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The effects of ovarian hormones on uterine phosphatases of the rhesus monkey (Macaca Mulatta).

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The Effects of Ovarian Hormones on Uterine
Phosphatases of the Rhesus Monkey
(Maccrca m uIaf ta)
JOHN P. MANNING,' FREDERICK L. HISAW? BERNARD G. STEINETZ *
AND ROBERT L. KROC
Department of Physiology, Warner-Lambert Research Institute, Morris
Plains, New Jersey and Biological Laboraturies, Harvard University,
Cambridge, Massachusetts 2
ABSTRACT
Endometrial acid and alkaline phosphatases were studied histochemically in rhesus monkeys treated with various combinations of estrogen (E, 178estradiol and/or estriol), progesterone (P) and relaxin ( R ) or a low potent relaxin
control preparation (NRF). In the cells of the uterine glands of the E-treated animal,
the apical cytoplasm showed intense activity of both phosphatases. This estrogenic
response was depressed in the stratum functionale by P and in the stratum basale by
PR. With E, acid phosphatase-staining granules appeared in scattered stromal cells
with eccentric nuclei. Addition of P or PR increased the number of acid phosphatase
positive stromal cells, especially in the stratum functionale. With the exception of the
sinus-like channels and superficial vessels of the stratum functionale of monkeys
treated with ER and EPR, all endothelium of capillary and precapillary vessels was
rich in alkaline phosphatase activity. Thus, acid alkaline phosphatases appear to be
metabolically important components of the endometrium which undergo cyclic variation and reflect specific influences of the ovarian hormones; estrogens, progesterone
and relaxin. The implications to human menstrual physiology are discussed.
Although relaxin ( R ) has been detected
in blood and tissues of pregnant women,
its function in human reproduction remains unknown (Zarrow, Holmstrom and
Salhanick, '55; Dallenbach and DallenbachHellweg, '64; Steinetz, Beach, Tripp and
DeFalco, '64). However, in subhuman primates (Macaca mulatta) two important
effects on uterine structure have been
ascribed to R acting in concert with estrogens (E) and progesterone (P): (1) preferential stimulation of endometrial growth;
(2) dilatation of the capillaries of the
stratum functionale into sinus-like channels with hypertrophy and hyperplasia of
endothelial cells (Hisaw and Hisaw, '64;
Dallenbach-Hellwig, Dawson and Hisaw,
'66). The latter effect, termed "endotheliod cytomorphosis" (ECM) , is normally
seen in endometrial vessels just below developing implantation sites in pregnant
rhesus monkeys (Wislocki and Streeter,
'38). Very limited sinus-like channels may
also be found in ovariectomized monkeys
treated with E and P (Cleveland, '41), and
R has been detected in blood of similarly
treated animals (Hisaw and Hisaw, '64).
ANAT. REC.,157: 465-480.
ECM also occurs in non-pregnant monkeys when the luted phase of the cycle is
prolonged by injections of human chorionic
gonadotrophin (Hisaw, '44), but the reaction is not nearly as extensive as that produced by EPR (Dallenbach-Hellweg, Dawson and Hisaw, '66). These findings suggest possible roles of R in the processes of
menstruation, implantation and placentation. Variations in endometrial acid and
alkaline phosphatases which occur during
the human menstrual cycle are thought to
reflect the pattern of secretion of endogenous ovarian hormones (Atkinson and
Engle, '47; Arzac and Blanchet, '48; Hall,
'50; McKay, Hertig, Bardawil and Velardo,
'56; Long and Doko, '59; Levine, '63;
Boutselis, de Neef, Ullery and George, '63;
Gross, '64). Exogenous E and P alter the
distribution of alkaline phosphatase in the
simian endometrium (Atkinson and Engle,
'47), but there is no information about
either alkaline or acid phosphatase in uteri
of cycling monkeys or in monkeys treated
with hormonal combinations which induce
ECM. In the persent study, the pattern of
endometrial phosphatases in rhesus mon465
466
J. P. MANNING, F. L. HISAW, B. G. STEINETZ AND R. L. KROC
keys treated with EPR has been found to
differ from that obtained with EP alone,
and in many ways resembles closely the
distribution seen in the surface epithelium,
glands and stroma of pre-menstrual human endometria.
extent, in the sub-nuclear region near the
basement membrane (figs. 12, 14). In
the arterioles, precapillary arterioles and
capillaries of the uterus, the alkaline enzyme was limited to the intima area. In
some instances, an enzyme reaction was
demonstrable within the lumen of the
MATERIAL AND METHODS
uterine glands.
