trans-Resveratrol, a Natural Antioxidant from Grapes, Increases Sperm
Output in Healthy Rats1
M. Emı ´lia Juan, Eulalia Gonza ´lez-Pons, Thais Munuera, Joan Ballester,*
Joan E. Rodrı ´guez-Gil,* and Joana M. Planas2
Departament de Fisiologia, Facultat de Farma `cia, Universitat de Barcelona, E-08028 Barcelona, Spain and
*Unit of Reproduccio ´ Animal, Department of Medicina i Cirurgia Animal, Universitat Auto `noma de Barcelona,
tection against cardiovascular disease. One of the mechanisms by which it exerts its action is through modulating
the estrogen response systems. Because estrogen is involved in male reproductive biology, we investigated the
effect of trans-resveratrol on testis and spermatogenesis. Adult male rats were divided into 2 groups. The treated
group was administered by gavage 20 mg/(kg ? d) of trans-resveratrol suspended in 10 g/L of carboxymethylcel-
lulose for 90 d, whereas the control group received only carboxymethylcellulose during the same period. The
relative weight of testes did not differ between the groups. However, the diameter of the seminiferous tubules was
significantly reduced from 437.5 ? 0.1 ?m in the controls to 310.9 ? 0.1 ?m in the resveratrol–treated rats. This
decrease was accompanied by a significant increase in tubular density, from 3.20 ? 0.18 in controls to 6.58 ? 0.18
tubules/mm2in the treated group. Moreover, sperm counts were significantly greater in the resveratrol-treated rats
(24.8 ? 3.30 ? 107) than in the control group (14.1 ? 0.80 ? 107), but sperm quality did not differ. Serum
concentrations of gonadotrophins and testosterone were significantly higher in the resveratrol-treated group. We
identified a novel activity of trans-resveratrol. The daily oral administration of this phytochemical to adult male rats
enhanced sperm production by stimulating the hypothalamic-pituitary-gonadal axis, without inducing adverse
effects.J. Nutr. 135: 757–760, 2005.
trans-Resveratrol was reported to have health benefits including anticarcinogenic effects and pro-
● trans-resveratrol ● phytochemical ● spermatogenesis ● testis ● rats
trans-Resveratrol (trans-3,4?,5-trihydroxystilbene) is a nat-
ural antioxidant that is widely consumed in the Mediterranean
Diet in the form of peanuts, grapes, and wine. The interest in
compounds present in wine increased when epidemiologic
studies indicated an inverse correlation between red wine
consumption and the incidence of cardiovascular disease. This
finding prompted considerable interest in the possible effects
of trans-resveratrol, leading to the description of several ben-
eficial effects on health. In addition of being an antioxidant
and a vasorelaxing agent, it modulates lipoprotein metabolism,
inhibits platelet aggregation, and exerts cancer chemopreven-
tive and therapeutic activity (1,2). In eliciting these actions,
trans-resveratrol triggers a variety of established cellular and
molecular effectors, the most remarkable of which is the es-
trogen response systems. Given the structural similarities of
trans-resveratrol to diethylstilbestrol (DES)3and estradiol, and
its activity as a modulator of the estrogen-response systems, it
has been classified as a phytoestrogen (1,2).
Estrogens, which are traditionally considered female hor-
mones, are also involved in the male reproductive system,
which is classically thought to be controlled mainly by andro-
gen hormones and their receptors. Estrogens, derived either
from local aromatization of androgens or produced by the
testes, can exert feedback action on the neuroendocrine com-
ponents of the male reproductive axis. They also have para-
crine actions within the testes (3–5). trans-Resveratrol mod-
ulates the estrogen-response systems and may therefore be
involved in male reproduction. Consequently, the aim of the
present study was to investigate the effect of a 90-d trans-
resveratrol treatment [20 mg/(kg ? d)] on testes and spermato-
genesis of adult rats. Because spermatogenesis involves a
complex interplay between the structural elements of the
testes and the endocrine system, the serum concentrations of
the reproductive hormones, luteinizing hormone (LH), folli-
cle-stimulating hormone (FSH), and testosterone were mea-
1Supported by grant AGL2000-0918 from the Ministerio de Ciencia y Tec-
nologı ´a and grant 2001-SGR-00142 from the Generalitat de Catalunya, Spain.
