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Binucleate Cells in Insects: Mitotic Division and Chromosome Configuration, Lecture notes of Cell Biology

The presence and mitotic division of binucleate cells in the follicular epithelium and sperm duct of insects, specifically in Notonecta. the stages of mitosis, including the resting stage, prophase, metaphase, anaphase, and telophase, and the possible variations in spindle arrangement. The document also mentions the chromosome composition of binucleate cells in oogonial and follicle cells of Notonecta and other insects.

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MITOTIC DIVISION OF BINUCLEATE CELLS.
ETHEL BROWNE HARVEY.
Attention has been called to the presence of binucleate cells
in the follicular epithelium of insects .by several observers
(Korschelt, ‘¿87,Preusse, ‘¿95,De Bruyne, ‘¿97,Gross, 0!, et al.).
In Notonecta, binucleate cells are characteristic of the follicular
epithelium and also of the tissues lining the sperm duct and in
vesting the spermatogonial cysts. These binucleate cells undergo
normal mitotic division, both nuclei being involved. As a
usual rule, both nuclei pass through the stages of division syn
chronously, though rarely one finds one nucleus in advance of
the other. (Fig. ii, upper cell.)
In the resting stage (Fig. I), each nucleus is exactly similar to
the single ftucleus of a spermatogonial or oogonial cell, a con
spicuous karyosphere being present (Browne, ‘¿13).In the pro
phase, the two nuclei remain distinct and thick chromatin strands
are present in each nucleus (Fig. 2, upper cell, Fig. 3). When
the nuclear wall breaks down in the late prophase (Fig. 4), the
two nuclei may undergo a certain amount of fusion, or they may
remain entirely distinct and this is true also of the metaphase.
In some cases, two distinct plates may be seen in polar view
(Fig. 5) or two distinct spindles in side view (Fig. 6); in fact,
the two spindles may be so independent as to lie in different
planes, so that one is seen in side view and the other in polar
view (Fig, 7). In other cases, the two spindles become so in
timately combined as to appear as one giant spindle. In polar
view a perfect equatorial plate is seen, but there are twice as
many chromosomes present as the diploid number for the species.
Each chromosome of the spermatogonial or oögonial metaphase
is represented by two exactly similar chromosomes in these
binuclear plates, as may be seen by comparing Fig. 8 from an
oogonial cell of Notonecta indica, with Fig. 2 (lower cell) from
a follicle cell of the same species. In the oogonial cell, there
are 6 large, 4 small and 16 intermediatu sized chromosomes, in
96
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MITOTIC DIVISION OF BINUCLEATE CELLS.

ETHEL BROWNE HARVEY.

Attention has been called to the presence of binucleate cells in the follicular epithelium of insects .by several observers (Korschelt, ‘¿ 87,Preusse, ‘¿ 95,De Bruyne, ‘¿ 97,Gross, 0!, et al.). In Notonecta, binucleate cells are characteristic of the follicular epithelium and also of the tissues lining the sperm duct and in vesting the spermatogonial cysts. These binucleate cells undergo normal mitotic division, both nuclei being involved. As a usual rule, both nuclei pass through the stages of division syn chronously, though rarely one finds one nucleus in advance of the other. (Fig. ii, upper cell.) In the resting stage (Fig. I), each nucleus is exactly similar to the single ftucleus of a spermatogonial or oogonial cell, a con spicuous karyosphere being present (Browne, ‘¿ 13).In the pro phase, the two nuclei remain distinct and thick chromatin strands are present in each nucleus (Fig. 2, upper cell, Fig. 3). When the nuclear wall breaks down in the late prophase (Fig. 4), the two nuclei may undergo a certain amount of fusion, or they may remain entirely distinct and this is true also of the metaphase. In some cases, two distinct plates may be seen in polar view (Fig. 5) or two distinct spindles in side view (Fig. 6); in fact, the two spindles may be so independent as to lie in different planes, so that one is seen in side view and the other in polar view (Fig, 7). In other cases, the two spindles become so in timately combined as to appear as one giant spindle. In polar view a perfect equatorial plate is seen, but there are twice as many chromosomes present as the diploid number for the species. Each chromosome of the spermatogonial or oögonial metaphase is represented by two exactly similar chromosomes in these binuclear plates, as may be seen by comparing Fig. 8 from an oogonial cell of Notonecta indica, with Fig. 2 (lower cell) from a follicle cell of the same species. In the oogonial cell, there are 6 large, 4 small and 16 intermediatu sized chromosomes, in 96

MITOTIC DIVISION OF BINUCLEATE CELLS. 97

the binucleate follicle cell there are 12 large, 8 small and @ intermediate ones. Similar double chromosome plates have been found in the follicle cells of other insects, e.g., Anasa (Wil son, ‘¿ oó)and are probably likewise due to the division of a binucleate cell. In the anaphase (Fig. 9), the two sets of chromosomes cahnot be distinguished, and the two, spindles appear always to be fused. It is difficult to tell exactly what happens in the telophase. It would seem that a membrane formed around the group of chro mosomes at each pole, (Fig. io), each daughter nucleus becoming elongated and that it later separated off into two nuclei (Figs. II and 12). It seems entirely possible that in such a separation into two nuclei, which would resemble a nuclear amitosis, the chromosome content might be so distributed that each nucleus would maintain the chromosome constitution characteristic of the species. It is possible that the separation into two nuclei occurs only in those cases where the original sister nuclei did not become entirely fused and that when they are fused, each daughter nucleus remains as an abnormally large nucleus, really double in nature. It is evident, however, from the series of figures that two binucleate cells may arise by the mitotic division of a binucleate cell. Although the binucleate cells of the follicular epithelium have been believed by some observers (Preusse, ‘¿ 95,De Bruyne, ‘¿ 97,Gross, ‘¿ oi),to arise by an amitotic division, I have found no evidence of such an origin, and would believe it probable that they arise by the failure of cell division after a nuclear divi sion by mitosis. Macklin (‘16)has described a process of amito tic nuclear division in his study of binucleate cells in living tissue, but the figures he gives might also be interpreted as the last stages in the mitotic division of a binucleate cell as described above. If binucleate cells do arise as Macklin believes and his interpretation of them as potentially mononuclear, is correct, there must be two types of binucleate cells. In one type, each of the two nuclei has a complete set of chromosomes, as described above. In the other type, the two nuclei taken together have a complete set of chromosomes. Macklin was, however, unfor tunately unable to verify his assumption as to the chromosome

100 ETHEL BROWNE HARVEY.

EXPLANATION OF PLATE I. Fig.i from the sperm duct of Notonectashooierii;Fig.8 from oogonia!cellof N. indica, other figures from follicle cells of N. indica. FIG. i. Binucleate cells from sperm duct of N. shooteris, resting stage. FIG. 2. Upper cell, prophase. Lower cell, metaphase showing a perfect double chromosome group, 52 chromosomes. FIG. 3. Prophase. FIG. 4. Late prophase. FIG. 5. Metaphase, polar view, two plates separate. FIG. 6. Metaphase, side view, two plates separate. FIG. 7. Metaphase, two spind!es in different planes. FIG. 8. Metaphase of odgonial cell of N. indica, showing 26 chromosomes. FIG. 9. Anaphase. FIG. 10. Telophase. FIG. ii. Lower cell, telophase, sister plates dividing into two. Upper cell one nucleus in prophase. one in metaphase. FIG. 12. Late telophase showing two daughter binucleate cells.

BIOLOGICAL BULLETIN, VOL XXXVI. (^) PLATE I

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