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Pineal influence on the Harderian glands of female golden hamsters.

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Pineal Influence on the Harderian Glands of
Female Golden Hamsters
JEANNE W. CLABOUGH AND JOHN E. NORVELL
Department of Anatomy, Medical College of Virginia, Health Sciences
Division, Virginia Commonwealth University,
Richmond, Virginia 23298
ABSTRACT
The effects of blinding on cellular structure and porphyrin pigment content were studied in Harderian g1and.s of pinealectomized and nonpinealectomized adult female golden hamsters. Uterine weights were used as a
criterion for pineal activity. Five weeks post-operatively, no significant changes
were observed in the Harderian glands or in the uterine weights of animals in
either group. Eleven weeks post-operatively, the blinded hamsters with intact
pineal glands had significantly lowered uterine weights and Harderian glands
that exhibited diminished amounts of porphyrin pigments and cellular characteristics of the male hamster. Harderian glands and reproductive organs from
the 11-week blinded-pinealectomized hamsters did not differ from those of untreated controls.
During the last three years there has
been a renewal of interest in the mammalian Harderian gland regarding possible
function, metabolism, and interaction with
other organs (Wetterberg, Geller and
Yuwiler, '70a; Wetterburg, Yuwiler, Ulrich,
Geller and Wallace, '70b; Kennedy, '70;
Hoffman, '71; Reiter and Klein, '71; Norvell and Clabough, '72; Clabough and Norvell, '73; and others). Although it has been
suggested that these orbital glands supplement the lacrimal lubricant in non-aquatic
vertebrates, the presence of porphyrin pigments in Harderian glands of rodents
(Davidheiser and Figge, '58; Kennedy, '70)
suggests a more complex function. Additionally, in the adult golden hamster, these
glands show a sexual dimorphism related
to porphyrin content and cellular structure
(Christensen and Dam, '53; Woolley and
Worley, '54; Paule, Hayes and Marks, '55;
Bucana and Nadakavukaren, '72). Microscopically, Harderian glands of female
hamsters are characterized by the presence
of reddish-brown or dark amber pigment
granules in the lumina, and acini which
consist of simple columnar epithelial cells
filled with minute lipid droplets of uniform
size (type I cells). According to Paule and
co-workers ('55) the pigment granules are
a porphyrin compound. This corresponds
to the findings in other rodents in which
ANAT. REC., 178: 119-126.
i t has been demonstrated that the pigment
accumulations in the lumina of the
Harderian glands have the properties of
porphyrins (Grafflin, '42; Cohn, '55; Woodhouse and Rhodin, '63). Additionally, the
porphyrin of Harderian glands exhibits a
red fluorescence in ultraviolet light
(Kennedy, '70). In female hamsters, a
fast-f ading red fluorescence is exhibited by
the alveolar cells and a persistent red
fluorescence, by the luminal pigment granules (Hoffman, '71; Clabough and Norvell,
'73). In males, porphyrin granules and red
fluorescence are lacking, and the acini are
composed of approximately equal numbers
of two1 types of cells : one, indistinguishable
from that of the female (type I cells) and
a second having a swollen appearance and
containing numerous large cytoplasmic
droplets of various sizes (type I1 cells).
Castration of male hamsters results in a
conversion to the female type of Harderian
gland within 20-30 days (Woolley and
Worley, '54; Hoffman, '71; Clabough and
Norvell, '73). Hoffman ('71) and Clabough and Norvell ('73) have demonstrated that this effect of castration is
greatly reduced or abolished if the animals
are also blinded. Additionally, data from
our study ('73) indicate that in blindcastrated male hamsters, the pineal gland
Received April 30, '73. Accepted July 30, '73.
119
120
JEANNE W. CLABOUGH AND JOHN E. NORVELL
is responsible to preventing the conversion of the male type of gland into the
female type.
The present study was conducted to determine whether or not the pineal gland
has a similar effect on porphyrin pigment
content and cellular type i n Harderian
glands of female hamsters, i.e., does the
pineal gland in the blinded female effect
the disappearance of porphyrin granules
and the appearance of cells characteristic
of the male?