The acid phosphatase activity was prinFive mature ovariectomized and 15 immature intact or castrated female rhesus cipally at the apical border of the epithemonkeys received by subcutaneous injec- lial cells of the endometrum, however, it
tion either estrogen (E, 17P-estradiol and/ had a more diffuse localization in the rest
or estriol), progesterone (P) or relaxin of the cytoplasm than did the alkaline en
(R) or a combination of these hormones. zyme (figs. 5, 6). Depending on the horThe presence of ovaries in immature ani- mone treatment, acid phosphatase was
mals did not seem to influence the results. found in endometrial stromal cells with
The castrated monkeys, after post-operative eccentric nuclei (figs. 4, 6, 8, 9). Hemableeding, were usually primed with 17p toxylin and triosin preparations of uteri,
estradiol before receiving the several com- demonstrated similar interstitial cells conbinations of estrogen, progesterone and taining kidney-shaped nuclei and numerrelaxin. Steroids were injected once daily ous triosinophilic granules (fig. 10).
in 0.1 ml propylene glycol while relaxin
The myometrial phosphatases were
was injected twice daily; in 0.85% saline weakly reactive and unaltered throughout
in the early morning and in 20% gelatin the whole experiment and will not be
in the late evening, the total daily dose considered further.
being 6000 guinea pig units (GPU).
In most animals treated with either ER
Monkeys 298 and 299 received only the or EPR the stratum functionale demonaqueous relaxin (total dose, 6000 GPU) strated large sinus-like channels with a
stratified round to oval endothelium (figs.
twice daily.
Treatments, body weights and types of 7, 13, 14). These structuers (endotheliod
cytomorphosis) are similar to those demonkeys are shown in table 1.
The animals were killed 6-18 hours after scribed previously (Dallenbach-Hellweg,
the last hormone injection. The uteri were Dawson and Hisaw, '66).
removed rapidly and small pieces fixed in
Control ( C J - 1
a 10% neutral formalin and 1% CaCL
solution for 24 hours at 4°C. After fixaThe phosphatases were weakly reactive
tion, the tissues were washed in running in the epithelial cells of the surface epitap water for two hours and immersed in thelium and of some uterine glands (table
a 0.88 M sucrose solution containing 1% 2, figs. 1, 2 ) . The alkaline enzyme was
gum acacia for 48 hours at 4°C. Six mi- positive in the blood vessels of the myocron cryostat sections were subjected to metrium and endometrium (table 2, fig.
the histochemical methods for acid and 1). No endothelioid cytomorphosis or apalkaline phosphatase using naphthol AS- preciable stromal phosphatases were obTR and naphthol AS-MX, respectively, as served.
substrates and fast red violet LB as a
Estrogen (33) - 270
coupIing reagent (Burstone, '62). Parallel
The cells of the glands of the endosections were stained with hematoxylin
metrium of this E-treated, adult, castrated
and triosin.
monkey demonstrated an intense apical reRESULTS
action for both phosphatases (table 2, fig.
In the glandular and surface epithelial 5). The vascular alkaline phosphatase and
cells of the endometrium, the alkaline scattered acid phosphatase-positive stromal
phosphatase was localized primarily near cells with eccentric nuclei were noted. No
the luminal border and, to a much lesser endotheliod cytomorphosis was present.
3265
6980
3570
5460
5665
5000
3850
3425
2980
3585
3985
3790
4360
3560
3135
3735
3875
4230
4030
4000
gm
Body
wt.
