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The structure of Bowman's capsule as an index of age and sex variations in normal mice.

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T H E STRUCTURE O F BOWMAN'S CAPSULE AS AN
INDEX O F AGE AND S E X VARIATIONS
I N NORXAL MICE
CHARLOTTE CRABTREE
Department of Zoology, Smith College, Northampton, Massachzi.setts
SEVEN FIGURES
I. INTRODUCTION
Among mammals secondary sex characteristics include variations in a large number of unrelated structures. The size
and shape of the masseter muscles, the character and distribution of hair, subcutaneous f a t distribution, the weights
of various endocrine organs, and the shape of the glottis
are a few of the many possible variations noted between males
and females of the same species. A number of statements
in recent articles have indicated the possibility of sexual variations in the normal kidney. I n 1927 MacKay and R'IacKay
reported that the renal weights, in albino rats of all ages, bore
a direct and constant relationship to body surface measurements. I n a second paper published the same year (MacKay
and MacKay, '27 b) they modified this statement, indicating
that sex was a factor influencing renal weights. I n the normal
female albino rat the renal tissue per unit of body surface averages 27 mg. less than in the normal male. Hall and NcGregor
('3'7) found a sex difference in the renal weights of cats:
females and young males maintain a constant kidney weight/
body weight ratio while in adult males the ratio is markedly
increased owing to the deposition of fat in the tubular walls.
Contributions from the Zoology Department, Smith College, No. 192. A thesis
subinitted to the faculty of Smith College in partial fulfillinelit of t h e degree of
Master of Arts.
395
396
CHARLOTTE CBABTREE
An analagous condition has been demonstrated in the human
kidney by Wald ( '37) who found the mean absolute weight of
normal kidneys of the human female to be 40 to 75 gm. less than
those of a similar representative group of males.
There also exists a certain amount of evidence indicatiiig
that renal weights a r e under hormonal influence and can be
increased in both normal and gonadectomized animals by injections of sex hormones. Korenchevsky et al. found that
young castrate rats treated with testosterone showed a definite
and significant increase in renal weight (Korenchevsky, Dennison and Kohn-Speyer, ' 3 3 ) . I n the renal weights of castrated
rats, the same workers found an increase of 6% to 24% when
oestriii alone was injected, of 10% to 15% when male and
female sex hormones were injected simultaneously (Koreiichevsky and Dennison, '34). Selye observed a comparable
renal increase in normal female mice injected daily with small
amounts of testosterone propionate. Histological study of
these kidneys showed that the increase in weight was owing
to hypertrophy of the secreting tubules and also to the preeence of numerous modified Bowman's capsules of the type
to be described shortly (Selye, '39). These workers' results
indicate that sex hormones have a stimulating effect on kidney
weight.
The vertebrate Bowman's capsule generally pictured in
textbooks of normal histology is a goblet-shaped structure
consisting of a double-walled sac of flattened cells. The inner,
o r visceral layer of the capsule is continuous and non-syncytial
in nature (Rensley and Beiisley, '30) and is closely applied to
the capillary loops of the glomerular tuft. The outer, o r
parietal layer is usually composed of a continuom, flat layer
of epithelial cells which may or may not be separated from
the visceral layer by the presence of filtrate in the lumen between the two layers. The entire capsule is surrounded by
a structureless basement membrane which is deflected inward
at the vascular pole and separates the cells of the glomerular
endothelium from those of the visceral epithelium. The basement membrane is also continuous a t the urine pole with the
STRUCTURE OF BOWMAN’S CAPSULE IN MICE
397
membrane surrounding the urinifcrous tubule. This area of
juncture of the capsule arid the urinifcrous tubule is commonly
designated as the neck. I n many cold-blooded animals a group
of characteristically flattened ciliated cells make up a definite,
fairly long, thin-walled neck seapent of the uriniferous tubule
(von Rlollendorff, ’29). In reptiles, birds, and mammals,
however, the neck is either much reduced or entirely absent,
‘’LOW“ CELL CAPSULE
A
‘H\6YmCEU-CAPSULE
a
Fig. 1 A. Diagram of a ‘ L l o w ” cell capsule of the type usually described
:IS characteristic of niamnialinii kidneys. B. Diagraui of a “high ” cell capsule
with the squanious cells of tlie parietal layer replaced by cuboidal cells similar
to those of tlie proxiirial convoluted tubule.
and the transition from flat, squamous parietal cells of the
capsule t o cuboidal cells of the tubule is usually very abrupt
(see fig. 1 A ) .
