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BASIC RESEARCH
Effects of ovariectomy and resistance training on oxidative stress markers in the rat liver
Maria Fernanda Cury RodriguesI,
Corresponding author
mafe_cury@hotmail.com

Tel.: 55 16 9724-3669
, Uliana Sbeguen StotzerI, Mateus Moraes DomingosI, Rafael DeminiceII, Gilberto Eiji ShiguemotoI, Luciane Magri TomazI, Nuno Manuel Frade de SousaI, Fabiano Candido FerreiraI, Richard Diego LeiteIII, Heloisa Sobreiro Selistre-de-AraújoI, Alceu Afonso Jordão-JúniorII, Vilmar BaldisseraI, Sérgio Eduardo de Andrade PerezI
I Federal University of São Carlos, Department of Physiological Sciences, Laboratory Exercise of Physiology, São Carlos/SP, Brazil
II Universidade de São Paulo, Faculdade de Medicina de Ribeirão Preto, Laboratory of Nutrition and Metabolism, Ribeirão Preto/SP, Brazil
III University of Rio de Janeiro, Laboratory Biovasc, Rio de Janeiro/RJ, Brazil
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          "en" => "<p id="spara70" class="elsevierStyleSimplePara elsevierViewall">Levels of Vitamin E in liver homogenates 48h after the last session of the resistance training program in sham-operated sedentary &#40;Sham-Sed&#41;&#44; ovariectomized sedentary &#40;Ovx-Sed&#41;&#44; sham-operated resistance training&#44; &#40;Sham-Rt&#41; and ovariectomized resistance training &#40;Ovx-Rt&#41;&#46; Values are presented as mean &#177; SDM&#59; <span class="elsevierStyleItalic">p &#8804; 0&#46;05&#46; a</span> statistically significant difference compared to Sham-Sed&#46;</p>"
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    "textoCompleto" => "<span class="elsevierStyleSections"><span id="cesec10" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle60">INTRODUCTION</span><p id="para10" class="elsevierStylePara elsevierViewall">Menopause &#40;ovariectomy in animals&#41; is associated with increased food intake and body weight&#44; metabolic dysfunction&#44; loss of bone mineral density&#44; diabetes&#44; impairment of muscle function&#44; and increased inflammatory markers and oxidative stress &#40;OS&#41; &#40;<a class="elsevierStyleCrossRefs" href="#bib1">1&#8211;4</a>&#41;&#46; OS may result in lipid peroxidation of cell membranes&#44; damage to proteins&#44; and DNA and stellate cell activation&#46; These processes in turn lead to fibrosis&#44; chronic inflammation&#44; and apoptosis in the liver &#40;<a class="elsevierStyleCrossRef" href="#bib5">5</a>&#41;&#46; The measurement of multiple biomarkers&#44; such as enzymatic and non-enzymatic antioxidants&#44; lipid peroxidation&#44; and cellular redox balance&#44; is required to confirm the presence of oxidative stress in tissues&#46; One of the most commonly measured markers of the cellular redox status is the reduced glutathione &#40;GSH&#41; and oxidized glutathione &#40;GSSG&#41; ratio &#40;<a class="elsevierStyleCrossRef" href="#bib6">6</a>&#44;<a class="elsevierStyleCrossRef" href="#bib7">7</a>&#41;&#46;</p><p id="para20" class="elsevierStylePara elsevierViewall">Estrogen has anti-oxidative properties related to the A-ring phenolic hydroxyl group&#44; which acts as an effective electron donor and a free radical scavenger and interrupts the lipoperoxidation reaction &#40;<a class="elsevierStyleCrossRef" href="#bib8">8</a>&#41;&#46; Estrogen also protects females by up-regulating the expression of antioxidants such as glutathione peroxidase &#40;GSH-Px&#41; and manganese superoxide dismutase &#40;MnSOD&#41; &#40;<a class="elsevierStyleCrossRef" href="#bib9">9</a>&#44;<a class="elsevierStyleCrossRef" href="#bib10">10</a>&#41;&#46; The influence of metabolic disturbances caused by ovariectomy on the liver is of clinical interest because these disturbances may aggravate liver diseases&#44; such as non-alcoholic fatty liver disease &#40;NAFLD&#41; and non-alcoholic steatohepatitis &#40;NASH&#41;&#44; through the generation of reactive oxygen species &#40;ROS&#41; &#40;<a class="elsevierStyleCrossRef" href="#bib11">11</a>&#44;<a class="elsevierStyleCrossRef" href="#bib12">12</a>&#41;&#46;</p><p id="para30" class="elsevierStylePara elsevierViewall">Nguyen et al&#46; &#40;<a class="elsevierStyleCrossRef" href="#bib13">13</a>&#41; demonstrated that women with breast cancer treated with tamoxifen&#44; a non-steroidal anti-estrogenic drug&#44; developed a higher level of intra-abdominal fat and increased incidence of hepatic diseases&#46; Yasuda et al&#46; &#40;<a class="elsevierStyleCrossRef" href="#bib14">14</a>&#41; analyzed the effect of estradiol treatment on hepatic fibrosis in young male and female rats and found that estradiol treatment effectively promoted a dose-dependent suppression of hepatic fibrosis&#44; with reduced collagen content and lower levels of protocollagen type I and II mRNAs&#46; Nevertheless&#44; hormone replacement therapy &#40;HRT&#41; is not universally accepted&#44; mainly due to its contraindication in some patients&#44; low compliance&#44; fear of and aversion to the side effects&#44; long-term risks of some types of cancer &#40;<a class="elsevierStyleCrossRef" href="#bib15">15</a>&#41;&#44; and