326
L. DONOASTBR AND H. a. CANNON.
EXPLANATION OF PLATE .16,
Illustrating Prof. L. Doncaster's and Mr. H. G\ Cannon's paper
on "The Spermatogenesis of the Louse (Pediculus
corporis and P. capitis), with some Observations on
the Maturation of the Egg."
DKSCIUL'TION Ob' FlGUBES.
[The fignres are from freehand drawings, made with Zeiss apochromatic N.A. 140, 3 mm. or 2 mm. objectives, and compensating ocular 12.
They are not all exactly to the same scale ; in most of the figures the
magnification is about 3000 diameters ; in figs. 1-4, 6-9, 38-50 it is about
twice as great. All figures are from preparations of P . c o r p o r i s
except when it is stated in the description that the figure is from
P . c a p i t i s . All figures except Nos. 7, 8, 15, 16, 17, 25 are from
drawings by H. G. Cannon.]
Figs. 1-14 illustrate chiefly the chromosomes in sperniatogonia and
spermatocytes.
Fig. 1.—-Equatorial plate from follicle cell of Stage II testis ; twelve
chromosomes. (Fleniming, acetic x 2.)
Fig. 2.—Equatorial plate from body-cell of Stage II female; twelve
chromosomes. (Flemming.)
Fig. 3 (P. capitis).—Equatorial plate from follicle cell of. Stage I I
testis; twelve chromosomes. (Flemming.)
Fig. 4 (P. capitis).—Equatorial plate from body-cell of Stage III
female; twelve chromosomes. (Flemming.)
Fig. 5.—Spermatogonial prophase, Stage I I I ; chromosomes partially
double and nucleolus. (Flemming.)
Fig. 6.—Typical spermatogonial equatorial plate, Stage I I I ; six
chromosomes. (Flemming.)
Fig. 7.—Spermatogonial equatorial plate from young Stage I nymph j
six chromosomes, all double. (Flemming.)
Fig. 8.—Spermatogonial equatorial plate from young Stage I nymph -f
six chromosomes. (Flemming.)
Fig. 9 (P. capitis).—Spermatogonial equatorial plate from Stage I I
male ; six chromosomes. (Flemming.)
Fig. 10.—Prophase of spermatocyte division, Stage I I male; six
double chromosomes; acroblast near the mitochondrial body. (Flemming,
acetic X 2.)
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Fig. 11.—Metaphase of spermatocyte division, Stage III male;
spindle at one side of the cell. (Flemming.)
Fig. 12.—Early anaphase of spermatocyte division, Stage III male;
the finger-shaped process has hardly begun to appear; almost certainly
six chromosomes at each pole. (Flemming.)
Fig. 13 (P. capitis).—Later anaphase of spermatocyte division,
. Stage II male ; appearance of finger-shaped process ; six chromosomes
are clearly visible in the body of the cell; the pair nearest the centrosome in the process are not quite certainly separate. (Flemming.)
[Note.—The acroblast and mitochondrial body are not preserved in
figs. 11-13.]
Fig. 14.—Late anaphase of spermatocyte division, Stage I I male;
the process has become long and narrow and the chromosomes are
aggregated in its tip; the large cell retains the niitochondrial body and
acroblast. (Flemming.)
[Note.—The complete separation of the small cell at the close of the
spermatocyte division is shown in figs. 28-30.]
Figs. 15-32 illustrate the development of the mitochondria! body,
acroblast and axial filaments in the spermatocyte and young
spermatid.
Fig. 15.—Young spermatocyte with first appearance of mitochondria.
Stage I I male. (F.w.a. J.)
Fig. 16.—Older spermatocyte ; mitochondrial mass showing vacuolation. Stage II male. (F.w.a. J.)
Fig. 17.—Older spermatocyte ; mitochondrial mass much vacuolated.
(F.w.a. followed by potassium bichromate.)
Figs 18-21.—Coalescence of mitochondrial vesicles intolarge vacuoles;
the appearance represented in fig. 20 is that most freq\iently seen. Fig.
18, from section preserved in Flemming; figs. 19-21, from smear
preparations.
Figs. 22, 23.—Coalescence of vacuoles to form two large vacuoles
enclosing a stained mitochondrial mass. (Smear preparations, F.w.a.)
Fig. 24.—Young spermatid, with acroblast next to the nucleus, double
centrosome, and typical mitosome of simple structure. (Flemming.)
Fig. 25.—Similar stage, from the same individual, with mitosome of
more complex structure. (Flemming).
Fig. 26.—Beginning of outgrowth of axial filaments; mitosome in
optical section. (Flemming.)
Fig. 27.—Further growth of axial filaments, with " distal centrosome " at posterior edge of cell. Mitosome seen from the side, so that
its bilateral structure is not visible. (F.w.a. £.)