E.G.-P. was a recipient of a Beca FPI from Ministerio de Ciencia y Tecnologı ´a and
T.M. was a recipient of a Beca para la Formacio ´n de Investigadores from the
Gobierno Vasco, Spain.
2To whom correspondence should be addressed. E-mail: firstname.lastname@example.org.
3Abbreviations used: AR, androgen receptor; DES, diethylstilbestrol; ER,
estrogen receptor; FSH, follicle stimulating hormone; LH, luteinizing hormone;
ROS, reactive oxygen species.
0022-3166/05 $8.00 © 2005 American Society for Nutritional Sciences.
Manuscript received 29 October 2004. Initial review completed 24 November 2004. Revision accepted 13 January 2005.
by on March 25, 2009
MATERIALS AND METHODS
Chemicals and reagents. trans-Resveratrol (Sigma, Tres Cantos)
was chemically pure. Before use, its purity was assessed by HPLC
coupled to a diode-array UV detector; a chromatogram that showed
a single peak at 306 nm, its maximum absorbance, was obtained. The
use of this detector allowed confirmation of the identity of the peak
by its spectrum (data not shown). Dose preparation, administration,
and sample treatments were performed in dim light to avoid photo-
chemical isomerization of trans-resveratrol to the cis form. All other
reagents were commercially available, analytical-grade chemicals.
Animals. Male Sprague-Dawley rats, weighting 210–220 g, were
purchased from breeding colonies (Harlan Ibe `rica) and quarantined
for 1 wk. They were housed in cages (3 rats/cage) at 22 ? 3°C, with
40–70% relative humidity and controlled lighting that provided a
12-h light:dark cycle. Water and a solid diet (Rodent Toxicology
Diet, B&K Universal) were freely available.4No traces of trans-
resveratrol were detected in the commercial diet or in the plasma
from control rats, as revealed by the analyses performed using the
method of Juan et al. (6). Rats were handled and killed following the
European Community guidelines for the care and management of
laboratory animals. The studies were approved by the Animal Exper-
imentation Research Committee of the University of Barcelona. All
rat manipulations were performed in the morning to minimize the
effects of circadian rhythm.
Experimental design and dose preparation. Rats were randomly
divided into 2 groups, a resveratrol group (n ? 12) and a control
group (n ? 12). The resveratrol group was orally administered, by
gavage, 20 mg/kg of trans-resveratrol at a constant volume of 10
mL/kg body weight, every day for 90 d. Because of its low solubility in
water, trans-resveratrol was suspended in 10 g/L carboxymethylcellu-
lose. The dose was adjusted according to the rat’s weight to ensure a
constant dose level and was freshly prepared immediately before each
administration. The control group was administered only carboxy-
methylcellulose during the same period. The dosage selected corre-
sponded to 1000 times the amount consumed by a 70-kg person who
drinks 250 mL of red wine a day containing 1.4 mg of trans-resvera-
trol, a dose that is not harmful to rats (7). A period of 90 d was chosen
to cover the complete spermatogenesis cycle of rats.
Skin, eyes, mucous membranes, respiratory system, autonomic and
central nervous system conditions, somatomotor pattern, and behav-
ior were examined daily. Body weight and food and water consumption
were recorded daily. The growth rate was calculated as the difference
between the final weight and the initial weight divided by 90 d.
At the end of the study, rats were food deprived overnight and
anesthetized with ketamine (90 mg/kg) and xylazine (10 mg/kg).
Blood samples were collected by cardiac puncture; 2 mL was trans-
ferred to a tube without anticoagulant for hormone analysis and 1 mL
was placed in EDTA for trans-resveratrol determination. Whole
blood was centrifuged at 1500 ? g (model TJ-6 centrifuge, rotor TH-4
with buckets, Beckman Coulter) for 15 min at room temperature.
Plasma trans-resveratrol and its conjugates. At 24 h after the
last oral administration, the trans-resveratrol concentration was mea-
sured using the method described by Juan et al. (6), which allows the
determination of the free compound as well as its sulfate and gluc-
uronide conjugates in plasma samples.
Testicular morphometry. Testes, epidydimae, and ducti deferens
were carefully dissected and the testes wet weight was recorded.
Results were expressed as testes weight relative to 100 g of body
weight. Subsequently, testes were fixed in 1.23 mol/L buffered form-
aldehyde pH 7.4 (Sigma Diagnostics) and kept at 4°C until analysis.