risk of myocardial infarction &#40;<a class="elsevierStyleCrossRef" href="#bib16">16</a>&#41;&#46;</p><p id="para40" class="elsevierStylePara elsevierViewall">There is evidence that regular physical activity up-regulates the anti-oxidative defense system&#44; attenuates the age-related increase in the concentration of cellular ROS in the rat liver&#44; and confers protection against oxidative stress-associated diseases &#40;<a class="elsevierStyleCrossRef" href="#bib17">17</a>&#41;&#46; However&#44; the beneficial effects of exercise are lost upon exhaustion&#46; Due to limited metabolic adaptation&#44; strenuous exercise could generate increased levels of ROS and lead to a down-regulation of the antioxidative defense system &#40;<a class="elsevierStyleCrossRef" href="#bib18">18</a>&#41;&#46;</p><p id="para50" class="elsevierStylePara elsevierViewall">Among the exercise interventions discussed in the literature&#44; resistance training appears to be the most efficient modality in the attenuation of sarcopenia&#44; osteopenia&#44; hepatic steatosis&#44; and body composition changes promoted by menopause and ovariectomy &#40;<a class="elsevierStyleCrossRef" href="#bib19">19</a>&#44;<a class="elsevierStyleCrossRef" href="#bib20">20</a>&#41;&#46; Previous studies conducted by our group have shown that resistance training is able to decrease fat accumulation and deposition in the liver of ovariectomized &#40;Ovx&#41; rats &#40;<a class="elsevierStyleCrossRef" href="#bib21">21</a>&#41; and to reduce the gene expression of molecules related to lypogenesis &#40;<a class="elsevierStyleCrossRef" href="#bib22">22</a>&#41;&#46; Recent studies in humans have demonstrated that resistance training yields results similar to aerobic training with respect to the up-regulation of the antioxidative defense system&#44; reduction in lipid peroxidation&#44; and protection against oxidative damage based on serum analyses &#40;<a class="elsevierStyleCrossRef" href="#bib23">23</a>&#41;&#46; However&#44; there is evidence that high-intensity resistance exercise increases ROS production and causes oxidative damage &#40;<a class="elsevierStyleCrossRef" href="#bib24">24</a>&#41;&#46; Among other organs&#44; the liver is one of the most sensitive targets of exercise-induced oxidative stress &#40;<a class="elsevierStyleCrossRef" href="#bib25">25</a>&#41;&#44; but the effect of resistance training on OS in the liver of ovariectomized rats remains unclear&#46; Thus&#44; the aim of this study was to examine the effect of ovariectomy and resistance training on OS biomarkers in the liver of ovariectomized rats&#46; In view of the results described above&#44; our hypothesis was that resistance training may attenuate the oxidative stress caused by ovariectomy in the rat liver&#46;</p></span><span id="cesec20" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle70">METHODS</span><span id="cesec30" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle80">Animals</span><p id="para60" class="elsevierStylePara elsevierViewall">Thirty-two female Sprague&#8211;Dawley rats from the breeding colony of the State University of S&#227;o Paulo &#40;UNESP&#44; Araraquara&#44; SP&#44; Brazil&#41;&#44; with initial weights of approximately 220&#177;12 g&#44; were used in this study&#46; The animals had free access to water and chow and were kept in collective cages &#40;four rats per cage&#41; at a controlled temperature of 22&#177;2 &#176;C with a 12-h light&#47;12-h dark cycle &#40;light from 1 a&#46;m&#46; to 1 p&#46;m&#46;&#41;&#46; All of the animals were fed balanced food for laboratory rats and mice &#40;Biobase&#174;&#44; &#193;guas Frias&#44; Brazil&#41;&#44; and food intake was monitored daily during the experimental period&#46;</p><p id="para70" class="elsevierStylePara elsevierViewall">This study was approved by the Committee of Experimental Animals of the Federal University of S&#227;o Carlos &#40;Protocol no&#46; 008&#47;2010&#41;&#46; All animal procedures were conducted in accordance with the Guide for Care and Use of Laboratory Animals &#40;<a class="elsevierStyleCrossRef" href="#bib26">26</a>&#41;&#46;</p></span><span id="cesec40" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle90">Experimental groups</span><p id="para80" class="elsevierStylePara elsevierViewall">The rats were randomly distributed into the following four experimental groups &#40;eight animals per group&#41;&#58; &#40;I&#41; sham-operated sedentary &#40;Sham-Sed&#41;&#44; &#40;II&#41; ovariectomized sedentary &#40;Ovx-Sed&#41;&#44; &#40;III&#41; sham-operated resistance training &#40;Sham-Rt&#41;&#44; and &#40;IV&#41; ovariectomized resistance training &#40;Ovx-Rt&#41;&#46; The sedentary animals &#40;Sham-Sed and Ovx-Sed&#41; were kept in their cages throughout the entire experimental period without any type of exercise&#46; The ovaries were removed from the Ovx animals &#40;Ovx-Sed and Ovx-Rt&#41;&#46; The trained animals &#40;Sham-Rt and Ovx-Rt&#41; underwent a 10-week resistance training program&#44; which was initiated at the same time for each group and is described