ANNACS OF ANATOMY
Morphological characterization of the female prostate
(Skene's gland or paraurethral gland)
of Lagostomus maximus maximus
Mirta A. Flamini 1, Claudio G. Barbeito a'2, Eduardo J. Gimeno 2 and
Enrique L. Portiausky 2
1Department of Histology and Embryology and 2 Institute of Pathology,
School of Veterinary Sciences, National University of La Plata, R O. Box 2 96,
1900 La Plata, Argentina
Summary. The Skene's (paraurethral) gland is the histolo-
gic homologue to the male prostate. Much experimental
work has been done on women and on various rodents.
In this study we describe for the first time the anatomical
and histological characteristics of the paraurethral gland
in the plain viscacha (Lagostomus maximus maximus).
This gland is formed by tubuloalveolar adenomers sur-
rounded by connective tissue and smooth muscle fibers.
The adenomers are lined by cuboidal or columnar epithe-
lium. In the lumen of the adenomers, PAS positive secre-
tions could be detected. The conduits are lined by
bi-stratified epithelium. In some aspects this gland is simi-
lar to that of other mammals.
Key words: Paraurethral gland - Skene's gland - Female
Prostate - Lagostomus rnaximus maximus - Histricog-
nathi - Morphometry
Introduction
In 1672, de Graaf for the first time described the presence
of a gland located around a woman's urethra, which be-
came known as the "female prostate" due to its similarity
with the masculine prostate (de Graaf 1672). In 1853,
Virchow described the presence of prostatic concretions
as "amylaceous bodies" present in this female gland
(Virchow 1853). Years later, Alexander Skene published
a work describing the presence of two glands closely re-
lated to the urethra (Skene 1880). There onwards, this or-
Correspondence to: M. A. Flamini
E-mail: aflamini@fcv.unlp.edu.ar
Ann Anat (2002) 184:341-345
© Urban & FischerVerlag
http:llwww.urbanfischer.deljournalslannanat
gan became known as Skene's gland (Zavia~i~ and Ablin
2000). Further studies (Zavia~i6 1999) detailed the anato-
my and the histological structure of both the ducts and
the characteristics of the glandular secretion.
For a long time this gland was considered to be a vesti-
gial organ without much functional importance for the
woman. It possesses a microscopic structure similar to the
masculine prostate, presenting numerous ducts, adeno-
mers and a rich stroma of smooth muscular tissue (Za-
via~i~ et al. 2000). The presence of an acidophilic
secretion inside the adenomers and ducts, which has been
studied with enzymatic labeling techniques and immuno-
histochemistry, was also described (Tepper et al. 1984; Za-
via~i~ 1985, 1997; Zavia~i~ and Ablin 2000). These studies
showed that a marked enzymatic activity exists in the fe-
male prostatic gland and that its cells produce an exo-
crine and neuro-endocrine secretion, producing a "female
ejaculation" (Zavia~i~ 1993).
Not only have structural and physiological descriptions
point to this gland, but different pathological alterations
have also been described; such as glandular hyperplasia
(Folsom and O'Brian 1943), adenocarcinomas (Dodson
1994; Zavia~i~ et al. 1993; Sloboda et al. 1998), cysts (Ishi-
gooka et al. 1995; Romero P6rez et al. 1999) and calculi
induced by Ureaplasma urealyticum (Gellman and Der-
shewitz 1986).
The presence of this gland is not constant in the differ-
ent animal species but has been described in some insecti-
vorous species (Grass6 1969) and in different rodents:
Rattus rattus (Shehata 1972), Avicantis nUoticus, Meriones
lybicus (Shehata 1975), Rattus norvegicus (Shehata 1980),
Mastornys erytholeucus (Ichihara 1976), Paraomys (Mast-
omys) natalensis (Smith et al. 1978; Gross and DiDio
1987).
0940-9602/02/184/4-341 $15.00/0
Fig. 1. Anatomical localization of paraurethral gland (PG). Two PG glands are surrounding the urethra (U). Vagina (V) and urinary
bladder (UB) are also observed. Bar represents 0.3 cm.
Fig. 2. Section of Lagostomus maxirnus maximus parauretral gland (PG) and its two ducts (D) entering the urethra (U). HE. Bar re-
presents 400 pm,
Fig. 3. Section of the paraurethral gland showing lobular aspect. HE, × 40.
Fig. 4. Section of the paraurethral gland showing smooth muscle fibers (arrows). CF = connective fibers. Massons's Trichrome, × 40.

342
 The plains viscacha (Lagostomus maximus maximus), is
a mammal belonging to the order Rodentia, sub-order
Histricognathi, family Chinchillidae. They inhabit the
fields of Argentina, forming colonies in deep caves called
viscacha's burrows. The light governs their habits: during
the day they remain in the caves and leave their burrows
in the evening to carry out their outdoor activities. There
is a marked size and weight difference between the males
and females of the species (Llanos and Crespo 1952).