The development of the capsule in the human fetus and in
iiifant,s in the first 2 years of postiiatal life has been worked out
in detail (Risak, ’28; Gruenwald and Popper, ’40). The cells
which develop into Bowman’s capsule are of mescnchpmal
origin and appear first as a rounded sac of cuboidal and/or
398
CHARLOTTE CRABTREE
columnar epithelium. As the adjaceiit glomerulus develops, it
invaginates this sac to form a primitive capsule. The epithelium of the parietal layer is first to flatten. This process may
occur even in the first half of embryonic life at which time
the loops of‘ the glomerulus a r e closely adherent and the entire
tuft is still covered with a layer of simple columnar epithelium
cliaractcrized by large numbers of closely-grouped nuclei.
a.
d.
b.
e.
C.
f.
CIS.
DEVELOPMENT
OF BOWMAN’S
CAPSULE
Fig. 2 Diagram showing the development of the human Bowman’s capsule.
Based 011 the description of Gruenwald and Popper ( ’40).
After birth the glomcrular loops fill with blood and expand,
while the epithelial cell layer ruptures. Deep fissures are
formed between the loops, thus exposing portions of naked
endothelial malls which appear to be covered gradually by a
thin layer of cells, the “glomerulothelium” of Randrath (’32)
forming the visceral layer of the adult Bowman’s capsule (see
fig. 2 ) . Whethei. the visceral layer of the capsule thus formed
STRUCTURE OF BOWMAN'S
CAPSULE IN MICE
399
is composed of cells modified from the primitive columnar
epithelium originally present, as claimed by Zimmerman ('33)
and others, or whether it is reticular in origin and develops
from cells carried into the capsule via the vascular pole, a s
suggested by von Mollendorff ( '27) and others, is a question
beyond the scope of this paper. It is sufficient here to point
out that, in the normal course of events, the parietal layer
of Bowman's capsule in the human is invariably observed to
be the first to flatten. These observations have been confirmed
by the author in a series of kidney sections from premature
infants from the Boston Lying I n Hospital.
This description applies to the structure and development
of the type of capsule most frequently observed and commonly
described for mammals. As early a s 1887, however, Benda
noted in mice kidneys a second type of capsule which differs
from the description just given. The two types of capsules are
essentially the same in structure save for the histological
details of the neck and the parietal lamina of the capsule. I n
the second type (in mice) the orifice from the capsule to the
uriniferous tubule is wider than usual, and the neck is entirely
absent. Instead of a n abrupt transition in this region from
squamous to cuboidal epithelium, the cuboidal cells of the
proximal convoluted tubule continue upwards into the capsule
to varying extents, replacing the usual squamous cells of the
parietal lamina. These cuboidal cells can frequently be found
encroaching on the equator of the capsule, and occasionally
extending as far a s the vascular pole. I n well-stained sections,
brush borders are sometimes visible. At other times the cell
outlines appear indistinct and irregular, with processes which
protrude into the lumen (see fig. 1B).
The observations of von Mollendorff ( '29), Rytand ( '38),
and Carpenter ('39) confirm this finding. I n 1935 Bemliolz
reported the existence of capsules of this type in a number
of different species of mammals. I n the rabbit, pig, horse,
guinea pig, fox and coon he observed no encroachment of
cuboidal cells on the capsule. I n the mouse, dog, mink and
woodchuck, however, he found large numbers of these modified
400
C H A R L O T T E CRABTREE
capsules which were particularly numerous in the kidneys
of four Pekinese dogs. He omitted mentioning tlie sex of the
animals studied. Selye ('39), in the paper already cited,
fouiid large numbers of this type of capsule in the hypertrophied kidneys of his testosterone-treated female mice.
Helmholz ('35) noted in a human kidney from an infant with
pyelonephyitis a similar continuation of cuboidal epithelium
into the parietal layer of the capsule. I n a second case, that
of a 9-year-old boy, he reported that some of the capsules
showed development of cuboidal cells in the parietal lamina,
while in others squamous cells similar to those of the typical
human capsule made up the first part of the proximal convoluted tubule. Risak ('28) reported the presence of cuboidal
cell capsules in an adult woman with chronic pyelonephritis,
renal calculus, and hydroneplirosis.