below&#46;</p></span><span id="cesec50" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle100">Ovariectomy and sham operation</span><p id="para90" class="elsevierStylePara elsevierViewall">Ovariectomies and sham operations were performed when the rats reached a weight of 250 g&#44; according to the technique described by Kalu &#40;<a class="elsevierStyleCrossRef" href="#bib27">27</a>&#41;&#46; For the surgery&#44; the rats were anesthetized with a mixture of ketamine-xylazine &#40;61&#46;5-7&#46;6 mg&#47;kg&#44; intraperitoneally&#41;&#46; The sham-operated rats underwent the surgical procedure&#44; but the ovaries were not removed&#46; The ovaries were removed from the Ovx animals&#46; All animals that underwent surgical procedures had 3 weeks of recovery&#44; and all animals were euthanized 92 days after the surgical procedure&#46;</p></span><span id="cesec60" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle110">Resistance training protocol</span><p id="para100" class="elsevierStylePara elsevierViewall">During the 10 weeks of resistance training&#44; climbing sessions were performed 3 times per week&#46; Initially&#44; the rats were adapted to the resistance training protocol&#44; which required them to climb a vertical ladder &#40;1&#46;1&#215;0&#46;18 m&#44; 2-cm grid&#44; 80&#176; incline&#41; with weights attached to their tails&#46; The size of the ladder required the animals to perform 8-12 movements per climb&#46; The load apparatus was attached to the tail by wrapping the proximal portion of the tail with a self-adhesive foam strip&#46; A Velcro strap was wrapped around the foam strip and fastened&#46; With the load apparatus attached to the tail&#44; each rat was placed at the bottom of the ladder and familiarized with climbing&#46; If necessary&#44; a stimulus was applied to the animal&#39;s tail with tweezers to initiate the movement&#46; At the top of the ladder&#44; the rats reached a housing chamber &#40;20&#215;20&#215;20 cm&#41;&#44; where they were allowed to rest for 120 s&#46; This procedure was repeated until the rats voluntarily climbed the ladder three consecutive times without stimuli&#46;</p><p id="para110" class="elsevierStylePara elsevierViewall">The first training session started three days after this familiarization and consisted of four to eight ladder climbs with progressively heavier loads&#46; For the initial climb&#44; each animal carried a load that was 75&#37; of its body mass&#46; Subsequently&#44; weight increments of 30 g were added until the load did not allow the animals to climb the entire length of the ladder&#46; The highest load successfully carried over the entire length of the ladder was considered the rat&#39;s maximal carrying capacity for that training session&#46;</p><p id="para120" class="elsevierStylePara elsevierViewall">The next training sessions consisted of four ladder climbs with 65&#44; 85&#44; 95&#44; and 100&#37; of the rat&#39;s previous maximal carrying capacity&#44; as determined in the previous session&#46; During subsequent ladder climbs&#44; an additional 30-g load was added until a new maximal carrying capacity was determined&#46; The resistance training protocol was adapted from that of Hornberger and Farrar &#40;<a class="elsevierStyleCrossRef" href="#bib28">28</a>&#41;&#44; according to the needs of the current investigation&#46;</p></span><span id="cesec70" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle120">Euthanasia and preparation of tissue samples</span><p id="para130" class="elsevierStylePara elsevierViewall">The animals were euthanized by decapitation 48 h after the last resistance exercise session&#46; Immediately post-mortem&#44; the livers were removed&#44; washed with saline solution&#44; weighed&#44; frozen in liquid nitrogen&#44; and stored at 80 &#176;C until analysis&#46; The right lobe of the liver was used for all analyses&#46; The liver tissue was homogenized in a phosphate buffer &#40;0&#46;1 mol&#47;L&#44; pH 7&#46;4&#41; using a Turrax dispenser &#40;MA 102 MINI E &#8211; Marconi Model&#41; and centrifuged for 15 min at 6&#44;500 &#215; g &#40;3000 rpm&#41;&#46; The supernatant was used for the measurement of thiobarbituric acid reactive substances &#40;TBARS&#41; and for catalase and superoxide dismutase &#40;SOD&#41; activity assays&#46;</p></span><span id="cesec80" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle130">Reduced and oxidized glutathione levels</span><p id="para140" class="elsevierStylePara elsevierViewall">Reduced and oxidized glutathione levels were measured in the liver as previously described &#40;<a class="elsevierStyleCrossRef" href="#bib29">29</a>&#41;&#46; The assay is based on the reaction of GSH with 5&#46;5&#8216; dithio-bis-2-nitrobenzoic acid &#40;DTNB&#44; Sigma&#44; cat&#46; no&#46; D-8130&#41;&#46; This reaction produces 5&#8216; thio&#8211;2&#8211;nitrobenzoic acid &#40;TNB&#41;&#44; which has a maximal absorbance at 412 nm&#44; and oxidized glutathione&#8211;TNB adducts &#40;GS&#8211;TNB&#41;&#46; The rate of TNB formation is proportional to the concentration of total glutathione