1-14
1-13
1-24
1-24
1-15
1-15
IN
IN
IN
1-20
1-44
1-20
1-41
1-44
1-53
1-40
1-20
1-20
1-20
1-20
1-20
lOpg
16-35
0.25mg
21-51
16-35
21-52
0.5mg
Estriol
0.25mg
Estrogen (E)
17 j3-Estradiol
IC
IC
IN
IN
IN
AC
AC
AC
AC
AC
IN
IN
IN
IN
IN
IN
IN
Type 1
15-24
14-23
21-33
21-33
21-52
21-51
21-55
21-55
1653
21-40
2.0mg
15-24
25-44
34-53
2.5mg
24-43
34-53
5.0mg
Progesterone ( P )
Days
J
31-52
31-51
35-55
35-55
2544
3452
25-49 5
34-52 5
22-41 3
1-20 5
16-35
16-35
25-40 5
24-39 5
15-24 6,e
15-24 5,e
6000 GPU
Relaxin (R)
33-53
1.4 GPU
Non:
relaxin
fraction
( NRF )
* Relaxin
1 IN,
immature, non-castrate; AC, adult, castrate; IC, immature, castrate.
(W1279-8, 0.23 udts/mg, Warner-Lambert). This is the non-relaxin fraction (NRF) prepared by starch gel electrophoresis of W1279-7.
3 Relaxin (W1279-7. 1000 units/mg, Warner-Lambert). This extract contains approximately 30% of the relaxin component by starch gel electrophoresis.
4 Relaxin (W1279-9, 2000 units/mg, Warner-Lambert). This is the relaxin fraction prepared by starch gel electrophoresis of W1279-7.
5Re!aXin (W1164, 4BE2103a, lo00 units/mg, Warner-Lambert). This preparation contains about 30% of the relaxin component by starch gel electrophoresis.
6Receiyed aqueous relaxin only; 3000 GPU in the morning and 3000 GPU at night. All other animals were treated with 3000 GPU aqueous relaxin in
the mornmg and 3000 GPU gelatm relaxin at night.
1
235
237
270
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
Animal
no.
Treatment schedule
TABLE 1
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J. P. MANNING, F. L. HISAW,
B. G . STEINETZ AND R
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HORMONES ON UTERINE PHOSPHATASES
Estrogen (E), progesterone ( P ) - 237
Most uterine glands of the stratum functionale were only weakly reactive whereas
the glandular epithelial cells of the stratum basale showed intense apical activity
for both the acid and alkaline enzymes
(table 2, fig. 6). The number of interstitial cells with acid phosphatase activity
was increased especially in the functional
layer. Vascular alkaline phosphatase was
unchanged and no endothelioid cytomorphosis was present.
469
DISCUSSION
Hormonal effects on endometrial
alkaline phosphatase
Following E treatment of the ovariectomized monkey, an intense alkaline phosphatase reaction was observed in the surface and glandular epithelium of the endometrium, confirming Atkinson and Engle
('47). The distribution and intensity were
similar to that described for the stratum
functionale of the human endometrium
during the follicular phase of the cycle
Estrogen (E), progesterone ( P ) , non(Atkinson and Engle, '47; Arzac and
relaxin fraction (NRF) - 235
Blanchet, 48; Hall, '50; McKay et al., '56;
This animal was similar to monkey 237 Levine, '63). The human and simian
(EP) except no increase in the number of endometrial responses to estrogen appear
acid phosphatase-positive stromal cells was to differ, however, in that the glandular
epithelium of the stratum basale of the
noted, (table 2).
monkey is intensely phosphatase positive
Estrogen ( E ) , progesterone ( P ) , relaxin
while that of the human being contains
(R) - 284, 285, 286, 287, 289, 291,
only traces of reactive enzyme.
296, 297, 299
When P was combined with E, we obExcept for animal 299, alkaline and acid served a marked reduction in epithelial
phosphatases were significantly depressed alkaline phosphatase with variable quanin the uterine glands of both the stratum tities of luminal activity appearing in the
functionale and basale of all monkeys stratum functionale, again confirming Attreated with EPR (table 2, figs. 3, 4). The kinson and Engle ('47). The glands of
interstitial cells with acid phosphatase ac- the basale were uninfluenced by P. These
tivity were numerous in the stratum func- changes in the functionale are similar to
tionale (table 2, figs. 8, 9, 10). The blood those described in the human luteal phase
vessels and the sinus-like channels (fig. where there is a progressive decrease in
7 with endothelioid cytomorphosis) of the epithelial alkaline phosphatase and a shift
functional layer were negative for the in location of the enzyme to the luminal
alkaline enzyme. Animal 299 (received secretions (Atkinson and Engle, '47; Arzac
PR for only 10 days instead of the usual and Blanchet, '48; Hall, '50; McKay et al.,
30-40 days) was similar to monkey 270 56; Levine, '63; Boutselis et al., '63; Gross,
which received only E (table 2). It is '64).