As an explanation of the occurrence of this type of mammalian capsule with high cuboidal cells in the parietal lamina,
Risak ('28) suggested that they might be the result of arrested development, but was forced t o reject the hypothesis.
Since the parietal layer is the first to flatten in the human (see
fig. 2 D ) arrested development would be characterized by the
presence of cuboidal cells in the visceral instead of the parietal
lamina. Since one can also find in sections of young mouse
kidney embryonic capsules with squamous cells composing
the parietal lamina but columiiar cells still adhering to the
glomernlar loops similar to those observed in the human, as
well as cuboidal cell capsules such as those originally described by Henda (188?), it does not seem probable that these
cuhoidal cell capsules represent arrested development in either
the human or in the mouse.
The findings of Bertelli ('30) are the converse of the results of the workers just cited: he claims to have observed
in the normal frog, salamander, guinea pig, and cat, capsnles
with the high columnar cells occurring more frequently in the
visceral than in the parietal layer. I have been unable to
confirm his observations in adult frogs or guinea pigs of
either sex.
STRUCTURE OF BOWMAN'S CAPSULE IN MICE
401
There is adequate evidence, a s indicated in this brief survey,
that three facts have been established concerning variations in
thc kidney in certain mammals: the kidneys of the two sexes
vary in weight ; weight variations can be produced experimentally by castration or by injection of sex hormones; two
types of Bowman's capsules occur normally. Indication that
a correlation exists between the age and sex of the animal and
the relative numbers of the two types of Bowman's capsules
liere described has been reported in a preliminary publication
(Crabtree, '40). Proof, based on diffcrciitial counts of the
two types of capsules in a series of kidneys of mice of known
age and sex, is here presented.
11. MATERIAL AND METHODS
Twenty mice from five diffei*ent litters and two separate
strains were used for this experiment : seven pairs were from
the mixed stock of the Smith College animal colony, a hybrid
stock exhibiting a large variety of external characteristics ;
three pairs were pure albino from the Carworth F a r m stock.
Males and females were placed in separate cages at the time
of weaning and maintained on the regular stock diet. I n
order to minimize the possibility of influencing genetic factors,
litter mates were always used f o r comparisons between sexes
of mice of any one age.
The animals were killed by a blow on the head, and the
kidneys removed immediately, washed in physiological salt
solution, fixed for 6 hours in Susa, and dehydrated in Dioxan.
Penetration of the fixative was insured by cutting the tissue into small pieces a t the end of the first hour of fixation.
After fixation the tissues were transferred for 24 hours
to Dioxan to which a little crystalized iodine had been added
to remove the mercuric chloride of the fixative. Both kidneys
of each animal were weighed together (correct to the
nearest milligram) 24 hours after killing. Following dehydration for several days in Dioxan, the tissues were embedded in 56'48" paraffin and sectioned a t 6 u. One kidney of
each animal was sectioned longitudinally and one transversely,
and some sections of each kidney were stained with iron
liaematoxylin and some with Mallory 's triple stain. One sec-
402
CHARLOTTE CRABTREE
tioii of each stain from each kidney, making four slides in all,
were selected for detailed study of each animal.
Immediately following removal of the kidneys to fixative
the remainder of the animal body was weighed directly, disregarding intestinal contents and blood loss. Thus the computed body weights and percentage weights are not accurate
beyond a ienth of a gram, but are included in the report
merely for comparison.
All the Bowman’s capsules in four slides from each animal
were counted and classified either as cuboidal o r “high” cell
capsules or as syuamous or “low” cell capsules, as described
above. Capsules were regarded as “high” if cuboidal cells
of the parietal lamina extended around some part of the
glomerulus o r if the cuboidal cells, in sections where the urine
pole was visible, were definitely seen to encroach on the
parietal lamina of the capsule. Capsules obviously embryonic,
i.e., in which the two layers had not reached adult form, were
disregarded.
111. RESULTS
Tlie following table summarizes the actual numerical results
obtained in counting and classifying the capsules in these
mouse kidneys.
