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Nuclear morphology according to sex in Macacus rhesus.

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NUCLEAR SIORPHOLOGP, ACCORDISG TO SEX,
I N IfACACUS RHESUS
EIGHTEEN FIGURES
INTRODUCTIOK
A distinction between iritcrniitotic nuclei of inales and fenialcs has been described in several species of the orders
Carnivora and Artiodactyla and, more recently, in man ( R a r r ,
Bertram and Lindsay, '50 ; Graham and Ran-, '52 ; Moore and
Barr, '53, '54). In brief, a special mass of chromatin, the sex
chromatin, is clearly visible only in females. Several lines of
indirect evidence suggest that the sex chromatin of female
cells represents heterochromatic portions of the XX sex
chr orno soine complex.
The present investigation consists of a comparison of
nuclei in inales arid females in a primate, N n c a c u s rhesus. It
will be shown that cell nuclei of the monlcey a r e similar to
those of such species of the orders C'arnivora and Artiodactyla
a s have been studied, and to nuclei of the human, in that a
sex difference in nuclear structure is clearly demonstrable.
They differ from nuclei of rodents where the neurons (these
cells alone have been studied in rodents) have multiple
chromatin inasses which obscure any sex difference which
may be present.
This work mas made yossiblc by grants-iii aid from K:itional JIealth Grants
(Mental IIealtli Pivision) of Canada a n d tlic National Research Couiiril of Ca11:ida.
Medical Fellow, Xational Research Council.
153
154
R. H. P R I N C E , M. A. G R A H A M A N D M. L. RARR
MATERIALS AND METHODS
Three male arid three fernale monkeys, all mature, were
used in this study. The brains were fixed by perfusion through
thc carotid arteries with a solution of 10% fornialin in normal
saline, after preliminary anesthesia with Nembutal. Blocks
which included the regions of the nervous system listed in
table 1 were fixed for a further period in isotonic 10% fornialiii, eniloedded in paraffiiri and sectioned at 10 p. Sections
stained with cresyl violet were used for the major portion
of the study. Acidification of the dye solution to approximately
pH 4 n’as found to be an advantage. V i t h this modification,
the chromatin is stained more darkly than the nucleolus and
the Nissl niaterial, and identification of the sex chromatin is
thereby facilitated. Sections were also examined after staining with methyl green-pyronin arid by the Feulgen method.
The noii-nervous tissues listed in table 2 were studied in
female monkeys. Corresponding tissues were studied in male
monkeys except for the substitutioii of the following elements
of the niale reproductive system - prostatic epithelium, Sertoli cells and interstitial cellh of the testis. Blocks were fixed
in a forniol-acetic-alcohol mixture for the niost part, limited
use being made of Zenlier’s fluid. Paraffin sections were cut
a t 5-7 cc and stained with IIari-is’ liematoxyliri and eosin,
cresyl violet, cresyl echt violet and hy the Feulgen method.
F o r the various regions of the nervous system in each
. .
aninial, 200 nuclei contmiiiiig a nucleolus were examined with
a Leitz fluorite oil inirnei*sionobjective, N. A. 1.32. T h e n the
nucleus contained a mass of sex chromatin in the plane of
the section, it was recorded as lying adjacent to the nucleolus,
frecx in the nucleoplasni, or adjacent to the nuclear membrane.
The WX chromatin of a cwnsidernble piwportion of f erriale
cells was located between the nuclear membrane arid an eccentric nucleolus, touching on both. Such sex chromatin masses
wcre recorded a s being adjacent to the nuclear membrane.
hleasureiiieiits of the sex chromatin were niade a s notcd in
the following section. Kuclei in other tissues were studied in
a similar manner using, for the more intensive work, sections
stained with heniatoxylin and eosin.
OGSERVATlOXS
12'c~eou.s tissue
1. Female (table 1;figures 1, 3, 5, 7, 9 and 11)
The characteristic feature of female nuclei throughout the
nervous systeni is the presence of the sex chromatin. Considering the various regions together, sex chromatin is encountered in i 6 % of 1 0 thick
~
sections through neurons,
when each cell section contains thc nucleolus. A s shown in
table 1 and illustrated by the figures, the sex chromatin is
more likely to be a t the nuclear memhrane than in any other
position. However, its location varies from one type of neuron
to another, this feature lwing consistent in the three females.