RESULTS
No significant differences were observed
in the weights of the reproductive organs
or Harderian glands or in the appearance
of the Harderian glands amcng any of the
groups of animals sacrificed five weeks
post-operatively (table 1). Harderian
glands of animals in all three groups exhibited moderate to heavy porphyrin concentrations and acini which were lined by
type I epithelial cells, typical of normal
female hamsters (fig. 1). An exception
was found in one animal from the blind,
sham-pinealectomized group. Although
MATERIALS AND METHODS
porphyrin content was normal in this aniTwenty-four female golden hamsters mal, a few type I1 cells were interspersed
(Mesocricetus auratus) weighing 95-1 10 among the type I cells throughout the gland
gm were used to establish the following (fig. 2 ) . It is noteworthy that this animal
three groups : ( 1) untreated, (2) blinded, also had the smallest ovaries and uterus in
sham-pinealectomized, ( 3 ) blind-pineal- the group (17.1 and 324.8 mg/100 gm
ectomized. Half of the animals i n each body weight, respectively).
group were sacrificed after five weeks and
After 11 weeks the blinded, sham-pinealthe remainder, after 11 weeks. Although i t ectomized group had significantly reduced
is well-known that pineal antigonadotropic uterine weights and Harderian glands with
activity is maximal in light deprived diminished porphyrin and various concenhamsters (Hoffman and Reiter, '65; Reiter, trations of type I1 cells (table 1, fig. 3 ) . Of
Hoffman and Hester, '66; Reiter, '67), the three 11-week groups, the mean
ovarian and uterine weights were used to ovarian weight was lowest in this group,
substantiate this fact in the present study. but the difference was not significant. The
Harderian glands were weighed, and one animal that exhibited the least amount of
or both glands were fixed in 10% buffered porphyrin also had the greatest concentraformalin and processed for light micros- tion of type I1 cells and the smallest uterine
copy. All sections were stained with hema- weight (67.2 mg/100 gm body weight). A
toxylin and eosin. Porphyrin content in the similar correlation, however, did not exist
Harderian glands was determined by gross for the remainder of the animals i n this
and microscopic examination and was group, e.g., the animal which exhibited
rated from light ( V -) to moderate ( V ) the heaviest concentration of porphyrin
to heavy ( V + ) . I Grossly the pigment had approximately equal numbers of type
granules appear as small brown flecks near I and type I1 cells and a relatively light
the surface of the yellow-white gland. uterus. Neither loss of uterine weight nor
With hematoxylin and eosin stain, the alteration toward the male type of
granules have a deep amber hue. Accord- Harderian gland occurred in any of the
ing to the above scale, porphyrin concen- blinded hamsters that were also pinealectrations i n normal female hamsters range tomized. I n fact, slight uterine hyperfrom moderate to heavy. Surgical proce- trophy was evidenced in this group.
Hoffman ('71) stated that in male
dures and housing conditions were the
same as those described in the earlier hamsters, type I1 cells appear to be transstudies on the Harderian glands of male formed from type I cells by accumulating
hamsters (Clabough and Norvell, ' 7 3 ) .
1 Ratings were assigned subjectively after examinserial sections of each Harderian gland. d- inSham-pinealectomy consisted of reflecting ing
dicates a few scattered small porphyrin granules
the skin, removing the plate of bone over- (usually in less than 20 lumina per longitudinal section); d indicates a porphyrin granule content interlying the pineal gland, and rupturing the mediate between d- and ++, i.e., sections were not
recognizable as either light or heavy. d+ inconfluens of sinuses in the manner done readily
dicates porphyrin granules of various sizes and numfor pinealectomy, without, however, dis- bers in approximately one-quarter or more of the
lumina in each longitudinal section (usually in 150
turbing the pineal gland.
lumina or more).
2
2
5
1. None (Control)
a. 5weeks
b. 11 weeks
sham2. Blinded
pinealectomized
a. 5weeks
5
5
3. Blinded
pinealectomized
a. 5weeks
b. 11 weeks
9826
10024
98a3
10226
10623
10620
Initial
124&7
12027
13028
120-1-3
13125
13223
Final
Body weight
3222
28&2
3020
2825
27a3
33k3
mg
26-1-2
2324
2320
22-1-5
2024
2523
bodywt.
mg/100 gm
558217
420&14
i8'22433
496k67
410k30
505-1-35
mg
456C7=
350212
i4Oiii3
414-t-55
313210
384+-35
mg/100 gm
body w t .