in the sample&#46;</p><p id="para150" class="elsevierStylePara elsevierViewall">Liver samples &#40;100 mg&#47;mL for total glutathione measurements and 200 mg&#47;mL for oxidized glutathione measurements&#41; were homogenized in ice-cold 5&#37; metaphosphoric acid solution &#40;Sigma-Aldrich 239275&#41; and centrifuged at 3&#44;000 &#215; g and 4 &#176;C for 10 min&#46; The clear upper layer was kept at 0-4 &#176;C for the assay&#46; KPE buffer &#40;0&#46;1 M potassium phosphate buffer with 5 mM EDTA disodium salt&#44; pH 7&#46;5&#41; was used as a diluent to stock solution of DTNB &#40;0&#46;67 mg&#47;mL prepared in KPE&#41; and GR &#40;40 &#181;L of GR &#91;250 units&#47;mL&#93; diluted in 3mL of KPE&#41;&#46; A total of 20 &#181;L of each blank&#44; standard&#44; or experimental sample and 120 &#181;L of a mixture of equal volumes of DTNB stock solution &#40;60 &#181;L&#41; and GR stock solution &#40;60 &#181;L&#41; was added to each well&#44; and the reaction was allowed to continue for 30s&#46; The absorbance was measured at 412 nm in a microplate reader &#40;SpectraMax&#174; M5 with SoftMax&#174; Pro Data Acquisition &#38; Analysis software&#44; Molecular Devices&#44; Sunnyvale&#44; CA&#44; USA&#41; every 30 s for a total of 5 min after the addition of 60 &#956;L of 0&#46;67 mg&#47;mL &#946;-NADPH in KPE&#46; The rate of 2-nitro-5-thiobenzoic acid formation was used to calculate the total glutathione concentration from a GSH standard curve&#46; For the GSSG assay&#44; 2 &#956;L of 2-vinylpyridine &#40;Sigma-Aldrich no&#46; 13229-2&#41; was added to the 200 mL of liver homogenate&#44; and the reaction was allowed to continue for 2 h in the dark&#46; Subsequently&#44; 20 &#956;L of the homogenate was added to each well as described above&#46; The GSSG concentration was calculated as the rate of 2-nitro-5-thiobenzoic acid formation based on a GSSG standard curve&#46; The reduced GSH was calculated as the total glutathione - GSSG values&#44; and the results were expressed as mmol&#47;g tissue&#46; The intra-assay coefficient of variation for reduced and oxidized glutathione was 1&#46;6&#37;&#44; and the sensitivity of the assay was 7&#46;8 &#956;M&#47;g of tissue&#46;</p></span><span id="cesec90" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle140">Vitamin E</span><p id="para160" class="elsevierStylePara elsevierViewall">Vitamin E &#40;a-tocopherol&#41; was measured by HLPC as described by Jord&#227;o &#40;<a class="elsevierStyleCrossRef" href="#bib30">30</a>&#41;&#46; Liver tissue was deproteinized with ethanol and then extracted with hexane&#46; The evaporated organic layer was reconstituted with the mobile phase and injected using a 4&#46;6&#215;25-cm C-18-type Shim-pack CLC-ODS column &#40;Shimadzu&#44; Kyoto&#44; Japan&#41; and a 4 mm X 1-cm precolumn at a flow rate of 2&#46;0 mL&#47;min<span class="elsevierStyleSup">-1</span>&#44; and the absorbance was read at 292 nm in a UV-Vis detector&#46; The results were expressed as nmol&#47;g tissue&#46; The intra-assay coefficient of variation for the measurement of vitamin E was 4&#37;&#44; and the sensitivity of the assay was 0&#46;18 &#956;M&#47;g tissue&#46;</p></span><span id="cesec100" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle150">Catalase and superoxide dismutase activities</span><p id="para170" class="elsevierStylePara elsevierViewall">CAT activity was assessed by measuring the rate of hydrogen peroxide &#40;10 mmol&#183;L<span class="elsevierStyleSup">&#8211;1</span>&#41; absorbance at 240 nm &#40;<a class="elsevierStyleCrossRef" href="#bib31">31</a>&#41;&#44; with the activity expressed as U&#47;mg protein&#46; The intra-assay coefficient of variation for the measurement of CAT activity was less than 32&#46;5&#37;&#44; and the sensitivity was 0&#46;2 units&#47;mg protein&#46; SOD activity was determined using a commercial EIA kit from Cayman Chemical&#174; &#40;706002&#41;&#44; which uses the xanthine and hypoxanthine method&#44; with absorbance read at 460 nm&#46; The SOD activity was determined based on standards of known SOD concentrations &#40;U&#47;mL&#41;&#44; and the activity was expressed as U&#47;g tissue&#46; The intra-assay coefficient of variation for the SOD assay was 7&#46;2&#37;&#44; and the sensitivity was 0&#46;025 units&#47;g tissue&#46;</p></span><span id="cesec110" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle160">Lipid peroxidation</span><p id="para180" class="elsevierStylePara elsevierViewall">The TBARS assay was adapted from that of Costa &#40;<a class="elsevierStyleCrossRef" href="#bib32">32</a>&#41;&#46; The liver samples were mixed with 1 mL of a solution containing 15&#37; &#40;w&#47;v&#41; trichloroacetic acid&#44; 0&#46;38&#37; &#40;w&#47;v&#41; thiobarbituric acid&#44; and 0&#46;25 N HCl&#46; The mixture was heated at 100 &#176;C for 30 min&#46; The TBARS concentration was measured by the absorbance at 535 nm&#44; using 1&#44; 1&#44; 3&#44; 3-tetramethoxypropane &#40;Sigma-Aldrich&#41; as an external standard&#44; and the results were expressed as &#956;mol&#47;g protein&#46; The intra-assay coefficient