probable that the duration of treatment
When a combination of EP and R (but
was too short. Evaluation of the functional not ER alone) was administered only scatlayer and of cytomorphosis in monkey 286 tered granules of alkaline phosphatase
was impossible because breakthrough were seen in all epithelia of the endobleeding occurred the day before the ani- metrium. This picture is strikingly like
mal was killed.
that observed in the pre-menstrual human
endometrium
(Atkinson and Engle, '47;
Estrogen ( E ) , relaxin ( R ) - 288, 290,
Arzac and Blanchet, '48; Hall, '50; McKay
292, 294, 295, 298
et al., '56; Levine, '63).
Glandular and stromal activity for the
Epithelial alkaline phosphatase in the
acid and alkaline phosphatase was similar simian endometrium therefore is clearly
to animal 270 which received E alone influenced by hormones in a way similar
(table 2, figs. 11, 12, 13, 14). The blood to that observed in the human menstrual
vessels and the proliferated endothelial cycle. We postulate that E induces, accells of the stratum functionale did not tivates or accelerates synthesis of the endemonstrate the characteristic vascular zyme in epithelia of both endometrial
alkaline phosphatase reaction (fig. 14).
strata. This effect is specifically reversed
470
J. P. MANNING, F. L. HISAW, B. G. STEINETZ AND R
by P in the functionale and by a combination of P and R in the basale.
The endothelium of capillary and precapillary vessels of the stratum basale was
rich in alkaline phosphtase in all monkeys regardless of treatment. However, the
blood vessels of the stratum functionale,
which were strongly phosphatase-positive
in control, E, EP or R-treated monkeys,
underwent a striking metamorphosis when
combinations of ER or EPR were administered. Following these latter treatments,
both the superficial vessels and the sinuslike channels (ECM) which were formed,
were devoid of alkaline phosphatase. Thus,
R may exert unique effects on the E-primed
uterus (independent of P) which may be
important to the process of menstruation:
i.e.; development of ECM and loss of alkaline phosphatase from the vessels in the
functionale.
L. KROC
the human menstrual cycle (Goldberg and
Jones, '56). The enzyme-laden interstitial
cells observed in the present study may be
similar or identical to the simian endometrial granulocytes described previously
(Dallenbach-Hellweg, Dawson and Hisaw,
'66). In addition to a similarity in location and morphology, the granulocytes increase in number under the same hormonal conditions as do the interstitial cells.
Granulocytes are also found in human endometria, and evidence has been obtained
with immunofluorescent techniques that
their granules contain relaxin (Dallenbach
and Dallenbach-Hellweg, '64). Relaxin has
also been detected by bioassay in serum of
ovariectomized monkeys treated with EP
(Hisaw and Hisaw, '64) at about the same
time that granulocytic hyperplasia occurs.
If our acid phosphatase-positive interstitial cells are in fact identical with endometrial granulocytes, an intracellular relationship between relaxin and the synthesis
and/or release of acid phosphatase and
other catabolic lysosomal enzymes might
exist. Such a relationship would be in
keeping with the postulate of Dallenbach
and Dallenbach-Hellweg ('64) that relaxin
may be responsible for the dissolution and
shedding of the menstrual mucosa.