The accuracy of this method is indicated by the uniform
results obtained. I n only one slide of one animal (D7 0 ) was a
percentage of low capsules found which varied more than
7% from the total average for that animal, o r which differed
more than 10% from a corresponding average for any other
single section from the same animal. As indicated in the above
chart, a further increase in accuracy was sought in some animals of representative ages by counting a larger number of
sections. Whenever serial sections were used, only every
fourth or more section was studied so that the same capsule
should not be counted twice.
Since a capsule with cuboidal cells extending only partly
around the parietal lamina can conceivably be cut in a number
of planes which do not include the cuboidal cells, the figures
quoted in the above table may be somewhat lower than the
TABLE 1
Resilts of chsificatiow of capsules
AGE I N
mRIES
NUMBER
WEEKS
D16 0
Birth
D21 J
Birth
D15 0
2
D14 2
2
D24 0
D25 8
3
3
D13 0
4.5
D12 8
4.5
D28 9
7
D29 d
7
D60
9.5
s
9.5
D9
D26 0
D27 8
10.5
10.5
D70
11.5
D11 d
11.5
D8?
20
D10 8
20
s. Mal.
No. low cap.
Total no. cap.
yolow cap.
No. low cap.
Total no. cap.
yolow cap.
No. low cap.
Total no. cap.
yolow cap.
No. low cap.
Total no. cap.
yolow cap.
No. low cap.
Total no. cap.
yolow cap.
No. low cap.
Total no. cap.
yolow cap.
No. low cap.
Total no. cap.
% low cap.
No. low cap.
Total no. cap.
yolow cap.
No. low cap.
Total no. cap.
% low cap.
No. low cap.
Total no. cap.
% low cap.
No. low cap.
Total no. cap.
% low cap.
No. low cap.
Total no. cap.
Yo low cap.
No. low cap.
Total no. cap.
yo low cap.
No. low cap.
Total no. cap.
70low cap.
No. low cap.
Total no. cap.
% low cap.
No. low cap.
Total no. cap.
yolow cap.
No. low cap.
Total no. cap.
% low cap.
No. low cap.
Total no. cap.
% low cap.
xs. Mal.
Is. FeH.
xs. FeH.
TOTALS
85%
Figure based on a count of
every 4th slide of a serial
91%
83%
83%
93%
99
82%
87%
59
90%
27
82%
Figure based on a count of
30 sections from a serial
Figure based on a count of
10 sections from a serial
56
59
38
55
82
54
79
79
70%
75%
68%
70%
37
53
31
50
53
88
78
49
64%
70%
60%
63%
56
76
103
65
73
137
91
106
77%
72 %
75%
71%
17
14
24
27
70
109
57
97
25%
24%
25%
25%
73
50
57
87
113
74
85
127
6996
6770
65%
68%
30
20
18
19
115
79
65
77
23%
26%
29%
25%
Figure based on a count of
1 2 serial sections
Figure based on a count of
12 serial section
40
31
85
107
37%
36%
29
13
84
196
15%
15%
42
36
94
88
45yo
41%
9
6
84
174
11%
3%
403
8770
114
131
87%
189
223
85%
1343
1603
84%
754
819
92 70
208
294
71%
171
270
63%
300
40 7
74yo
82
333
25%
267
399
67%
87
336
26%
967
1396
7070
216
1086
20%
124
385
32 %
80
513
16%
136
347
39%
31
503
6%
404
CHARLOTTE CRABTREE
actual condition warrants. This possibility is equally present
in all the sections examined, but for this reason the figures
must be regarded not as absolute values, but as denoting a
general trend toward a specific condition.
There is little doubt that the capsules of both types are
fairly equally distributed throughout the cortex ; since, in
cross, longitudinal, and serial sections, uniform results were
obtained.
The following table correlates the numerical data from
table 1with other data derived from the study of these animals.
The weight ratios obtained in this study are in general
agreement with results of other workers already cited. Although in prepnbertal groups the kidneys of the female mouse
may be slightly larger than those of the male of correspondTABLE 2
Summary of rrsults of the study of normal kirln~ysin mice
SERIES
NUMBER
SEX
D16 0
D21dz
D15 0
D14 d
D24ly2
D25ldZ
D13 ?!
D12 6
D2S19
D29 *
D60
D9
D26
D27
s
0
8
D7P
nll d
D80
n1o 6
'ER CENT WEIGHT
AGE I N
WEEKS
KIDNCY
\\EIGHT
Birth
Birth
1.1
3
0.0151
0.0149
0.0582
0.0929
0.1929
0.1773
4.5
4.5
0.0881
0.0828
1.6
7
7
9.5
9.5
10.5
10.5
11.5
11.5
20
30
0.3198
0.5130
...