F o r example, in such regions as thc putamen, caudate nucleus,
subthalamic nucleus, inferior olirary nucleus, Purkiiije cells,
and others, the s e s chromatin is usually at the nuclear membrane, while in the motor nuclei of the oculoniotor, trochlear,
facial and hypoglossal nerves i t is more likely to be found
adjacent t o the nucleolus. Further, the sex chromatin is commonly a t the nucleolus in giant pyramidal 01' Retz cells, as
illustrated in figure 5, while it is inore likely to be situated
a t the nuclear nienihimie in the sniallclr neurons of the motor
cortex. Sex chroniatin n i n q w s are rarely seen free in the
nucleoplasni.
The real incidence of sex chromatin in ncrvc~cells of the
female moiikcy must be higher than that shown in table 1,
since tlie section may iiot include tlie whole thickness of the
iiucaleus. The sex chroniatin will be excluded from the plane
of the sectioii in a n appreciable nurribcr of these cells, especially in regions wherc it is commonly located a t the
nuclear nieriibrane. Cell sections without c2 nucleolus but with
the sex chromatin at the nuclear membrane arc, in fact, encountered frequently.
156
B. H. PRINCE, M. A. GRAHAM A N D M. L. BARR
TABLE 1
Incidence, position and size of the sex claromatin in nervous system of the monkey
(1) adjacent to nucleolus ( 2 ) free in nucleoplasm
( 3 ) adjacent t o nuclear incmbrane
(average of tlirec animals of each sex)
REGION OB
NERVOUS
SYSTEM
SEX
P O S I T I O N O F SEX
CHROMATIN ( % )
SIZE O F F E M A L E
SEX C H R O M A T I N
(1)
(2)
(3)
15
1
1
2
70
2
86
d
13
1
0
1
72
2
?
d
21
2
1
1
61
0
3
8
2
0
P
(%)
Occipital cortex
?
s
Parietal cortex
Motor cortex
Frontal cortex
Teinportnl cortex
0
s
(P)
0.8
x
1.1
85
4
0.8
x
1.1
83
10
0.8 X 1.1
7
1
1
79
1
88
4
0.8 X 1.1
6
I
0
2
72
2
78
5
0.7 X 1.1
52
3
58
5
0.8
x
1.1
x
1.0
5
Jfitral cells,
olfactory bulb
s
5
1
1
Pyramidal cells
of hippocampus
0
d
2
0
1
1
51
54
2
0.7
1
Polymorphic cells
of fascia deiitata
0
3
5
1
0
0
47
1
52
2
0.6 X 1.1
Caudate nucleus
0
3
2
1
2
0
74
6
78
7
0.7 X 1.1
Putamen
0
3
4
2
2
3
71
6
77
11
0.7 X 1.1
Globus pallidus
P
3
18
2
2
54
1
71
0.8 X 1.2
1
?
13
3
1
1
68
2
82
6
0.9
x
1.2
24
1
1
61
1
86
0.8
x
1.1
71
3
88
0.8
x
1.0
Medial thalainic
nuclcus
3
L a t m r l thalamic
nucleus
3?
I,att,ral geniculate
bodp
?