Testes were dehydrated in an alcohol gradient and placed in xylene.
They were then embedded in paraffin and cut into small sections (1
mm3). The tissue blocks were oriented so that the seminiferous
tubules could be sectioned transversely into 5-?m slices, thereby
giving round or rounded tubules, which were stained using the he-
matoxylin-eosin technique. Morphometrical measures were per-
formed on 5 distinct areas and a total of 250–300 seminiferous
tubules were measured in each rat (8). The diameter of the testicular
tubules was determined by projecting the slides at 50X; measurements
were systematically made from a set of 5 regions with presized areas.
The smallest minor axis diameter of each of the 5 randomly selected
sections of each rat was measured. Images were taken in a Nikon
Eclipse E800 optical microscope (Nikon Europe) linked to a digital
Sony 3CCD camera (Sony Inc. Europe). The digitalized images were
then processed using the ANALYSIS 2.1 imaging package (Soft-
Imaging Systems GmbH).
Testicular histology. Formaldehyde-fixed testes were embedded
in paraffin and sliced on silane-precoated slides (slice thickness: 3–4
?m). These slices were further deparaffined with xylol, and histolog-
ical observations were performed after staining using the hematoxy-
lin-eosin method. For long storage, slides were mounted using a
commercial mounting medium (Adh CLINIC©, Clinic Services).
The samples were observed under an Olympus microscope (model
BX50) coupled to a photographic camera.
Sperm counts and morphology. Sperm contained in the epidy-
dimae/ducti deferens complex from both testes of each rat were
released together into 1 mL of buffered formaldehyde. The sperm
count was measured in a hemocytometer chamber. A 10-?L aliquot
of sperm suspension was stained using the vital Eosin-Nigrosin stain-
ing technique (9). The percentage of sperm morphological abnormal-
ities was calculated in these stained spermatozoa after counting 200–
300 in an optical microscope at 1000X augmentations in a bright
field. The percentage of abnormalities was calculated, first as a total,
and then further classified in relation to the specific location of each
abnormality in the sperm cell. Consequently, total abnormalities
were classified as head abnormalities, neck and midpiece abnormali-
ties, distal cytoplasmic droplets, and tail abnormalities.
Hormone assays. After centrifugation, serum was removed from
the clot, immediately frozen in liquid N2and stored at ?80°C until
analysis. The serum concentrations of LH, FSH, and testosterone
were measured by ELISA. Specific commercial kits for rats were used
to quantify FSH (RPN 2560), LH (RPN 2562) (Amersham) and
testosterone (EIA-1559, DRG Instruments) concentrations.
Results were expressed as means ? SEM.
Means were compared by the Student-Neuman-Keuls Test (STATIS-
TICA 6.0 for Windows software, StatSoft). Differences with P ? 0.05
were considered significant.
Body weight and food and water consumption.
Resveratrol did not have adverse effects during the experimen-
tal period. The body weight gain of the resveratrol-treated
group did not differ from that of the controls; both groups grew
steadily throughout the study. Thus, the body weight of the
resveratrol group increased from 219 ? 2 g (n ? 12) on d 1 to
397 ? 9 g on d 90, whereas in the control group, it increased
from 216 ? 2 g (n ? 12) on d 1 to 394 ? 6 g (n ? 12) on d
90. Furthermore, the consumption of food and water did not
differ between the 2 groups.
Plasma trans-resveratrol and its conjugates. The trans-
resveratrol plasmatic concentrations determined by HPLC
with diode-array detection was 66.7 ? 12.0 nmol/L. Free
trans-resveratrol was the only compound detected in plasma
24 h after the oral administration in food-deprived rats. No
traces of sulfate or glucuronide conjugates were found.
Testicular histology and morphometry. trans-Resveratrol
did not affect testicular gross anatomy or wet weight. The
relative weights of the testis in the control and resveratrol
groups were 0.52 ? 0.02 and 0.49 ? 0.01, respectively. More-
over, histological examination did not reveal microscopic le-
sions such as cytoarchitectural alterations or disorganization of
the tubular elements in either group (Fig. 1). However, struc-
tural changes in the density of the seminiferous tubules were
observed (Fig. 1). In the resveratrol group, the tubes were of
reduced diameter but of increased length, which doubled the
tubular density (P ? 0.05).