Uterine weight
102k3
82k2
6822
7?&3
9323
82k7
68&4
61k3
74-1-4
mg/100gm
1
2
d+/5
d /2
d+/3
-J /2
d+/1
d- /2
d+ /5
d+ / 2
\ I t - /2
Harderian glands
Porphyrin 4
8125
8024
9825
mg
Weight
I/5
I/5
I = II/1
I/4
> II/l
I1 > I / 2
1 > II/2
I
I/2
I/2
Cell type
on the Harderian glands of adult female hamsters after 5 and 11 weeks
Ovarian weight
+ pinealectomy
Weights are recorded +- standard error and are compared with the appropriate control according to Fisher's t-test.
Value differs from control value; P f 0.05.
3 Value differs from control value; P f0.001.
4 Arabic numerals represent number of animals showing the indicated porphyrin content or cell type.
+
5
b. 11weeks
+
No. of
animals
Treatment
and group
Effects of blinding and blinding
TABLE 1
X
122
JEANNE W. CLABOUGH AND JOHN E. NORVELL
more lipid, thus giving rise to larger droplets with less uniformity of size. In the
blinded, sham-pinealectomized females of
the present study, apparent transition
stages between type I and type I1 cells also
indicate a transformation of type I cells
into type I1 cells. In all of the hamsters in
this study, porphyrin granules and cellular debris, including pyknotic nuclei, were
found in the lumina of ducts and of
numerous acini (fig. 4). Porphyrin granules often appeared to have a lamellar configuration or to contain small spherical
sub-units within a single granule. Similar
small spherical granules were commonly
seen in the cells which line large ducts in
the Harderian glands (fig. 4). The abundance in this location was much greater
than that observed in castrated males (Clabough and Norvell, '73). Also of common
occurrence in the female were mast cells.
Of less frequent occurrence in females
than in males were mitotic figures and
small intracellular pigment granules in
the acinar cells.
enhancing pineal antigonadotropic activity, prevents this transformation.
Our studies on both male and female
hamsters provide evidence that the pineal
gland in the blinded animal is responsible
for the presence of type I1 cells and for the
decrease in the formation or storage of
porphyrin granules in the Harderian
glands. Such responses to blinding do not
occur if the animals have also been pinealectomized. In the female with the pineal
gland intact, this response is not evident
morphologically until between five and
eleven weeks after blinding. This also corresponds to the time required for significant changes in uterine weight to occur.
The presence of a few type I1 cells in the
Harderian glands of one blind, shampinealectomized female after five weeks
suggests that morphological changes are
just beginning to occur at this time. In
both male and female hamsters, definitive
morphological changes in the Harderian
glands and in porphyrin content in response to blinding and other experimental
procedures appear to require five weeks or
more (Hoffman, '70; Clabough and Norvell,
DISCUSSION
'73). Longer time periods may provide
The first reports linking the pineal gland even more conclusive data, such as whether
to the Harderian gland indicated that in or not in blind, sham-pinealectomized
blinded suckling rats the Harderian glands female hamsters all porphyrin pigment
act as photoreceptor organs mediating the would eventually disappear.
effects of light on pineal biochemical activQuestions still remain about the sigity (Wetterberg et al., '70a,b). Reiter and nificance of porphyrin in the Harderian
Klein ('71) were unable to find evidence that glands of rodents and the sexual dimorthe Harderian gland has such a function phism peculiar to the Harderian glands of
in adult female rats. Several observations, hamsters. Biochemical, histochemical and
considered collectively, have prompted the morphological evidence indicates that pordifferent suggestion that in the hamster phyrin is synthesized in the Harderian
the pineal gland has a regulatory effect on glands of mice and rats (Kennedy, '70).
the Harderian glands (Clabough and The presence of small reddish-brown granNorvell, '73): in this species Harderian ules in the acinar cells of castrated males
glands exhibit a sexual dimorphism; castra- and in the duct cells of females may be
tion of males results in Harderian glands related to stages of porphyrin synthesis
of the female type; this effect of castration and release in the hamster. Possibly the
is prevented by blinding but still occurs in sexual dimorphism in the hamster is rethe Harderian glands of both orbits after lated to the LH surge in the female since
unilateral enucleation; blinding leads to i t appears likely that porphyrin formation
elevated pineal antigonadotropic activity. is dependent on elevated gonadotropic horBased on these observations, it was further mones in this animal.
suggested that following castration, transLITERATURE CITED
formation of Harderian glands of the male
C . D., and M. J . Nadakavukaren 1972
type into the female type is probably a Bucana,
Fine structure of the hamster Harderian gland.
consequence of elevated levels of gonado2. Zellforsch., 129: 178-187.
trophic hormone and that blinding, by Christensen, F., and H. Dam 1953 A sexual
HARDERIAN GLANDS OF FEMALE HAMSTERS
dimorphism of the Harderian glands in
hamsters. Acta Physiol. Scand., 27: 333-336.