0.3049
0.3499
0.3628
0.3829
1.5
......
...
0.3276
0.4423
1.5
2
2
3
......
PER CENT HIGH
PER CENT L O W
P
87
1.1
85
1.3
92
63
25
26
1.6
20
16
Carworth Farm stock.
Figures based on counts of serial sections.
6
S T R U C T U R E O F BOWMAN’S
CAPSULE I N MICE
405
ing age, in the older, mature animals the males’ kidney weight
exceeds that of the females’. Except for the pairs killed at
birth and at 10; weeks, the kidney in the male always constitutes a greater percentage of the total body weight than
in the female.
The relative numbers of the two types of capsules form a
striking and significant unit of measurement f o r age and sex.
I n young mice, 43 weeks or less in age, the percentage of
cuboidal cell capsules is uniformly low in both sexes. With
the onset of sexual maturity, which occurs somewhere between the fifth and sixth weeks in these strains of mice, a
marked change is noted. I n the female the percentage of
cuboidal cell capsules increases steadily but slowly until by
114 weeks more than half of the total number of capsules
may be included in the high cell group. The greatest increase
(50%) occurs between lo& and 11; weeks. I n the male, however, the increase is far more rapid and extensive. From 43
weeks through the twentieth week the male kidney always contains a higher percentage of cuboidal cell capsules than the
kidney of a female litter mate. At the seventh week a large
increase of high cell capsules suddenly occurs in the male;
the kidney attains at this time a percentage of cuboidal cell
capsules higher than is ever reached by the female. The
relative numbers of cuboidal cell capsules continues to increase in the male until by the twentieth week 94% of all the
capsules observed have some cuboidal cells in the parietal
lamina. As already pointed out, the remaining 6% may have
contained cuboidal cells, but were cut so that none of these
cells were included in the section.
Thus the presence of cuboidal cells in the parietal lamina is
a normal occurrence in mice. The relative numbers of the
two types of capsules is dependent on the age and sex of the
animal. Renal sexual differentiation occurs earlier in the
male than in the female, but in both sexes the percentage of
cuboidal cells in the capsules tends to increase with age. The
postpubertal male kidney, however, always has a higher per-
406
CHARLOTTE CRABTREE
centage of cuboidal cell capsules than does the female of
corresponding age.
IV. DISCUSSION
The exact nature of the cells which invade the parietal
lamina is still not determined. Gorer (’40) listed these cells
as characteristic of one of three types of renal lesions common
to mice, but the above results indicate that, on the contrary,
their occurrence is normal to the species. His statement:
“None of the lesions appear to occur more frequently in either
sex” is probably not based on statistical methods of examination. His observations of their frequent occurrence “in old
members of the CBA strain” is in agreement with my findings.
I n appearance the cuboidal cells of the capsules are continuous with and identical with the cells of the proximal convoluted tubule ; the cytoplasm is granular in appearance with
brush borders usually demonstrable ; frequently basal granules can be seen. Pfeiffer, Emmel and Gardner (’40) state
that mitochondria of the capsular cells are similar in structure and in intracellular location t o the mitochondria of the
tubule cells. Still further evidence of the structural and
physiological similarity of the cuboidal cells of the capsule
and those making up the proximal convoluted tubule can be
established by the use of vital staining. Trypan blue granules
are characteristically collected and stored in the form of
large crystals by the cuboidal cells of the proximal end of the
proximal convoluted tubule. The kidney of a young mature
male mouse (exact age unknown) treated with trypan blue
was found t o exhibit characteristic storage of the large blue
granules in the cells of the proximal convoluted tubule as
expected, but also storage of stain in the cuboidal cells of
the capsule whenever these cells were present (see fig. 3 ) .
‘ F o r 3 days the mnuse was given daily intraperitoneal injertions of about
1.5 cc. of a 1% trypan blue solution made up in 0.8% NaC1. A t the end of the
third day the animal was killed, and the kidneys removed, preserved, and sectioned
in the same manner as the others of this series. Mayer’s Paracarmine in 70%
alcohol was used a8 a rounterstain, thus avoiding the use of water which tends
to dissolve the trypan blue.