Hormonal effectsof endometrial
acid phosphatase
Lining epithelium. The pattern of acid
phosphatase in the epithelia of both the
stratum basale and stratum functionale
closely paralleled that of alkaline phosphatase under the several hormonal treatments: with E, the enzyme increased in
the epithelia of both strata; with P, there
ACKNOWLEDGMENT
was a reduction of acid phosphatase in the
We wish to express our appreciation to
functionale, while with PR the enzyme
also disappeared from the basale, in E Mrs. Margaret C. Butler, Miss Sara Priester
primed monkeys. These changes contrast and Miss Shirley G. Carter for their techwith reports of an increased acid phos- nical assistance.
phatase reaction in the epithelium of huLITERATURE CITED
man endometrial glands during the luteal
phase of the menstrual cycle (Goldberg Arzac, J. P., and E. Blanchet 1948 Alkaline
phosphatase and glycogen in human endoand Jones, '56; McKay et al., '56; Boutselis
metrium. J. Clin. Endocrinol., 8: 315-324.
et al., '63; Gross, '64; Garcia-Bunuel and
Atkinson, W. B., and E. T. Engle 1947 Studies
Brandes, '66).
on endometrial alkaline phosphatase during the
human menstrual cycle and in the hormoneStroma
treated monkey. Endocrinology, 40: 327433.
The stromal cells of the stratum basale Boutselis, J. G., J. C. de Neef, J. C. Ullery and
0.0.George 1963 Histochemical and cytowere nearly devoid of acid phosphatase
logic observations in the normal human endoactivity regardless of hormonal treatment.
metrium. Obstet. Gynecol., 21: 423-434.
Scattered acid phosphatase-positive stro- Burstone, M. S. 1962 In: Enzyme Histochemma1 cells with eccentric nuclei were obistry and its Application in the Study of Neoplasms. Pp. 160-292. Academic Press, New
served in the functional layer of the endoYork and London.
metrium following treatment with E.
These cells became hyperplastic in mon- Cleveland, R. 1941 Cytologic and histologic
observations on the epithelial, connective and
keys treated with P or PR in addition to
vascular tissues of the endometrium of macaE. These changes in stromal acid phosques under various experimental conditions.
Endocrinology, 28: 388-405.
phatase duplicate those recorded during
HORMONES O N UTERINE PHOSPHATASES
Dallenbach, F. D., and G. Dallenbach-Hellweg
1964 Immunohistologische untersuchunger ZUI
lokalisation des relaxins i n menschlicher placenta und decidua. Virchows Arch. Path. Anat.,
337: 301-316.
Dallenbach-Hellweg, G., A. B. Dawson and F. L.
Hisaw 1966 The effect of relaxin on the
endometrium of monkeys. Histological and
histochemical studies. Am. J. Anat., 119: 6177.
Garcia-Bunuel, R., and D. Brandes 1966 Lysosomal enzymes in human endometrium. Am.
J. Obstet. Gynecol., 94: 1045-1055.
Goldberg, B., and H. W. Jones 1956 Acid phosphatase of the endometrium. Histochemical
demonstration in various normal and pathologic
conditions. Obstet. Gynecol., 7: 542-546.
Gross, S. J. 1964 Histochemistry of normal and
abnormal endometrium. Am. J. Obstet. Gynecol.,
88: 647-666.
Hall, J. E. 1950 Alkaline phosphatase i n human endometrium. Am. J. Obstet. Gynecol.,
60: 212-216.
Hisaw, F. L. 1944 The placental gonadotropin
and luteal functional in monkeys (Macaca
mulatta). Yale J. Biol. Med., 17: 119-137.
Hisaw, F. L.,Jr., and F. L. Hisaw 1964 Effect
of relaxin on the uterus of monkeys (Macaca
mulatta) with observations on the cervix and
471
symphysis pubis. Am. J. Obstet. Gynecol., 89:
141-155.
Levine, B. 1963 Sex steroids, alkaline phosphatase, and endometrial carcinoma. Obstet.
Gynecol., 22: 563-566.
Long, M. E., and F. Doko 1959 Cytochemical
studies on nonmalignant and malignant human
endometrial. Ann. N. Y. Acad. Sci., 75: 504-
523.
McKay, D. G.,A. T. Hertig, W. A. Bardawil and
J. T. Velardo 1956 Histochemical observations on the endometrium. 1. Normal endometrium. Obstet. Gynecol., 8: 22-29.
Steinetz, B., V. L. Beach, L. V. Tripp and R. J.
DeFalco 1964 Reactions of antisera to porcine relaxin with relaxin-containing tissues of
other species in viuo and in uitro. Acta Endocnnol., 47: 371-384.