328
L. DONCASTER AND H. G. CANNON".
Fig. 28.—Axial filaments beginning to protrude from cell; mitosome
seen obliquely. The small cell separated at the unequal spermatocyte
division is near t,he anterior end of the cell. (F.w.a, £.)
Fig. 29.—Similar cell from same testis, with further growth of axial
filaments. (F.w.a. i.)
Fig. 30.—Similar cell from same testis ; axial filaments united outside
cell (probably intertwined), and distal centrosomes at the end of short
processes. This condition is not general, and is possibly due to artefact. Small " polar " cell still attached to spermatid. (F.w.a. i.)
Fig. 31.—Beginning of elongation of spermatid; same testis as figs.
28-30. (F.w.a. h)
Fig. 32.—Transverse section of same stage as last, from same testis,
showing two axial filaments cut across just outside the mitosome, the
structure of which is well shown. The position of the nucleus, at a,
different focus, is indicated.
Figs. 33-50 illustrate the conversion of the spermatid into the sper• matozoon.
Fig. 33.—Elongation of the spermatid; mitosome still vacuolated.
(F.w.a. i.)
Figs. 34-37.—Continued elongation, with development of a swelling
at the posterior end, into which the greater part of the mitosome
becomes concentrated. The structure of the mitosome is no longer
distinguishable. (F.w.a. J.)
Fig. 37A is an oblique section of the head at the stage of fig. 37r
showing acroblast, two centrosomes and nucleus.
Figs. 38-50 illustrate the development of the head of the spermatozoon.
Figs. 38-42 (P. capitis).—Application of acroblast to the nucleus
(figs. 38, 39), its spreading forward like a cap (fig. 40) until it forms a
projection in front of 'the nucleus (fig. 41, face view, fig. 41A side-view).
Fig. 42 shows the deeply staining body at the posterior side of the
nucleus, apparently connected with the acrosome by a band. (Flemming.)
Figs. 43-49.—Elongation of the head and formation of the projecting
acrosome from the anterior portion of the acroblast. The stained body
at the posterior end of the nucleus is shown especially in figs. 45-47, and
figs. 45A and B represent optical transverse sections (A) through it, and
(B) in front of it. (All F.w.a. i.)
Fig. 48A repsesents a complete spermatozoon at the stage of fig. 48,
from a smear preparation. (Smaller scale.)
Fig. 50.—The head of a nearly mature spermatozoon; smear
preparation.
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SPERMATOGENESIS OF LOUSE.
342
H. M. OARLETON.
EXPLANATION OF PLATE
17.
I l l u s t r a t i n g Mr. H . M. Carleton's p a p e r on " Observations on
an Intra-nucleolar Body in Columnar Epithelium Cells
of the I n t e s t i n e . "
CE. Chromatin reticulum. CS. Chromosomes. GA. Golgi apparatus.
IF. Interzonal fibres. MI. Mitochondria. KY. Karyosome. PL. Plasmosome. SB. Striated border of columnar cells. SP. Spireme.
X. Nucleolinus.
[All the figures were outlined with camera lucida, pa,per at table level,
with Koritska -j-^th oil immersion and No. 12 compensating eye-piece.
Present magnification of fig. 1 is 1720 diameters, of figs. 2-16 3580
diameters approximately.]
Pig. 1.—Columnar epithelium cells from duodenum of cat, showing
Golgi apparatus, mitochondria and nucleolinus. Cajal-safranin.
Figs. 2 and 3.—Nuclei from same material, showing nucleolini
within the karyosome. Cajal-pyronin-methyl green technique. In the
original preparation the nucleolini are black, the karyosomes and the
chromatin network green.
Figs. 4 and 5.—Same as figs. 2 and 3. These figures show the
variations in the number of nucleolini, etc. Cajal-safranin.
Figs. 6 and 7.—Nuclei from intestinal epithelium of frog. Modified
Cajal-safranin.
Figs. 8 and 9.—From intestinal epithelium of frog. Modified Cajal
technique counterstained with pyronin-methyl green. In the original
preparations the nucleolini are black, the nucleoli (plasmosomes) bright
red, and the chromatin green.
Figs. 10 to 16 (with the exception of No. 14) all represent mitotic
figures from the columnar cells of the frog's intestine. With modified
Cajal-safranin technique employed, the nucleolini alone were impregnated black, chromosomes being stained red and the cytoplasm pink.
Fig. 10.—Early prophase; spireme beginning to segment; nucleolini
apparently about to divide.
Fig. 11.—A later prophase, section somewhat oblique, showing
equatorial plate and two pairs of nucleolini. Arrows in this ajid
succeeding figure indicate direction of mitosis.