of variation for the measurement of TBARS was 16&#46;9&#37;&#44; and the sensitivity was 0&#46;055 &#956;M&#47;g protein&#46;</p></span><span id="cesec120" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle170">Protein determination</span><p id="para190" class="elsevierStylePara elsevierViewall">Total proteins were measured using a commercially available kit &#40;LABTEST&#174;&#59; Labtest Diagn&#243;stica&#44; Lagoa Santa&#44; Minas Gerais&#44; Brazil&#41;&#46;</p></span><span id="cesec130" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle180">Isolation of RNA and quantitative real-time &#40;RT&#41; polymerase chain reaction &#40;PCR&#41;</span><p id="para200" class="elsevierStylePara elsevierViewall">Total RNA was extracted from frozen liver tissue using the Trizol reagent &#40;Invitrogen Corporation&#44; California&#44; USA&#41; according to the manufacturer&#39;s specifications&#46; The integrity and quality of the purified RNA were analyzed using formaldehyde denaturing agarose gel electrophoresis and by measuring the A<span class="elsevierStyleInf">260</span>-to-A<span class="elsevierStyleInf">280</span> ratio&#46; For the removal of genomic DNA&#44; total RNA &#40;1 &#956;g&#41; from each sample was treated with deoxyribonuclease I &#40;DNase I&#59; Invitrogen Corporation&#44; CA&#44; USA&#41; according to a standard protocol recommended by the manufacturer&#46; The treated RNA was reverse-transcribed into cDNA using M-MLV reverse transcriptase &#40;Promega Corporation&#44; Madison&#44; WI&#44; USA&#41;&#46; The quantitative RT-PCR employed 25 &#956;L-volume reactions containing SYBR Green PCR Master Mix &#40;Fermentas&#174;&#41;&#44; 20 ng of cDNA&#44; and 0&#46;5 &#956;M of each primer&#46; The gene-specific primers were purchased from Invitrogen Life Technologies&#44; Inc&#46;&#44; and are listed in <a class="elsevierStyleCrossRef" href="#tbl1">Table 1</a>&#46; The thermal cycling program consisted of 10 min at 95 &#176;C&#44; followed by 40 cycles of 94 &#176;C for 15 s&#44; 57-61 &#176;C for 30 s&#44; and 72 &#176;C for 60 s&#46; Following PCR&#44; the melting curve was analyzed to ensure that only one PCR product was amplified per reaction&#46;</p><elsevierMultimedia ident="tbl1"></elsevierMultimedia><p id="para210" class="elsevierStylePara elsevierViewall">Glyceraldehyde-3-phosphate dehydrogenase &#40;GAPDH&#41; was used as an endogenous control&#46; The relative expression of the quantitative RT-PCR products was determined using the &#916;&#916;Ct method&#44; which calculates the relative expression using the following equation&#58; fold induction &#61; 2<span class="elsevierStyleSup">-&#91;&#916;&#916;Ct&#93;</span>&#44; where Ct &#61; the threshold cycle &#40;i&#46;e&#46;&#44; the cycle number at which the sample&#39;s relative fluorescence surpasses the background fluorescence&#41; and &#916;&#916;Ct &#61; &#91;Ct gene of interest &#40;unknown sample&#41; &#8211; Ct GAPDH &#40;unknown sample&#41;&#93; &#8211; &#91;Ct gene of interest &#40;calibrator sample&#41; &#8211; Ct GAPDH &#40;calibrator sample&#41;&#93; &#40;<a class="elsevierStyleCrossRef" href="#bib33">33</a>&#41;&#46; The primers used in this study were as follows&#58; GSH-Px Fwd&#58; 5-GGGCAAGGTGCTGCTCATTG-3 and Rev&#58; 5-AGAGCGGGTGAGCCTTCTCA-3&#46; The results were normalized to the housekeeping gene GAPDH through the use of the following primers&#58; Fwd&#58; 5- GATGCTGGTGCTGAGTATGTCG-3 and Rev&#58; 5- GTGGTGCAGGATGCATTGCTGA-3&#46; The following primers are available on the NCBI website&#58; GSH-Px &#40;NM&#95;10336022&#41; and GAPDH &#40;NM&#95;017008&#46;3&#41;&#46; The intra-assay coefficient of variation for GSH-Px was 6&#46;6&#37;&#46;</p></span><span id="cesec140" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle190">Statistical analysis</span><p id="para220" class="elsevierStylePara elsevierViewall">All data are presented as the means and &#40;&#177;&#41; standard deviation of the mean &#40;SDM&#41;&#46; The normality and homoscedasticity of the data were first tested by the Shapiro-Wilk and Levene&#39;s tests&#44; respectively&#46; Two-way analysis of variance &#40;ANOVA&#41; was used to compare the variables of resistance training and ovariectomy&#46; Fisher&#39;s post-hoc test was used in the event of a significant &#40;<span class="elsevierStyleItalic">p&#8804;</span>0&#46;05&#41; ratio&#46; SPPS version 17&#46;0 &#40;Somers&#44; NY&#44; USA&#41; software was used&#44; and the sample size was determined using G&#8727;Power version 3&#46;1&#46;3 &#40;Kiel&#44; Germany&#41; with the following settings&#58; level of significance at <span class="elsevierStyleItalic">p</span> &#61; 0&#46;05&#44; power &#40;1&#8211;&#946;&#41; &#61; 0&#46;80&#44; and a large effect size &#40;f2 &#61; 0&#46;40&#41;&#46; Based on these a priori calculations&#44; the final sample size was n &#61; 39&#46; Due to methodological and logistical aspects&#44; the sample size used in the study was 32&#44; and the sample power was therefore reduced to 0&#46;74&#46;</p></span></span><span id="cesec150" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle200">RESULTS</span><p