STRUCTURE
OF BOWMAN’S
CAPSULE IN MICE
407
These granules were distributed in the cytoplasm around the
nucleus in a n identical manner by both capsular and tubular
cells. Save for an occasional granule in a macrophage either
of the interstitial tissue of the medulla or in the connective
tissue around a blood vessel, trypan blue crystals were found
nowhere else in the kidney, indicating that this specific storage
ability is characteristic only of this one type of renal epithelium. It would seem evident, therefore, that cuboidal cells
of the capsule, if present, a r e identical with those of the
.05rna
Fig. 3 Camera lucida drawing showing the deposition of trypan blue granules
in the cuboidal cells of the Bowman’s capsule and of the proximal cnd of the
proximal convoluted tubule.
proximal end of the proximal convoluted tubule. As the
animal grows older the total number of capsules increases, but
this increase is not in proportion to the rapid change in the
percentage of high cell capsules. Thus in a single capsule a
change from the squarvious to the cuboidal type is inferred, but
there is no direct evidence t o prove whether the cuboidal cells
arise from migration of tubular cells or metaplasia and differentiation of preexisting squanious epitlielium of the parietal
lamina.
408
CHARLOTTE CRABTREE
An interesting and probably significant correlation between
the development of the mouse kidney and of the adjacent
adrenal gland also exists. The origin, development, and subsequent degeneration of the androgenic or x-zone of the mouse
adrenal has occupied the attention of a number of workers
TABLE 3
Coniparison of the decelopment of the x-zone of the adrenal gland witA the number
of cuboidal cell capsules in t h e kidney
DIALE
.4GE
1-3
weeks
4-5
weeks
7-8
weeks
10-12
weeks
I
FEMALE
x-zone of tlie
adrenal gland
Early developmentSame as in f e
of x-zone. Same male (8% t o
as in fcmale
15%)
X-zone begins t o Number begins t c
atrophy
increase (37%)
X-zone entirely
75% high capsules
gone
.?yo to 10% in
No change
crease (80% tc
84%)
No. of high re11 r a p -
sules in the kidney
Early develop- Same a s in male
ment of x-zone. (13% t o 15%)
Same as in malc
X-zone continues Slight increase in
to develop
number (29%)
No change
No change
X-zone degener- Peak number of
ates
female series
(68%) reachcd
for the last 10 years. Table 3 compares the times of appearance and disappearance of the x-zone a s noted by
Dearisley (’28) with the times of sudden increase of cuboidal
cell capsules of normal male and female kidneys.
During the first 3 weeks of postnatal life both the kidneys and
the adrenal glands of the two sexes a r e essentially alike.
During the fifth week the x-zone of the male adrenal begins to
Figures 4 t o 7
Photomicrographs of various capsules from mouse kidneys.
Fig. 4 Capsule stained with Mallory’s triple stain t o show brush borders
of the cuboidal cells of the capsule. Male mouse, 9.3 weeks old. X 350.
Fig. 5 A “low” cell capsule with sqnamous cells making u p the entire parietal
layer. Male mouse, 7 weeks. Fc. Haem. X 3.50.
Fig. 6 A “high” cell capsules showing the continuity of the cuboidal cells
with those of the adjoining proximal convoluted tubule. Adult male mouse. Fe.
Haem. X 350.
Fig. 7 A and B. Cuboidal cell capsules from a 7 weeks old male mouse. C.
Similar capsule from the kidney of an adult male mouse. (a) Pen and ink schema
of A. showing the extent of cytoplasm of the capsular ruhoidal cells. Fe. Haem.
X 350.
410
CHARLOTTE CRABTREE
degenerate, arid a t tlie same time the number of cuboidal cell
capsules increases 29% as compared with a 13% increase of
the female during the same pci*iod. By tlie end of the seveiith
week, when degeneration of the male x-zone is nearing completion, the percentage of cuboidal cell capsules has made
a n additional increase of 38% and is approaching the adult
limit. I n the female the x-zone persists for 10 to 12 weeks,
and concurrently the percentage of cuboidal cell capsules
iaemains low until betweeii 10; and 11i weeks, when the relative
numbers of this type of capsule jumps abruptly from 30% to
68%. I n both sexes, then, the number of cuboidal cell capsules
of the noi*mal kidiiey remains low until the x-zone has disappeared, arid then rises abruptly to adult level.