Clabough, J. W., and J. E. Norvell 1973 Effects
of castration, blinding, and the pineal gland
on the Harderian glands of the male golden
hamster. Neuroendocrinology, 12: 344-353.
Cohn, S . A. 1955 Histochemical observations
on the Harderian gland of the albino mouse.
J. Histochem. Cytochem., 3: 342-353.
Davidheiser, R. H.,and F. H. J. Figge 1958
Comparison of porphyrin producing enzyme
activities in Harderian glands of mice and
other rodents. Proc. SOC.Exp. Biol. Med., 97:
775-778.
GraMin, A. L. 1942 Histological observations
upon the porphyrin-excreting Harderian gland
of the albino rat. Am. J. Anat., 71: 43-64.
Hoffman, R. A. 1971 Influence of some endocrine glands, hormones and blinding on the
histology and porphyrins of the Harderian
glands of golden hamsters. Am. J. Anat., 132:
463-478.
Hoffman, R. A., and R. J. Reiter 1965 Pineal
gland: influence on gonads of male hamsters.
Science, 148: 1609-1611.
Kennedy, G. Y. 1970 Harderoporphyrin: A
new porphyrin from the Harderian gland of
the rat. Comp. Biochem. Physiol., 36: 21-36.
Norvell, J. E., and J. W. Clabough 1972
Adrenergic and cholinergic innervation of the
hamster Harderian gland. Science, 178: 11021103.
Paule, W. J., E. R. Hayes and B. H. Marks 1955
123
The Harderian gland of the Syrian hamster.
Anat. Rec., 121: 349-350.
Reiter, R. J. 1967 The effect of pineal grafts,
pinealectomy, and denervation of the pineal
gland on the reproductive organs of male
hamsters. Neuroendocrinology, 2: 138-146.
Reiter, R. J., R. A. Hoffman, and R. J. Hester
1966 The effects of thiourea, photoperiod
and the pineal gland on the thyroid adrenal
and reproductive organs of female hamsters.
J. EXP. ZOO^., 162: 263-268.
Reiter, R. J., and D. C. Klein 1971 Observations on the pineal gland, the Harderian glands,
the r'etina, and the reproductive organs of adult
female rats exposed to continuous light. J.
Endocr., 51: 117-125.
Wetterberg, L., E. Geller and A. Yuwiler 1970a
Harderian gland: A n extraretinal photoreceptor influencing the pineal gland in neonatal
rats. Science, 167: 884-885.
Wetterberg, L., A. Yuwiler, R. Ulrich, E. Geller
and R. Wallace 1970b Harderian gland: Influence of pineal hydroxyindole-0-methyltransferase activity in neonatal rats. Science, 170:
194-196.
Woodhouse, M. A., and J. A. G. Rhodin 1963
The ultrastructure of the Harderian gland of
the mouse with particular reference to the
formation of its secretorv~- products. 5. Ultrastruct. Res., 9: 76-98.
Woollelr, G . W., and J . Worley 1954 Sexual
dimorphism in the Harderian gland of the
hamster (Cricetus auratus). Anat. Rec., 11 8:
416-,417.
PLATE 1
EXPLANATION OF FIGURES
124
1
A typical Harderian gland from a normal female hamster. Acini are
composed of type I epithelial cells and contain dark reddish-brown
porphyrin granules and cellular debris (arrows). X 285.
2
Harderian gland from a blinded, sham-pinealectomized female
hamster. Five weeks post-operatively, a few type I1 acinar cells
(arrows) were interspersed among the type I cells in this animal.
Porphyrin content is normal. X 285.
3
Harderian gland from a blinded, sham-pinealectomized female hamster, 11 weeks post-operatively. Type I and type I1 acinar cells are
approximately equal in number, representing an average response to
blinding for this group of animals. X 285.
4
A typical large duct in the hilar region of a Harderian gland from
a normal female hamster. The lumen of the duct is filled with
porphyrin granules and cellular debris. X 180. Epithelial cells bordering on the lumen contain small dark granules of the color and
density of porphyrin. Inset. X 400.
HARDERIAN GLANDS OF FEMALE HAMSTERS
Jeanne W. Clabough and John E. Norvell
PLATE 1
125
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