id="para230" class="elsevierStylePara elsevierViewall">Compared with the Sham-Sed rats&#44; the Ovx rats were found to have higher body weights and higher daily food intakes &#40;<span class="elsevierStyleItalic">p&#8804;</span>0&#46;05&#41;&#46; The Ovx rats also had lower femur and uterus weights than the Sham-Sed rats &#40;15&#37;&#59; <span class="elsevierStyleItalic">p</span> &#61; 0&#46;032 and 77&#37;&#59; <span class="elsevierStyleItalic">p &#61;</span> 0&#46;001&#44; respectively&#41;&#46; However&#44; the Sham-Rt group exhibited a significantly higher femur weight compared with the Sham-Sed &#40;8&#37;&#59; <span class="elsevierStyleItalic">p &#61;</span> 0&#46;035&#41; and Ovx-Rt &#40;17&#37;&#59; <span class="elsevierStyleItalic">p &#61;</span> 0&#46;036&#41; groups&#46; In addition&#44; the Ovx-Rt group showed higher femur weights &#40;9&#37;&#59; <span class="elsevierStyleItalic">p &#61;</span> 0&#46;028&#41; than the Ovx-Sed group &#40;<a class="elsevierStyleCrossRef" href="#tbl1">Table 1</a>&#41;&#46;</p><p id="para240" class="elsevierStylePara elsevierViewall">No group x time interaction was observed for the maximal workload during the 10-week training period&#44; indicating that the maximal load carrying capacity increased in a similar manner for both the Sham-Rt and Ovx-Rt groups during the training period &#40;<a class="elsevierStyleCrossRef" href="#fig1">Figure 1</a>&#41;&#46;</p><elsevierMultimedia ident="fig1"></elsevierMultimedia><p id="para250" class="elsevierStylePara elsevierViewall">The GSH levels&#44; GSSG levels&#44; and GSH&#47;GSSH ratios are shown in <a class="elsevierStyleCrossRef" href="#fig2">Figures 2A&#44; B&#44; and C</a>&#44; respectively&#46; The Ovx-Sed group displayed lower GSH levels than the Sham-Sed group &#40;49&#37;&#44; <span class="elsevierStyleItalic">p&#8804;</span>0&#46;0005&#59; <a class="elsevierStyleCrossRef" href="#fig2">Figure 2A</a>&#41;&#46; Similarly&#44; the Ovx-Rt group had lower GSH levels than the Sham-Sed group &#40;62&#37;&#44; <span class="elsevierStyleItalic">p&#8804;</span>0&#46;005&#59; <a class="elsevierStyleCrossRef" href="#fig2">Figure 2A</a>&#41;&#46;</p><elsevierMultimedia ident="fig2"></elsevierMultimedia><p id="para260" class="elsevierStylePara elsevierViewall">The GSSG levels were significantly higher in the Sham-Rt group than in the Sham-Sed group &#40;38&#37;&#44; <span class="elsevierStyleItalic">p</span> &#61; 0&#46;021&#59; <a class="elsevierStyleCrossRef" href="#fig2">Figure 2B</a>&#41;&#44; while the GSSG levels were significantly lower in the Ovx-Sed group than in the Sham-Sed group &#40;32&#37;&#44; <span class="elsevierStyleItalic">p</span> &#61; 0&#46;032&#59; <a class="elsevierStyleCrossRef" href="#fig2">Figure 2B</a>&#41;&#46; In addition&#44; the Ovx-Rt group had lower GSSG levels than the Sham-Rt group &#40;56&#37;&#44; <span class="elsevierStyleItalic">p</span> &#61; 0&#46;022&#59; <a class="elsevierStyleCrossRef" href="#fig2">Figure 2B</a>&#41;&#46;</p><p id="para270" class="elsevierStylePara elsevierViewall">The GSSG levels were significantly lower in the Ovx-Sed and Ovx-Rt groups than in the Sham-Sed group &#40;32&#37;&#44; <span class="elsevierStyleItalic">p</span> &#61; 0&#46;036 and 39&#37;&#44; <span class="elsevierStyleItalic">p</span> &#61; 0&#46;043&#44; respectively&#41; &#40;<a class="elsevierStyleCrossRef" href="#fig2">Figure 2B</a>&#41;&#46; Furthermore&#44; the GSSG levels were significantly higher in the Sham-Rt group compared with the other groups &#40;<span class="elsevierStyleItalic">p&#8804;</span>0&#46;05&#59; <a class="elsevierStyleCrossRef" href="#fig2">Figure 2B</a>&#41;&#46;</p><p id="para280" class="elsevierStylePara elsevierViewall">The GSH&#47;GSSG ratio was lower in the Ovx-Sed and Sham-Rt groups than in the Sham-Sed group &#40;28&#37;&#44; <span class="elsevierStyleItalic">p</span>&#8804;0&#46;005 and 19&#37;&#44; <span class="elsevierStyleItalic">p</span> &#61; 0&#46;001&#44; respectively&#59; <a class="elsevierStyleCrossRef" href="#fig2">Figure 2C</a>&#41;&#46; The GSH&#47;GSSG ratio was lower in the Ovx-Rt group than in the Sham-Rt group &#40;14&#37;&#44; <span class="elsevierStyleItalic">p</span> &#61; 0&#46;28&#59; <a class="elsevierStyleCrossRef" href="#fig2">Figure 2C</a>&#41;&#46;</p><p id="para290" class="elsevierStylePara elsevierViewall">The RT-PCR results demonstrated lower gene expression of GSH-Px in the Ovx-Sed group compared with the Sham-Sed group &#40;49&#37;&#44; <span class="elsevierStyleItalic">p</span> &#61; 0&#46;032&#59; <a class="elsevierStyleCrossRef" href="#fig2">Figure 2D</a>&#41;&#46; In contrast&#44; the gene expression of GSH-Px was higher in the Ovx-Rt group than in the Ovx-Sed group &#40;90&#37;&#59; <span class="elsevierStyleItalic">p</span> &#61; 0&#46;043&#59; <a class="elsevierStyleCrossRef" href="#fig2">Figure 2D</a>&#41;&#46;</p><p id="para300" class="elsevierStylePara elsevierViewall">The Sham-Rt group was found to have a higher mean concentration of the lipid peroxidation marker TBARS than the Sham-Sed group &#40;30&#37;&#59; <span