3
16
0
1
1
1
2
4
3
3
157
SEX CHROMATIN I N MACACUS RHESUS
TABLE 1 (continued)
-REGION O F
NERVOUS
SYSTEM
SEX
CHROMATIN ( 7 c )
POSITION O F SEX
SIZE O F FEMALE
SEX CHROMATIN
(2)
(3)
3
1
2
0
68
3
73
4
0.8 X 1.2
0
9
4
1
1
68
6
78
11
0.8 X 1.1
0
15
2
2
0
62
1
79
3
1.0
x
1.2
0
44
2
0
0
35
3
79
5
0.9
x
1.2
0
41
1
2
36
3
79
5
0.8 X 1.1
1
0
43
2
2
0
28
1
73
3
0.8 X 1.1
0
1
0
0
1
71
2
72
3
0.9
0
12
2
1
2
66
2
79
6
0.8 X 1.1
0
63
0
1
1
15
0
79
1
0.9
0
(1)
-
(FL)
(%o)
Subthalamic
nucleus
Tectum of midbrain
Substantia nigra
Oculomotor nucleus
Trochlear nucleus
Facial nucleus
Suclei pontis
0
6
s
s
s
s
s
s
x
1.2
Vestibular
nucleus
s
Hypoglossal
nucleus
s
Inferior olivary
nucleus
3
1
0
1
70
1
73
3
0.8 X 1.1
d
Reticular formation
of brain stem
s
P
16
1
2
58
2
75
8
0.8 X 1.2
4
P
12
3
3
1
64
5
79
9
0.9
x
1.2
0
2
3
3
3
69
3
74
0.9
x
1.1
d
Anterior horn,
cervical cord
P
d
26
2
2
1
51
2
79
5
0.9
x
1.2
Posterior root
ganglion, cervical
s
0
22
2
3
2
44
2
69
6
0.9
x
1.2
Dentate nucleus
Furkinje cells
s
x
1.1
9
158
R. I€. PRINCE, ill. A. G R A H A M
AND
ni.
L. BARR
I n a considerable proportion of cells, varying horn one region to another. the sex chroniatin is interposed between a n
eccentric nucleolus and the iiuclear membrane. This is especially characteristic of neurons in the pyramidal cell layer
of the hippocampus aiid the polymorphic cell layer of the
fascia dentata. The sex chroniatin is rather more difficult to identify under these conditions. This factor, together with the slightly smaller dimensions of the sex chromatin in the archipallium, compared with many other regions
(see below), accounts for the relatively low percentage of
cells in which it could be identified in the hippocampus and
fascia dentata.
The data in table 1 for the incidence of sex chromatin a t
the nuclear memhrane a r e considered to be reasonably accurate, since only rarely is there a comparable mass of
chromatin in male nuclei. Similarly, a mass of sex chroniatin
a t the nucleolus can be identified with confidence in large
neuroiis where other chroniatin particles a r e inconspicuous.
However, chromatin masses of variable numbcr and size
occur, usually in contact with tlie nucleolus, in the smaller
neurons of males aiid females. These a r e the paranucleolar
bodies to which Coidan ('52) drew attention i n spinal cord
neurons of the male monkey. They do not interfere seriou4y
with identification of the sex chromatin since puranucleolar
bodies a r e inconspicuous in large neurons throughout the
nervous system i n preparations which are good technically,
and since tlie sex chromatin is usually at the nuclear iiienibrane in this species.
Very small neurons such a s the granule cells of the cercbellar cortex, and neuroglial cells, have so many large cliromatin masses i n both sexes that such sex difference in their
morphology as may be present is largely obscured.
The sex chromatin is usually planoconvex, less frequently
triangular or some other shape, 117lien it is at the nuclear
membrane. It is irregular, round or planoconves in outline
when situated next to the nuclcolus. I n general, the sex
chromatin tends to conform in shape, or to be niouldcd
SEX C H R O N A T I N I K MACACUS RHESUS
159
against, an adjacent structure. A small clear area is seen in
a considerable proportion of sex chromatin masses.
Although it is difficult to obtain precise measurements f o r
such a small object, an attempt was made t o obtain size values
by measuring the least and the greatest diameters with a
filar micrometer ocular. The mean values of 30 sex chromatin
masses in each region of the nervous system are recorded in
table 1. Consideriiig the various regions together, the average
size is 0.8 p x 1.1p. TVe gained the impression, prior to
making the measurements, that the sex chromatin varies in
size from one region to another within relatively narrow
limits. This impression is borne out by the figures obtained
by measurements. The sex chromatin seems t o be largest
in the substantia nigra ( 1 . 0 ~x 1 . 2 ~ )and smallest in the
pyramidal cell layer of the hippocampus (0.7 p X1.0 p ) and
the polymorphic cell layer of the fascia dentata (0.6 p x 1.1p).
The sex chromatin is consistently Feulgen-positive and has
a strong affinity for methyl green. These reactions indicate
that the sex chromatin, like other chromatin and the chromosomes in general, contains desoxyribose nucleic acid.