Fig. 12.—Early anaphase, section rather oblique. The two chromosome groups are indicated, each with one pair of nucleolini.
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OBSERVATIONS ON AN INTRA-NCTOLEOLAR BODY.
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Fig. 13.—Prophase; equatorial plate in transverse section. Only two
nucleolini present.
Fig. 14.—Nucleus from intestinal epithelium of frog. Fixed in
Bouin and stained in toluidin blue-eosin. The nucleolinus ia plainly
visible.
!Fig. 15.—Late anaphase. Two groups of daughter-chromosomes
each with two nucleolini.
Fig. 16.—A still later (telophase) stage. Nuclear membrane reformed. Two nucleolini in each daughter-nucleus.
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408
D. AVARD CUTLER.
2. Dobell, C. C. (1914).—" Oytological Studies on Three Species of
Amcebee," 'Arch. Protistenk.,' Bd. xxxiv.
3.
and Jepps, M. (1918).—"A Study of Diverse Races of E nt amceba h i s t o l y t i c a distinguishable from one another by the
Dimensions of their Cysts," ' Parasit.,' vol. x.
4. Foa, A., and Grassi, B. (1904).—"Richerche sulla riproduzione dei
flagellati: 1. Processo di divisione della Jcenia," ' Rend. R. Ace.
Lincei,' vol. xiii.
5. Gatenby, J. B. (191V and 1918).—" Oytoplasmic Inclusions of the
Germ-cells," ' Quart. Journ. Micr. Sci.,' vols. 62, 63.
6. Grassi, B., and Foa, A. (1904).—"Richerche sulla riproduzione dei
flagellati: 1. Processo di divisione della Joenia e forme affini,"
' Rend. Atti. Accad. Lincei,' ser. 5, vol. xiii, fasc. 52 sem.
7.
(1911)-—"Intorno ai Protozoi dei Termitidi," 'Rend. R.
Accad. Lincei,' ser. 5, vol. xx (1).
*8.
(1917).—" Flagellati viventi nei Termiti," ' Mem. R. Accad.
Lincei,' ser. 5, vol. xii, fasc. 8.
•9. Hartmann, M. (1910).—" Untersuchungen iiber Bau und Entwicklung dei1 Trichonymphiden," ' Festchr. z. 60 Geburtstage Rich.
Hertwigs,' Jena, vol. i.
*1O. Imms, A. D. (1919).—" On the Structure and Biology of Archot e r m o p s i s , together with Descriptions of New Species of Intestinal Protozoa and General Observations iipon the Isoptera,"
' Phil. Trans. B.,' vol. ccix.
*11. Janicki, O. (1914-1915).—" Untersuchungen an parasitischen
Flagellaten," ' Zeit. f. wiss. Zool.,' Bd. cxii.
The papers marked * contain further references to the literature of
Termite parasites.
EXPLANATION OF PLATES 18 TO 21,
Illustrating Mr. D. Ward Cutler's paper on "Protozoa parasitic in Termites."
[All the figures, except figs. 1 and 2, are drawn from fixed and stained
preparations. The optical apparatus employed was as follows: Zeiss
apochromatic oil-immersion objective 2 mm. (N. A. 13) and compensating
oculars 6,12, 18. Critical illumination was always used. The method
PROTOZOA PAKASITIO IN TEIJMITES.
409
of fixation and staining and the appropriate magnification is given in
the case of each figure, except figs. 1 and 2.
The following abbrevations are employed: S.A. Schaudinn's sublimate-alcohol mixture. S.A. (D.J.) Schaudinn's sublimate-alcohol as
modified by Dobell and Jepps. Fl. (Gat.) Fleming's strong fluid as
modified by G-atenby. H.I.H. Heidenhain's iron-alum haematoxylin.
D.H. Dobell's iron-haematein.
The lettering of the figures on the plates is as follows: A.X. Axostyle. B.F. Body flagella. B.G. Basal granules. Fl. Flagella. G.B.
Groove. H.F. Head flagella. P.B. Parabasal body. P.T. Parabasal
thread. S.P. Supporting piece. S.T. Striations.]
PLATE 18.
Fig. 1.—Side view of living Joenopsis p o l y t r i c h a . Note the
supporting piece, parabasal thread and axostyle. The striations do
not appear in a right- and left-handed series as in the face view of the
animal seen in fig. 2.
Fig. 2.—Freehand drawing, as in fig. 1, of a living animal seen in face
view, showing the right- and left-handed arrangement of the striations,
also the groove dividing the animal into " head " and " body " regions.
The flagella in the living animal are more numerous than represented
in the drawing.
Fig. 3.—Stained preparation of J . p o l y t r i c h a , showing the supporting piece, parabasal threads and axostyle arising from the supporting piece. The basal granules and flagella are omitted from the
drawing. Parabasal bodies are not seen owing to the method of fixation.
x 450. S.A., D.H.