4The commercial diet contained (g/kg) 160.4 crude protein, 26.0 crude fiber,
467.4 carbohydrate, 29.5 lipid, and 42.2 ash.
JUAN ET AL.
by on March 25, 2009
Sperm counts and morphology. The sperm count was 76%
greater in treated rats than in controls (P ? 0.01) (Table 1).
Sperm abnormalities, including misshapen head, middle piece,
or principal piece and distal cytoplasmatic droplet were not
more frequent in resveratrol–treated rats (Table 1). Conse-
quently, the percentage of atypical forms was not affected by
the treatment, indicating that the overall sperm quality was
not impaired by trans-resveratrol.
Serum hormones. At the end of the study, serum concen-
trations of FSH, LH, and testosterone were greater in the
resveratrol-treated rats than in the control group (Table 1).
Our study is the first to describe a novel effect of trans-
resveratrol, namely, an increase in spermatozoa production in
healthy rats. Mammalian testes fulfill 2 main functions, the
synthesis of steroid hormones and the production of sperma-
tozoa, both of which are controlled by gonadotrophins and
testosterone as well as locally produced factors (10). Our
results show that these 2 functions were enhanced in male rats
by the daily oral administration of trans-resveratrol. These
changes were not accompanied by side effects related to the
compound. This lack of toxicity was not surprising because we
previously demonstrated that the oral administration of trans-
resveratrol at a dose of 20 mg/(kg ? d) for 28 d was not harmful
to male rats (7). Thus, the absence of symptoms observed through-
out the 90-d treatment, the lack of negative effects on devel-
opment, and the normal appearance of the vital organs in the
gross necropsy indicate that trans-resveratrol is nontoxic under
these experimental conditions. Moreover, our results are sub-
stantiated by a recent toxicological study that reports that the
exposure of male and female rats to a dose of 300 mg/(kg ? d)
of trans-resveratrol for 28 d did not have adverse effects (11).
The repeated oral administration of trans-resveratrol in-
duced a 71% decrease in the mean diameter of the seminifer-
ous tubules with a 100% increase in the testicular tubular
density. Together, these changes led to an overall increase in
the size of the spermatogenic tissue. Therefore, this enlarge-
ment may explain the increase in sperm production observed.
It is noteworthy that sperm collected from the epididymis had
matured properly because the morphological examination re-
vealed the same percentage of abnormalities between groups.
This is the first report of a stimulatory effect of trans-
resveratrol on the secretion of gonadotrophins, the major
endocrine regulators of spermatogenesis. The concentrations
of FSH, which acts within the tubules to stimulate spermato-
genesis, and LH, which signals the production of testosterone
in Leydig cells, were elevated in the resveratrol group com-
pared with the control rats. Testosterone, which is essential for
promoting spermatogenesis, was also enhanced. These results
indicate that the effect of trans-resveratrol on sperm count may
be caused by the hypophisary stimulation of testicular func-
tion. The endocrinal regulation of the hypothalamic-pituitary-
gonadal axis in the males is intricate and involves estradiol
and testosterone (12,13). The sites for the feedback regulation
include cells in the hypothalamus, which are in close proxim-
ity to gonadotrophin-releasing hormone neurons, and gonado-
trophins in the pituitary, which may respond directly to an-
drogens caused by the expression of the androgen receptor
(AR), whereas the presence of aromatase and the estrogen
receptor (ER) allows the conversion of androgens into estro-
gens, and the subsequent activation of ER signaling pathways
(12). A possible explanation for our findings could be attrib-
uted to the binding of trans-resveratrol to ER as a mixed weak
agonist/antagonist, without estrogenic properties (1,2,14,15).
Interestingly, we did not observe estrogenic activity of trans-
Sperm count, sperm morphology, and serum hormones of
control and trans-resveratrol-treated Sprague-Dawley male rats1
Sperm counts (?107)
Head abnormalities, %
Neck and midpiece
Tail abnormalities, %
14.1 ? 0.80 24.8 ? 3.30*
12.5 ? 1.21
0.62 ? 0.24
12.4 ? 1.13
0.60 ? 0.16
6.38 ? 0.98
1.78 ? 0.23
6.18 ? 0.70
1.54 ? 0.42
3.94 ? 0.694.68 ? 1.05
6.30 ? 0.77
109.0 ? 14.2
12.51 ? 1.35
10.65 ? 1.48*
244.3 ? 36.4*
33.25 ? 9.85*
1Values are means ? SEM, n ? 12. * Different from the control
group, P ? 0.05.
and those administered 20 mg/kg of trans-resveratrol (black bars) for
90 d. (A) Diameter; (B) tubular density; (C,D) histological appearance of
testis sections of control (C) and trans-resveratrol–treated rats (D).