2. Male (table 1; figures 2, 4, 6, 8, 10 and 12)
A mass of chromatin comparable in size, position, shape,
and internal structure with the sex chromatin of female nuclei
is seldom encountered in male nuclei. Table 1 shows the
incidence of chromatin masses in nerve cell nuclei of males
that may correspond with the sex chromatin of females. I n
general, less than 10% of male nuclei contain such chromatin
particles, and it is uncertain whether these represent a special
mass of chromatin derived from the sex chromosomes or
whether they are of autosomal origin.
Noit-tzerrmts t i s s u e s
The nuclei of the various tissues differ in their suitability
f o r this work, depending on the amount and coarseness of
the general particulate chromatin. Both male and female
160
R. H. PRINCE, M. A. GRAHAM A N D M. L. BARR
nuclei contain one to several nucleoli as well as perinucleolar
chromatin and varying numbers of chromatin particles scattered through the nucleoplasm. I n general, we find that the
nuclear chromatin is coarser and more abundant in the monkey, as compared with the cat (Graham and Barr, '52) and
man (Moore and Barr, '54).
1. Female (table 2 ; figures 13, 15 and 17)
The sex chromatin is particularly clear in the transitional
epithelium of the urinary bladder, where the general chromatin
is very fine. There are numerous, rather large, chromatin
particles in the pancreatic acini, uriniferous tubules, striated
muscle and germinal centers of lymph nodes. Considerable
experience is required to identify the sex chromatin in these
locations and in many nuclei it is impossible to do so. The
sex chromatin can be identified with considerable confidence
in nuclei of the other cell types listed in table 2.
Considering the various tissues together, the sex chromatin
can be identified in about 80% of female nuclei. It is generally
adjacent to the nuclear membrane, except for the transitional
epithelium of the bladder. Its location here is as follows
(average of three females) - adjacent to the nucleolus 41%,
free in the nucleoplasm 47'0, and adjacent to the nuclear
membrane 4470. The sex chromatin is typically planoconvex
in outline and frequently has a small clear area in its center.
As in the nervous system, it is Feulgen-positive and stains
with methyl green.
Thirty sex chromatin masses were measured in each location; the mean figures are recorded in table 2. The average
size of the sex chromatin f o r all the tissues is 0.8 p X 1.1p,
the same value that was obtained for the average size of
the sex chromatin in nerve cells. There is a variation in
size from one type of cell to another within relatively narrow
limits. The sex chromatin appears to be largest in cartilage
cells (1.0 p x 1.3 p ) and smallest in the surface epithelium of
the gastric mucosa (0.7 p x 1.0 p ) and in smooth muscle
74
82
i3
82
74
69
74
74
90
Adrenal cortex
Aclrenal niedulla
Islets of Tdnngerhans
Surface epithelium,
gastric niucosa
Parietal cclls,
gastric glands
Surface epithelium,
jejunal niucosa
Paiicrratic aciiii
T,iver
Renal convoluted
tubules
Epitlrcliuin of
urinary bladder
IXCIDENCE O F
SEX CHltOMATIN
(70)
86
TISSUE
1.1
1.2
x
x
0.8
0.8
0.9
0.8
0.8
0.8
1.2
x
x
1.1
1.2
1.0
x
x
1.2
1.1
x
x
0.7 X 1.0
1.0
0.8
0.9 X 1.3
(P)
SIZE
Germinal center,
lymph node
Cartilage
Epidermis
Smooth muscle,
stomach
Cardiac muscle
Skeletal muscle
Vaginal epithelium
Uterine glands
Ovarian stromal cells
Ovarian follieular
cells
TISSUE
58
86
68
84
70
66
84
91
86
x
x
x
x
1.1
1.1
1.1
1.2
1.2
x
1.1
- a
0.8
1.0 X 1.3
0.8
CI
CI
x
w
0
s
a
$
1.1
rn
rn
cl
x
0
!p-4
2
2
z
O
F
E
0
x
m
rn
0.8 X 0.9
0.8
0.9 X 1.3
0.8
0.8
0.8
0.9
(P)
(%)
83
SIZE
IXCIDENCE O F
SEX CHROMATIN
Incidence and size of the sex chromatin in tissues of the female monkey
TABLE 2
162
R. H. PRINCE, M. A. G R A H A M AND M. L. BARE
cells (0.8 p X 0.9 p ) . The sex chroniatin is large and easy t o
identify in tlie adrenal cortex and islets of Laiigerhans. These
tissues (and cartilage cells) a r e also among the most suitable
for identification of the female sex chromatin in the cat and
human.