Their reproductive cycle is characterized by a main zeal
in March-April and a secondary zeal after a gestation of
153.7 + 0.5 days (Weir 1971 a).
To our knowledge, there are no publications concerning
the presence of the paraurethral gland in any species of
the sub-order Histricognathi. In this study the presence of
the paraurethral gland or female prostate in Lagostomus
rnaximus rnaximus (plain viscacha, Blainville 1817) will
be described, for the first time, together with its anatomi-
cal localization and histological structure.
Materials and methods
Animals. Seventeen mature females of Lagostomus maximus
maximus weighing approximately 4.5 to 5 kg were used. They
were captured at different periods (December-January, March-
April and July-August), with the goal of including pregnant and
non-pregnant animals in the study. Animals were anesthetized
with intramuscular doses of ketamine-chlorohydrate (50 mg/kg
body weight) and sacrificed by bleeding. The complete urogenital
system was extracted and fixed in a 10% buffered formaldehyde
solution. The material was then processed for its embedding in
paraffin.
Histological processing. Three micron thick histological slides
were stained with the following techniques: routine Hematoxy-
lin-Eosine (HE); Masson's trichrome method to differentiate
muscular from connective tissue; Gomori's argentic impregnation
for the observation of reticular fibers; Pranter's orceine to identi-
fy elastic fibers, and PAS for histochemical demonstration of mu-
cin.
Morphometrical study. For the macroscopic study a stereo-
scopic microscope (SZ40, Olympus, Japan) was used.
To carry out the morphometric study, 5 histological images
(magnification 200 x) were captured from each slide (Olympus
BX50 microscope, Japan) and digitized using an attached analo-
gue RGB video-camera (Sony DXC-151A CCD) connected to a
frame grabber (Flashpoint 128, Integral Technologies, Inc, India-
napolis, IN, USA). Twenty-four bits, TIFF format images were
then analyzed using the Image Pro Plus v4.5 image analyzer soft-
ware.
Fig. 5. Female prostate section showing reticular fibers (arrows). Gomori, x 40.
Fig. 6. Elastic fibers in the connective tissue of the paraurethral gland (arrows). Orcein, x 200.
Fig. 7. Female prostate duct showing bi-stratified epithelium with eosinophilic secretion in the lumen. HE, x 200.
Fig. 8. a: Columnar epithelium showing PAS positive granules at the supranuclear area. PAS, x 1000. b: PAS positive prostatic con-
cretions in the lumen of a lobule. PAS, x 100.
343
The following measurements were registered: major and minor
diameters (length of the longest and shortest lines joining two
outline points and passing through the centroid) of adenomers
and excretory ducts; height (caliper length along a major axis of
the object) and width (caliper length along a minor axis of the
object) of the adenomers' epithelial cells.
Statistical analysis. All data were expressed as mean + SD. A
Student's t test was used to identify significant differences be-
tween adenomer and epithelial cell parameters.
Statistical significance was accepted at the level of p < 0.05.
Results
Macroscopic analysis. All the animals studied presented
two paraurethral glands. Anatomically, the glands were
located laterally in the vicinity of the female urethra, near
the urinary bladder - one at each side of the urethra,
without completely surrounding it (Fig. 1). The size of
each gland ranged from 0.6 to 1 m m long. Each unit ter-
minated at the urethra, in independent form, through a
main duct.
Microscopic analysis. Microscopically, a lobular pattern
was observed (Figs. 2, 3). The main ducts had an oval
shape with a major axis of 220.18 + 67.77 gm and a minor
axis of 103.91 + 42.39 gm (Fig. 2). The lobules were sepa-
rated by trabeculae of connective tissue with abundant
smooth muscular fibers, seen with Masson's trichrome
technique (Fig. 4). The connective tissue presented reticu-
lar fibers (Gomori's technique) that were more evident
around the adenomers and ducts (Fig. 5) The orceine
technique showed the presence of elastic fibers in the
perilobular tissue surrounding the adenomers (Fig. 6).
Each lobule presented a main duct with a wide lumen
tlaat occasionally showed an irregular pattern. It was gen-
erally covered by a bi-stratified epithelium (Fig. 7). In its
lumen, a secretion of acidic nature similar to that found
in the male prostate could be observed.
The adenomers had a tubuloalveolar pattern. T h e dia-
meter of the adenomers did not vary statistically between
pregnant and non-pregnant females (Table 1). The epithe-
lium of the adenomers was simple, being cuboidal in the
non-pregnant females and cylindrical in the pregnant vis-
cachas (Table 2). The cells presented a rounded or oval
nucleus located in the basal region. The apical portion of
the cytoplasm had a granular aspect. The small and scarce
granules present in the apical portion of the cytoplasm
and the secretion found in the lumen of the adenomers
were positive to the PAS technique (Fig. 8).