Fig. 4.—Highly magnified drawing of two striations with the basal
granules and flagella. X 1800. Fl. (Gat.), H.I.H.
Fig. 5.—Head region of J . p o l y t r i c h a with supporting piece and
its basal granules (flagella omitted). Parabasal threads and bodies
seen. Note the resting nucleus with its chromatin bodies embedded in
the plastin ground-work, x 950. Fl. (Gat.), H.I.H.
Fig. 6.—Highly magnified drawing of a parabasal thread with its
attached parabasal bodies, x 1000. Fl. (Gat,), H.I.H.
PLATE 19.
Fig. 7.—Resting nucleus of J . p o l y t r i c h a . x 950. S.A., D.H.
Fig. 8.—Section through the body of J . p o l y t r i c h a , showing the
410
D. WARD CUTLER..
folded surface (striations) and basal granules. X 1000. S.A. (D.J.),
H.I.H.
Tig. 9.—" Head " region of animal showing what is probably the first
stage of division. The organella are in most cases reduplicated, but the
nucleus is still in the resting condition, x 950. Fl. (Gat.), D.H.
Fig. 10.—" Head " region with dividing nucleus and double set of
organella. Parabasal bodies are not seen due to method of fixation.
x 950. S.A. (D. J.), D.H.
Fig. 11.—Anterior end of animal with supporting piece and basal
grannies (flagella omitted). The basal grannies on the striations are
seen, but the anterior ends of the striations are omitted. Note axostyle
arising from the supporting piece. Nucleus in the first stage of division
with the chromatin resolved into granules. X 950. S.A. (D.J.), H.I.H.
Fig. 12.—Nucleus with its so-called chromosomes; plastin in the form
of granules. X 1800. S.A., H.I.H.
Fig. 13.—Dividing nucleus with chromatin aggregated at each pole
and the plastin situated at equator. No Bpindle-fibres or centrioles
present, x 950. S.A. (D.J.), H.I.H.
Fig. 14.—Dividing nucleus in which membi-ane is constricting, x 950.
S.A. (D.J.), H.I.H.
Fig. 15.—Dividing J. p o l y t r i c h a in which two sets of organella are
seen. At the top of the animal the middle portion of the striations are
seen passing over the edge of the body. Axostyles omitted. X 450.
On the right of the animal the nuclei, more highly magnified, are drawn.
Note that the upper daughter-nucleus is in a more advanced stage of
reconstruction than the lower one. x 950. Fl. (Gat.), H.I.H.
Fig. 16.—Animal at the close of division; double set of organella,
with the exception of parabasal bodies. Striations and flagella omitted.
X 450. The daughter-nuclei, more highly magnified, are seen below.
X 950. S.A. (D.J.), H.I.H.
PLATE 20.
Fig. 17.—Dividing J . p o l y t r i c h a in which the parabasal bodies and
axostyles are not visible. X 450. As in the previous figures a more
highly magnified drawing of the nuclei is seen to the right. The lower
nucleus is more advanced in the reconstruction process than the upper
one. X 950. S.A. (D.J.), D.H.
Fig. 18.—Daughter-nuclei directly after separation, x 1800. S.A.
(D.J.), H.I.H.
Fig. 19.—A daughter-nucleus at the close of division. Note plastin
PROTOZOA
PARASITIC IN TERMITES.
411
granules and the large chromatin bodies, approximately thirty-two in
number. X 1800. Fl. (Gat), H.I.H.
Fig. 20.—Animal near the close of reproduction. X 450. The
daughter-nuclei below are magnified. X 950. Fl. (Gat.), H.I.H.
Fig. 21.—Drawing of a specimen of Jcenopsis c e p h a l o t r i c h a .
Note supporting piece with atriations and axostyle. Nucleus at
anterior end probably in the first stage of division. X 950. Fl.
(Gat.), H.I.H.
Fig. 22.—As above, but nucleus in resting stage; basal granules are
seen on the supporting piece. X 953. S.A., H.I.H.
PLATE 21.
Fig. 23.—J. c e p h a l o t r i c l i a seen from above. The striations are
observed radiating from the supporting piece and the nucleus between
the arms, x 950. S.A. (D.J.), H.I.H.
Fig. 24.—Animal at the end of division process, with two supporting
pieces and axostyles. X 950. Fl. (Gat.), H.I.H.
Fig. 25.—As above, x 950. S.A., D.H.
Fig. 26.—Stained specimen of Microjcenia axostylis. Note the
broad axostyle arising from tlie anterior extremity, also the thickened
anterior border with basal granules and flagella. x 950. S.A., D.H.
Fig. 27.—As above, x 950. Fl. (Gat.) H.I.H.
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