2. Males (figs. 14, 16 and 18)
The nuclei of niale tissues contain multiple particles of
chromatin, varying i n number arid size from one cell type
to another. It is impossible to distinguish a very small particle
of sex chromatin, such a s might he expected to occur in male
iiuclei, from these general chromatin particles. Ilowever,
it is certain that only a very few male nuclei contain a chromatin niass with characteristics like those of the sex chromatin
of female nuclei.
n'ith respect to the noii-nervous tissues of the monkey it
can be stated that a careful comparison between nuclei of
inales and feinales denionstrates the practicability of distinguishing the sex chromatin of females from other chromatin
particles mid from nucleoli. The distinction is based on the
tyipical size, shape, position and staining properties of the
sex chromatin and the niiiiute pale area which it often contains.
111RCTTSS I ON
S e r v e cells of the monkey h a w a sex difference in nuclear
morphology which is similar t o that found in the cat (Rarr
et al., 'SO), and i n the dog, mink, marten, ferret, racoon,
skuiik, goat and deer ( X o o r e and Barr, '53). The sex difference is perfectly clear in the larger neui-0x1s of the monkey,
where the general chromatin particles a r e inconspicuous, hut
is ol)scured to some extent in srnall neurons wliere the chromatin is clumped in larger niwsscs. The iieuroiis of the nlonli-ey
m e also comparable to those of tlie human frontal cortex
and sympathetic ganglia, which have been shown by hlylle
and Graham ('54) to liave a characteristic moq)liologp according t o sex. The nerve cells of all of tliesc species differ
from those of the guinea pig, rat, mouse, hamster, groundhog
SEX CHROMATIN I N MACACUS RHESUS
163
and the rabbit. The nerve cells of rodents and lagomorphs
contain multiple masses of basophilic material next to the
nucleolus i n both sexes, and the n u d e a r morphology is complex (Moore and Barr, '53). The nuclei of non-nervous
tissues have been investigated, with respect to their structure
according to sex, in the cat (Graham and Rarr, '52) and man
(Moore, Graham and Harr, '53; Davidson and Smith, '54;
Emery and Afchfillan, '54 ; Marberger and Nelson, '54 ; Moore
and Barr, '54), and a clear sex difference has been established in both species. The nuclei of non-nervous tissues
of the monkey a r e comparable to those of the cat and man
in this respect, although the chromatin particles are coarser
and tissues of the monkey a r e less satisfactory for such a
study on this account.
The position of the sex chromatin is similar i n non-nervous
tissues of monkey, cat and man, where it is usually found
adherent to the inner surface of the nuclear membrane. There
is, however, a species difference in the position of the sex
chromatin of neurons. F o r example, in carnivores the sex
chromatin is most likely to be found adjacent to the nucleolus ;
this juxtanucleolar position is perhaps rather less constant in
the goat and deer; while throughout the nervous system
generally of the monkey and in the frontal cortex and sympathetic ganglia of man the sex chromatin is located at the
nuclear membrane in a large proportion of cells. There is
also a variation in the position of the sex chromatin from one
type of neuron to another within the species.
These variations in the position of the sex chromatin
according to the species and type of cell a r e difficult to
interpret since so little is known concerning the forces which
operate within the nucleus. It may be pertinent that the
position of the sex chromatin is altered in chromatolytic
neurons ( B a n and Bertram, '51 ; Crouch and Rarr, '<54;
Lindsay and Rarr, '55). Further, the position of the sex
chromatin changes during development in the cat. The sex
chromatin of neuroblasts and neurons is usually located a t
the nuclear membrane during early embryonic stages. T h e w
164
R. H. PRINCE, M. A. GRAHAM A K D &I. L. BARR
is a gradual change as development proceeds, until the juxtanucleolar position characteristic of neurons of mature cats
is attained i n most regions shortly after birth. A less marked
shift in the reverse direction occurs in other tissues during
development of the cat embryo (Graham, '54). These observations suggest that metabolic factors influence the position of the sex chromatin within the nucleus. Hence, some
variations in cellular metabolism, depending on the species
and type of cell, may be responsible for the normal variations
which have been observed.