Values are means ? SEM, n ? 12. *Different from control, P ? 0.05.
Seminiferous tubules of control male rats (white bars)
trans-RESVERATROL ENHANCES SPERMATOGENESIS
by on March 25, 2009
resveratrol. Our results indicate that the daily oral administra-
tion of 20 mg/kg for 90 d did not affect body weight or food
and water consumption in the treated group compared with
the control rats. Given that growth inhibition is a sensitive
indicator of estrogenic effects (16), this lack of reduction in
body weight in the treated rats substantiates that trans-resvera-
trol does not act as an estrogen agonist, which is in agreement
with other in vivo studies (17,18). Consequently, this com-
pound could interact with the ER, thus increasing the secre-
tion of gonadotrophins, leading in turn to an increment in
testosterone and sperm output. Furthermore, the effects de-
scribed above may have been enhanced through androgen
antagonism because trans-resveratrol also antagonizes andro-
gen action in prostate cancer cells by inhibiting AR activity
and suppressing AR expression (19,20).
The behavior of trans-resveratrol as a mixed weak agonist/
antagonist, without estrogenic properties, was also confirmed by
the absence of a clear estrogenic effect on testes, as opposed to
DES, a structural analog of trans-resveratrol and a potent estrogen
agonist. In contrast to trans-resveratrol, DES has deleterious
effects on the male reproductive tract. Rats treated with DES
show adverse effects that include a reduction in testicular
weight associated with impaired seminiferous tubular morphol-
ogy (21,22). Exposure to DES also impairs spermatogenesis,
which is substantiated at least in part by a reduced testosterone
concentration (22,23). Although structurally similar, the dis-
tinct activity of trans-resveratrol and DES can be explained by
subtle differences in their molecules. Compared with trans-res-
veratrol, DES lacks the 3-OH and 5-OH groups, but possesses a
to ER similar to that of estradiol and acts as a potent agonist,
which would explain the harmful effects described.
There is no information available in the literature on the
activity of trans-resveratrol on male testes and spermatogenesis
in either growing or adult rats. To our knowledge, the only 2
studies that we found were conducted in mice; one confirms
the lack of harmful effects on testes (25), whereas the other
describes a protective role in the male reproductive tract (26).
Consequently, no data were provided concerning the effect of
trans-resveratrol alone on spermatogenesis.
The effects of trans-resveratrol could also be mediated by
counteraction of constitutive oxidative stress within the seminif-
erous tubules. It was shown previously that during spermatogen-
esis stages VI–VIII in rats, there is a significant increase in
superoxide dismutase mRNA expression coinciding with the
presence in the tubules of elongated spermatids with excess cy-
toplasmatic retention. The cytoplasm was shown to produce high
levels of reactive oxygen species (ROS) (27). trans-Resveratrol
was found to be an effective scavenger of hydroxyl, superoxide,
and metal-induced radicals as well as having antioxidant abil-
ities in cells producing ROS. trans-Resveratrol exhibits a protec-
tive effect against lipid peroxidation in cell membranes and DNA
damage caused by ROS (1,2,14). Therefore, trans-resveratrol
could be acting by decreasing the steady-state levels of ROS and
proinflammatory factors in the seminiferous tubules, thus increasing
account for the increase in sperm output observed in healthy rats.
Our findings indicate that trans-resveratrol merits further re-
search because this phytochemical may constitute a promising
new compound for the treatment of male infertility. In Western
society, infertility is a growing problem; its causes are diverse, and
considerable effort is being made to provide effective therapy. In
the case of male infertility, antioxidants, anti-inflammatories,
androgens, and antiestrogens are some of the treatments used.
However, a truly effective treatment has yet to be found (27).
In conclusion, we describe here a novel activity of the phyto-
chemical trans-resveratrol. The daily oral administration of this
compound to adult male rats enhanced spermatogenesis through
the stimulation of the hypothalamic-pituitary-gonadal axis, and
did not have adverse effects. These findings indicate that trans-
resveratrol may provide a treatment for male infertility.
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