class="elsevierStyleItalic">p&#8804;</span>0&#46;0005&#59; <a class="elsevierStyleCrossRef" href="#fig3">Figure 3</a>&#41;&#44; while higher levels of TBARS were observed in the Ovx-Rt group compared with the Ovx-Sed group &#40;23&#37;&#44; <span class="elsevierStyleItalic">p</span> &#61; 0&#46;004&#59; <a class="elsevierStyleCrossRef" href="#fig3">Figure 3</a>&#41;&#46; However&#44; no statistically significant differences in the concentrations of TBARS were observed between the Ovx-Sed and Sham-Sed groups&#46;</p><elsevierMultimedia ident="fig3"></elsevierMultimedia><p id="para310" class="elsevierStylePara elsevierViewall">The concentration of vitamin E was found to be lower in the Ovx-Sed&#44; Sham-Rt&#44; and Ovx-Rt groups compared with the Sham-Sed group &#40;45&#37;&#44; <span class="elsevierStyleItalic">p&#8804;</span>0&#46;0005&#59; 35&#37;&#44; <span class="elsevierStyleItalic">p&#8804;</span>0&#46;0005 and 41&#37;&#44; <span class="elsevierStyleItalic">p&#8804;</span>0&#46;0005&#44; respectively&#59; <a class="elsevierStyleCrossRef" href="#fig4">Figure 4</a>&#41;&#46; No statistically significant differences in enzymatic SOD and CAT activity were observed between the groups &#40;<a class="elsevierStyleCrossRef" href="#fig5">Figures 5A and B</a>&#41;&#46;</p><elsevierMultimedia ident="fig4"></elsevierMultimedia><elsevierMultimedia ident="fig5"></elsevierMultimedia></span><span id="cesec160" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle210">DISCUSSION</span><p id="para320" class="elsevierStylePara elsevierViewall">Regular exercise induces adaptations such as the hepatic down-regulation of ROS production &#40;<a class="elsevierStyleCrossRef" href="#bib34">34</a>&#44;<a class="elsevierStyleCrossRef" href="#bib35">35</a>&#41;&#46; However&#44; the beneficial effects of exercise are lost with exhaustion &#40;<a class="elsevierStyleCrossRef" href="#bib18">18</a>&#44;<a class="elsevierStyleCrossRef" href="#bib36">36</a>&#41;&#46; In the present study&#44; resistance training did not reverse the hepatic oxidative stress promoted by ovariectomy&#46; Additionally&#44; this resistance training protocol induced negative changes in the hepatic anti-oxidative&#47;oxidative balance&#44; as evidenced by the increased levels of TBARS &#40;30&#37;&#41;&#44; decreased GSH&#47;GSSG ratios &#40;19&#37;&#41;&#44; and decreased vitamin E concentrations &#40;35&#37;&#41;&#46;</p><p id="para330" class="elsevierStylePara elsevierViewall">Oxidative stress is an important mechanism involved in the progression of several ovariectomy-related pathogeneses&#44; such as NAFLD and NASH &#40;<a class="elsevierStyleCrossRef" href="#bib10">10</a>&#44;<a class="elsevierStyleCrossRef" href="#bib37">37</a>&#41;&#46; Regular physical exercise can promote antioxidative defenses and reduce OS &#40;<a class="elsevierStyleCrossRef" href="#bib23">23</a>&#44;<a class="elsevierStyleCrossRef" href="#bib34">34</a>&#41;&#46; Radak et al&#46; &#40;<a class="elsevierStyleCrossRef" href="#bib17">17</a>&#41; showed that regular treadmill running for 8 weeks resulted in a 40&#37; increase in the maximal oxygen uptake and caused a down-regulation of ROS production in the liver of trained rats&#46; However&#44; Ogonovszky et al&#46; &#40;<a class="elsevierStyleCrossRef" href="#bib18">18</a>&#41; demonstrated that the overtraining model&#44; consisting of 1 hour of swimming&#47;day for 5 days&#47;week for 6 weeks&#44; promoted oxidative damage to the hepatic nuclear DNA without influencing lipids or proteins&#46; In that study&#44; oxidative damage to the nuclear DNA was measured by 8-hydroxydeoxyguanosine &#40;8-OHdG&#41;&#46;</p><p id="para340" class="elsevierStylePara elsevierViewall">Recent studies have indicated that resistance training is beneficial for skeletal muscle&#44; bones&#44; and the liver &#40;<a class="elsevierStyleCrossRef" href="#bib3">3</a>&#44;<a class="elsevierStyleCrossRefs" href="#bib20">20&#8211;22</a>&#41;&#46; However&#44; limited data are available regarding oxidative stress in the liver after this type of training&#46; Margonis et al&#46; &#40;<a class="elsevierStyleCrossRef" href="#bib24">24</a>&#41; used a resistance training protocol with progressively increased and decreased volume&#47;intensity in humans and demonstrated that overtraining induced a marked increase in oxidative stress biomarkers&#44; which&#44; in some cases&#44; was proportional to the training load&#46; Although the magnitude of the adaptation appears to be partly dependent on exercise intensity in mice and humans &#40;<a class="elsevierStyleCrossRef" href="#bib24">24</a>&#44;<a class="elsevierStyleCrossRef" href="#bib38">38</a>&#41;&#44; a recent study with human subjects indicated that resistance training performed at intensities corresponding to hypertrophy and strength training yielded protective effects against OS similar to those provided by aerobic exercises&#44; and these effects did not seem to be dependent on the training intensity &#40;<a class="elsevierStyleCrossRef" href="#bib23">23</a>&#41;&#46;</p><p id="para350" class="elsevierStylePara elsevierViewall">Our