Wislocki, G. B., and G. L. Streeter 1938 On the
placentation on the macaque (Macaca mulatta),
from the time of implantation until the formation of the definitive placenta. Carnegie Contrib. to Embryol., 27: 1-66.
Zarrow, M. X., E. G. Holmstrom and H. A. Salhanick 1955 The concentration of relaxin
in the blood serum and other tissues of women
during pregnancy. J. CIin. Endocrinol., 15:
22-27.
PLATE 1
EXPLANATION OF FIGURES
1
The uterus from the control, untreated, intact monkey demonstrating
the localizatioii of alkaline phosphatase in the surf ace epithelium ( S 1,
uterine glands ( U ) and blood vessels (B). 300 X.
2
Acid phosphatase activity in the uterus of the normal, immature,
intact monkey (surface epithelium, S; uterine gland, U ) . 300 X.
3 The glandular alkaline phosphatase reaction of the stratum basale
from an EPR animal (uterine gland, U; blood vessels, B). 300 X.
4
472
A frozen section of uterus stained for the acid enzyme. All monkeys
treated with EPR demonstrated a depression of the phosphatases in
both the stratum functionale and basale (uterine glands, U ) . The
acid phosphatase positive stromal cells ( G ) were especially numerous
in the stratum functionale. 300 x.
HORMONES ON UTERINE PHOSPHATASES
John P. Manning, Frederick L. Hisaw, Bernard G. Steinetz and Robert L. Kroc
PLATE 1
473
PLATE 2
EXPLANATION OF FIGURES
5 The intense acid phosphatase reaction in uterine glands of the
stratum functionale of the E-treated animal (270). Alkaline phosphatase was similarly distributed. 400 X.
6
4 74
The uterine gland acid phosphatase activity of the monkey (237) receiving EP. There was a depression of the enzyme in the stratum
functionale (SF) whiIe the stratum basale ( S B ) was unaltered
(stromal cells, G ) . Monkey 235 (EPNRF) exhibited the same intense
glandular localization of the enzyme. 300 X.
HORMONES ON UTERINE PHOSPHATASES
John P. Manning, Frederick L. Hisaw, Bernard G. Steinetz and Robert L. Kroc
PLATE 2
475
PLATE 3
EXPLANATION O F F I G U R E S
7
A hematoxylin and triosin frozen section of uterus from EPR monkey exhibiting an area of endothelioid cytomorphosis ( C ) and uterine
glands ( U ). Similar structures (endothelioid cytomorphosis) were
described by Dallenbach-Hellweg, Dawson and Hisaw ('66) who reported that the endometrial capillaries and venules of the €unctional
layer became greatly dilated with hypertrophy and hyperplasia of
the endothelium. 300 X.
8
Uterine interstitial tissue demonstrating the acid phosphatase-positive cells ( G ) with eccentric nuclei which increase in number especially in the functional layer of the endometrium with P or PR
treatment. 750 X.
9
This is a stromal cell with acid phosphatase-staining granules and
the characteristic eccentric nucleus. 1900 x.
10 A section of endometrium from an area with a concentration of acid
phosphatase positive stromal cells. Note the cells have eccentric
nuclei and large numbers of triosinophilic granules. 1900 X.
476
HORMONES ON UTERINE PHOSPHATASES
John P. Manning, Frederick L. Hisaw, Bernard G . Steinetz and Robert L. Kroc
PLATE 3
477
PLATE
4
EXPLANATION O F FIGURES
11 Glandular acid phosphatase reaction in an animal treated with ER.
50 X.
12 Glandular alkaline phosphatase reaction in a n ER monkey, 50 X.
13 A section of uterus from a n ER monkey stained for acid phosphatase.
Note the glandular (U) activity in the presence of endotheloid cytomorphosis ( C ) ; thus P, although synergistic with R in altering the
intensity and distribution of the phosphatases is not required for the
blood vascular alteration. 300 X.
14 Same as above, except stained for alkaline phosphatase (cytomorphosis, C ; uterine gland, U). 300 X .
478
HORMONES O N UTERINE PHOSPHATASES
John P. Manning, Frederick L. Hisaw, Bernard G. Steinetz and Robert L. Kroc
PLATE 4
479
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