Table 1. Morphometrical pattern of the adenomers
Morphometrical Pregnant Non-pregnant Significance
parameter
Major diameter 93.05+ 15.57 92.14+ 13.26 NS
Minor diameter 52.77+ 9.59 50.32+ 3.90 NS
All the numbers are expressed in pm _+Standard Deviation. NS:
no significance.
Table 2. Morphometrical pattern of the adenomer ceils
Morphometrical Pregnant Non-pregnant Significanceb
paramenter
Height 12.66 + 2.28 11.24+ 3.70 NS
Width 6.34 + 0.55 6.65 + 0.94 NS
Significance a * NS
All the numbers are expressed in gm+Standard Deviation.
a. Significance between height and width of the same animal
group; b. Significance between either height or width of both
groups. * p <0.05, NS: no significance
Discussion
Few studies have been carried out on the reproductive
tract of the female Lagostomus maximus maximus (Weir
1971 a, b), and no previous observation has revealed the
presence of the female prostate or paraurethral gland in
this species. Some researchers have found this gland to be
present in different species of mammals of the order Ro-
dentia, although with some differences in the size, anato-
mical location and position of the organelles inside the
cells of the adenomers. The constant presence of a bilat-
eral well-developed female prostate in some species of
the Muridae family, for example Rattus rattus, Rattus nor-
vegicus and in Avicantis niloticus has been described
(Shehata 1972, 1975, 1980). In the Cricetidae family, the
Skene's gland could only be located in Meriones lybicus.
In the latter case this is also bilateral but much smaller
that in the other above mentioned species. Gross and Di-
Dio (1987) described the female prostate in Praomys
(Mastomys) natalensis, pointing out that it is formed by
two lobes extended medially, cranially to the urine blad-
der and can be entered in contact in the ventral portion
of the urethra. This anatomical position differs to what
was observed in Lagostomus maximus maximus. In our
observations, the viscacha paraurethral gland appearance
is constant, bilateral and without contact between the two
glands. Contrary to what has been described in the other
mammals, in the woman the gland is formed by very
ramified ducts (Huffman 1948).
In Rattus rattus, Rattus norvegicus, Avicantis niloticus
and Meriones lybicus the adenomers are covered by a sin-
gle layered cuboidal or low columnar epithelium, which
contains acidophilic secretion in its cytoplasm (Shehata
1972, 1975, 1980).
344
The stroma that supports the glands is formed by con-
nective tissue and smooth muscular fibers. In Praomys
(Mastomys) natalensis columnar cells and small basal
cells form the epithelium that cover the adenomers.
Gross and DiDio (1987) also described the presence of
PAS positive granules in the cells, as well as a secretory
product with the same staining characteristics. The histo-
logical structure described in this work for Lagostomus
maximus maximus coincides in some aspects with that
mentioned for other rodents; i.e. the form of the adeno-
mers, the tissue that forms the stroma and the character-
istics of the secretion product. With regard to the cell
types, we have observed that the epithelium that sur-
rounds the glandular units is single cuboidal or colum-
nar, similar to that described for Rattus rattus and Rattus
norvegicus (Shehata 1972, 1975, 1980), whilst we have
not found basal cells as in Praomys (Mastomys) nataIen-
sis.
In some rodents the Skene's gland maintains its histo-
logical structure throughout its lifetime (Gross and Di-
Dio 1987). It was also described that its size increases
during gestation and also during the nursing period in fe-
male rodents. We found differences in the form of the
glandular cells, being cylindrical or columnar in the preg-
nant females and cuboidal in the non-pregnant animals.
It is possible that such a difference has some relationship
to the hormonal changes that take place during the preg-
nancy.
The adenomers of mucous type that Shehata (1972) de-
scribed for Rattus rattus were not observed in Lagostomus
maximus maximus.
In the woman, the female prostate has been studied
from several aspects. It has been demonstrated that the
cells react positively to prostate specific antigen of the
male and also to alkaline phosphatase. It could also be
demonstrated that the histoenzymafic parameters of
the prostatic gland in the woman were comparable to
those of the male prostate (Zavia~i~ 1985). The rele-
vance that it acquires in human beings resides not only
in the lesions that the gland can manifest, but also
whether a relationship exists between the secretion of
the gland and female ejaculation as a product of sti-
mulation during coitus, of the so called "G spot". In
animals, the existence of that ejaculation has not yet
been determined, although the presence of a well de-
veloped gland could have a similar significance. The
discovery of a prostate of constant appearance in the
viscacha female would allow the use of this animal as
an experimental model for studying the physiology of
the gland.
Acknowledgements. The technical work done by Ruben C. Ma-
rio and the personnel of the Estaci6n de Cria de Animales Sil-
vestres (ECAS), Ministerio de Asuntos Agrarios de la Provincia
de Buenos Aires is acknowledged. We would like to thank Tamar
Ferguson for the proofreading of the manuscript. ELP and EJG
are Research Career Members of CONICET (National Scientific
Research Council).
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Accepted February 12, 2002