No new information has become available to modify the
suggestion concerning the origin of the sex chromatin, which
was offered in earlier papers from this laboratory. Painter
( '22, '24) established the chromosome number in Mncacus
rhesus a s 48, and showed that the female has X X sex chromosomes and the male X P sex chromosomes. H e described the
X chromosome a s a rod shaped structure and the T chromosome a s a small round body in the primary spermatocyte, the
XY complex having the appearance of an exclamation mark.
Similar morphological characteristics were described by Painter for the X and Y chromosomes of cells undergoing mitosis
in a male embryo, where the dot-like Y chromosome is about
one-sixth a s long as the X chromosome. Tissue culture studies
such as those of Hsu, Hooks and Pomerat ( '53) on human chromosomes may provide information a s to the fate of the sex
chromosomes when the nucleus enters the interini totic phase.
Until further evidence is available, it is inferred that the
sex chromatin of female cells represents adherent heterochromatic portions of the two X chromosomes. It must be
assumed then, that the X P complex of male cells fails to form
a chromatin mass of comparable size, possibly due in p a r t
to the small size of the Y chromosome, relative to its heterologous partner.
SUMNARY
1. Nuclei of various tissues of ilfacncus rhesus have a di
stinctive morpholo,gy, according to sex. I n females, the nuclei
SEX CHROMA'CIN I N MACACUS FLHESUS
165
contain a special mass of chromatin, the sex chromatin, which
is usually located at the nuclear menibram; a comparable
chroiiiatiii mass is seldoni seen in the nuclei of males.
2. The distinction, according to sex, is easy to recognize
in such cclls a s the larger neurons that have large vesicular
nuclei. It is difficult o r irripossihlc to identify the female sex
chromatin in cclls that have small, more pyknotic nuclei,
such as occur i n the pancreatic acirii and especially the neuroglial elements of the central neivous system. However, a
sex difference iii nuclear structure is demonstrable in most
tissucls and organs of the monkey.
3. It is irifcri-ed that the sex chromatin of female monkeys
represents adherent heterochroriiatic portions of the two
X chromosomes, while the XY sex chromosome complex of
males fails t o form a chromatin mass of comparable size.
___-
The authors wish to thank 31r. J. E. Walker and Rlr. C. E.
Jarvis €or valuable technical assistance.
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during depletion and restoration of Nissl matcrial in motor neurons.
J. Anat., 8 5 : 171-181.
DARR, M. L., L. F. 3h:RTK1hL 4ND 13. A. LINDSAY1950 The morphology of
the iieivt' cell nncleus, according to sex. Anat. Rec., 1 0 7 : 283-297.
COID4N, R. S. 1932 T h e p:minueleolar bodies in spinal neurons of niarnn~nls.
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CBOUCII, Y. F., AN11 &f. 1,. BAKR 1951 Brllnviour of the bex chronlatin dullllg
axon reaction. J . Neuropath. and Exper. Neurol., 13: 353-358.
DAVIDSON,
W. M., A N D 1).R. S m T H 1951 A morplioloyical sex difference in the
polyinorplionuclcar neutrophil leueocgtes. Brit. M. J., ,? : 6-7.
EMERY,
J. L., BND M. RICMILLAN 1931 Observations on the feinalc sex rhromatin
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sex. J. Path. and Bact., 68: 17-22.
GRAHAM,&I. A. 1951 Sex chromatin in nuclei of tlir cat from tlic early embryo
to matnrity. Anat. Rec., 1 1 9 : 4(i9-492.
O r t a ~ a x 31.
, A., A N D &I. L. B S K K 195% A sex cliffermce in the rnorpliology of
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Ilsu, T. C., C. A. IIOoKS 1KD C. &f. P O h Z E R 4 T 1953 Opportunities for detcrniining sex in Iiuninn tissues. Texas Rep. Riol. and Mcd., 11:
585-587.
166
R. FC. PRIK(?ll:, M . A. GRAHAM AND M. L. BARR
LINDSAY,
H. A.,
AXD Y.11. DARR 1955 3'iirt.her observations on the bchaviour of
nitclenr st.riic,turt:s tluriiig depletion aiid rest.orat,ion of Nissl niat.cria1.
