal fragment. 4) All the chromosomes divide transversely in both divisions, a diagram being enclosed to elucidate the question. 5) Spermatogonial chromosomes are provided with one kinetochore at each end, being curved toward the poles since the most beginning anaphase. 6) The following hypothesis is presented as an essay to explain the origin of microchromosomes: Since microchromosomes parallel sex chromosomes in most respects, as for instances in heteropycnosis and pairing modus, it seems highly probable that they originate from sex chromosomes. One may suppose that the ancestral form of a given species had a sex chromosome which used to lose a small centric fragment when it divided during meiosis. This fragment might well be at first an unstable one. Later, to compensate the effects of such a deficiency a mechanism arose through evolution which produced two useful results : a) the establishment of the fragment as a permanent structure of the cell nucleus and b) the acquirement by the sex chromosome of the faculty of passing to one pole without losing any of its ends.