The above correlation in development is not surprising.
Functional correlatioii between the adrenal and the gonads lias
been established for some time, although the exact nature
of the relationship is still f a r from being conipletely understood. I n view of this, and also in view of the embryological
origin and anatomical relationships betweeii the adrenal, the
genital, and tlie urinary systems, a direct correlatioii between
tlie development of the adrerial tmcl the kidney would seem
theoretically logical.
I n the mouse kidney the relative number of capsules coiitaiiiiiig cuboidal cells in the parietal lamina varies with the
age and sex of the animal, increases a t puberty, arid is at all
times thereafter higher in the male than in the female of
corresponding age. The rate of increase of the euboidal cell
capsules is closely correlated with the development of the
adrenal gland. These two facts point toward an endocrinological causative agent producing hypertrophy of the llarietal,
but not the visceral lamina of Bowman’s capsule. Pfeiffer et 31.
( ’40) have injected estradiol benzoate, est~.ac’lioldipropioiiate,
testosterone, and testosterone propionate into normal male
mice, and Selye (’39) used testosterone propionate and cholesterol in normal females. Both obtained marked renal hypertrophy a i d noted particularly hypertrophp of the cells of the
parietal lamina of Bowman’s capsule. A further study is
STRUCTURE OF BOWMAN’S
CAPSULE IN MICE
411
indicated to compare with the normals just established the
effects on the kidney of castration and the injection of sex
hormones. I have already collected data on this problem, and
will publish it in a second communication.
V. CONCLUSIONS
1. I n some species of mammals, including mice, the parietal
layer of Bowman’s capsule normally may be partly o r completely composed of cuboidal cells instead of the more usual
squamous epithelium.
2. Capsules of this type are not the product of arrested
development.
3. Cuboidal cells in these capsules appear to be identical with
those of the proximal convoluted tubule, and both exhibit the
highly specialized ability to store granules of trypan blue
vital stain, There is no direct evidence, however, to prove
whether the cuboidal cells of the capsule arise as a result of
migration of tubular cells or of metaplasia of preexisting
squamous capsular cells.
4. Indirect evidence indicates that single capsules caii
change from the low cell type to the high cell type.
5 . Cuboidal cell capsules are normally present in both sexes
in mice of all ages studied.
6. After puberty the percentage of cuboidal cell capsules
begins to increase. I n the male this occurs earlier than in the
female, and the rate of increase is f a r greater-tending to
approach 100% while that of the female of corresponding age
always remains decidedly lower.
7. I n both sexes the relative numbers of cuboidal cell
capsules remains low until degeneration of the x-zone of the
adrenal gland has occurred, and then rises rapidly t o the adult
level. This occurs at about 7 weeks of age in the male and
about 11weeks in the female.
8. It is suggested that the increase of cuboidal cells in capsules is a specific response of the mouse kidney to an endocrine
factor or factors.
412
CHARLOTTE CRABTREE
9. The numerical relationship between high cell and low cell
capsules appears t o be a reliable index of age and sex changes
in mice.
The author wishes to take this opportunity to express her
sincere appreciation to Dr. Myra M. Sampson f o r her helpful
suggestions concerning the foregoing work.
LITERATURE CITED
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R. R., AND R. D. BENSLEY 1930 The structure of the renal corpuscle.
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BERTELLI,
R. 1930 Sulla Presenza di Cellule Alte nel Foglietto Viscerale della
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CARPENTER,
E. C‘. 1939 Personal communications.
BENDA,C.
CRABTREE,C. E. 1940 Sex differences i n thc structure of Bowman’s capsule
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DEANSLEY,
R. 1928 A study of the adrenal cortex in the mouse and its relationship t o the gonads. Proc. Roy. Soc., vol. 103, p. 523.
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Baet., vol. 50, p. 25.
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P., AND H. POPPER1940 The histogenesis and physiology of the
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H. 1935 The presence of tubular epithelium within the glomerular
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STRUCTURE OF BOWMAN’S CAPSULE I N MICE
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RANDRATH,E. 1932 Zur normalen und pathologisehens Anatomie der Deckzellen des Nierenkorperchens. Zeit. f. Zell. u. mik. Anat., vol. 15, p. 182.
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MOLLENDORFF,
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