J. Allat,., S 9 : 47-62.
I~.IAKBEI~GI':K,
E., Imn W. 0. NELSOX1954 Srs differences in epidermal nuclei of
I i u i i i a i i skin. An:rt. Rcc., l18: 399.
~IIOORE, I<. r,., AN) 31.I>. R A I ~ K 1933 l;lorpliology of thc 1it~vecell liucleus in
uiiiiiiiii:i.ln, with a l d ; i l refrlcnre to tlic Yex chroniatin. d. Coiiip. Xciir.,
$ 8 : 21:1-2:11.
1 !)54 Niic.1ea.r iiiorpliology, :iceording t.0 sex, in liunian tissues.
Acts Aii:tt., :3: 19i-208.
MOOXE,I<. Id., 11. A. C~RhHAhf AND M. L. BARR 1953 The clc.t.ectioii o f clirouiosonial svx in Iicriiinphroditcs from n skin biopsy. Surg., G,yncc. n l i d
Obstct., 96: 841--618.
NYLI.13, %I., ANn 51.A. ~ R a a A N 19.54 Ses chroin:i.t,ini i i iieurons of hiininn froiit:il
cortc8.s :nit1 syiiiynhlietir ganglia. An:tt. Itec., I l S : 402.
PAINTER,
T. S. 1922 Tlie SC'S clironiosonies of tlic moiikcy. Sciuicc, 5 6 :
286-28'7.
.I924 St,udics iii 1iiiiiiiini11i:in spcriii:it,ogencsis. IT. Tlic sex cliroiiiosouics of monkeys. .I. Eq).Zool., 30: 433-462
PIAATE
1
E X PL1N.lTION 01" I.'IGUI<ES
Xerve eetls of feui:ile and mnlc monkeys. Crespl violet, X l(i00.
1 and 2
~~eiiro~
ini svrstibular niiclrus of the nicdulla oblongatn.
3 :iiicl 4 Motor iieuroiis hi tlic rriitrnl horn of the ecrvical regioii of tlie
spiiial cord.
5 :uid ti Giant pyr:iuiitl:il cells in l;i,vcr 5 of tlie iiiotor cort.c.s.
Tlic scx cliromntin is :it tlic iiiiclt.:u iiiciul)ra.iie in figures 1 :intl 3. !rliis is its
1oent.ioii throiigliout t.lic II(:TVOUH
sy:xt:w of tlit. iiioiikcy. Ti1 figiirv
5, tlic scs chromatin is :idjaecnt t o tlic Iiuclcolulc. I t is ( : ~ i i i ~ i i ~em
~ ~ i ill l~ t.liix
location ill the giant pyr:iuiid:il c d s , alt.liough the more t.ypical position, nc,st tliv
ituclcar ruwiibr:iii(~,oecurs most frcqiiciitlp in other neurons o f the ecrelJr:il cortvs.
iiicrst, con1111011
PLATE 1
167
PLATE
2
EXPLANATION OF FIGURES
Nerve cells of female and male monkeys. Cresyl violet, figures 9 and 10 are
X 2000, others X 1600.
7 and 8
Neurons of the substantia nigra.
9 and 10 Nerve cells of the caudate nucleus.
11 and 12
Mitral cells of the olfactory bulb,
The sex chromatin in figures 7, 9 and 11 is in its most common position,
adjacent t o the nuclear membrane. The sex chromatin is often interposed, more
frequently in some regions than in others, between the nuclear membrane and
an eccentric nucleolus, as illustrated by figure 9.
168
PLATE 2
169
PLATE 3
EXPLANATION O F FIGURES
13 and 14 Smooth muscle cells in the urinary bladder. 11. and E.,
x
1600.
15 and 16 Cartilage cells. H. and E., X 1600.
17 and 18 Parietal cells of gastric glands. H. and E., X 1600.
The sex chromatin is located a t the nuclear membrane, with few exceptions,
in various tissues of the monkey as illustrated by figures 13, 15 and 17.
170
SEX OHROXATIN IN XACACUS RHEBUS
I’LITE 9
B. H. PRINCE, Y. A. GRAIIAM AND If. 1,. B m B
171
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