study demonstrated that the ovariectomy procedure decreased the hepatic GSH levels in the sedentary and trained groups &#40;<a class="elsevierStyleCrossRef" href="#fig2">Figure 2A</a>&#41;&#46; Glutathione is an antioxidant that protects mammalian cells against oxidative processes&#44; and the liver is the largest producer and exporter of this antioxidant &#40;<a class="elsevierStyleCrossRef" href="#bib39">39</a>&#41;&#46; The relationship between the two glutathione isoforms &#40;GSH and GSSG&#41; is considered an index of the cellular redox state&#59; under increased OS conditions&#44; the GSH&#47;GSSG ratio decreases &#40;<a class="elsevierStyleCrossRef" href="#bib39">39</a>&#41;&#46; In our study&#44; resistance training and ovariectomy promoted a strong alteration of the liver redox state&#44; as evidenced by the decreased GSH&#47;GSSG ratio &#40;<a class="elsevierStyleCrossRef" href="#fig2">Figure 2C</a>&#41;&#46;</p><p id="para360" class="elsevierStylePara elsevierViewall">Furthermore&#44; ovariectomy markedly decreased the mRNA levels of GSH-Px &#40;<a class="elsevierStyleCrossRef" href="#fig2">Figure 2D</a>&#41;&#46; Estradiol appears to up-regulate the expression of Mn-SOD and GSH-Px by interacting with the membrane estrogen receptor and activating MAP kinases and NF-kappaB in the mammary gland tumor cell line MCF-7 &#40;<a class="elsevierStyleCrossRef" href="#bib56">56</a>&#41;&#46; However&#44; resistance training also led to an increase in the GSH-Px mRNA levels &#40;<a class="elsevierStyleCrossRef" href="#fig2">Figure 2D</a>&#41;&#44; corroborating the results of Chang et al&#46; &#40;<a class="elsevierStyleCrossRef" href="#bib57">57</a>&#41;&#44; who observed similar effects with endurance exercise&#46;</p><p id="para370" class="elsevierStylePara elsevierViewall">Some types of training&#44; such as moderate-intensity training&#44; have been shown to provide protection against oxidative stress by increasing the hepatic gene expression and activity of Mn-SOD and GPx and by increasing the GSH levels &#40;<a class="elsevierStyleCrossRef" href="#bib17">17</a>&#44;<a class="elsevierStyleCrossRef" href="#bib58">58</a>&#41;&#46; However&#44; the increase in hepatic GSH-Px gene expression induced by our resistance training protocol was not accompanied by increased GSH levels&#44; which are necessary to maintain the optimal catalytic condition of GSH-Px&#46;</p><p id="para380" class="elsevierStylePara elsevierViewall">The literature has clearly demonstrated that changes in the glutathione levels caused by ovariectomy are accompanied by increased lipid peroxidation and increased MDA production &#40;<a class="elsevierStyleCrossRef" href="#bib23">23</a>&#41;&#46; However&#44; the ovariectomy procedure was not found to promote an increase in the MDA levels in this study &#40;<a class="elsevierStyleCrossRef" href="#fig3">Figure 3</a>&#41;&#46; This result could be explained by the fact that the liver has a high mitotic potential and increased capacity for DNA repair &#40;<a class="elsevierStyleCrossRef" href="#bib40">40</a>&#41;&#46;</p><p id="para390" class="elsevierStylePara elsevierViewall">Resistance training did increase the concentration of TBARS&#44; regardless of the ovarian status &#40;<a class="elsevierStyleCrossRef" href="#fig3">Figure 3</a>&#41;&#46; These results demonstrated that the resistance training protocol used in our study induced an oxidative process in the liver of these animals&#46; The conversion of xanthine dehydrogenase &#40;XD&#41; into xanthine oxidase &#40;XO&#41; is among the mechanisms involved in the induction of oxidative damage in the liver during exercise&#46; XO reduces O<span class="elsevierStyleInf">2</span> instead of NADP<span class="elsevierStyleSup">&#43;</span>&#44; thereby producing superoxide ions &#40;O<span class="elsevierStyleInf">2</span><span class="elsevierStyleSup">&#8226;</span>&#41; &#40;<a class="elsevierStyleCrossRef" href="#bib4">4</a>&#41;&#46; Da Silva et al&#46; &#40;<a class="elsevierStyleCrossRef" href="#bib36">36</a>&#41; investigated the effects of different physical exercise protocols on OS markers in murine livers and found that continuous running and downhill running for 45 min daily&#44; 5 days&#47;week for 8 weeks&#44; decreased the lipid peroxidation levels&#44; as assessed by TBARS&#46; This result demonstrated that the liver did not suffer damage from localized eccentric contractions&#46; In addition&#44; Rosety-Rodriguez et al&#46; &#40;<a class="elsevierStyleCrossRef" href="#bib34">34</a>&#41; reported that an 8-week moderate swimming training program reduced oxidative damage induced by emotional stress&#46;</p><p id="para400" class="elsevierStylePara elsevierViewall">Oxidative stress and hepatic damage have been negatively correlated with vitamin E levels &#40;<a class="elsevierStyleCrossRef" href="#bib42">42</a>&#41;&#46; Vitamin E is a fat-soluble antioxidant in the plasma that is primarily stored in the liver and performs the function of eliminating peroxyl radicals &#40;ROO&#41; &#40;<a class="elsevierStyleCrossRef" href="#bib43">43</a>&#41;&#46; Moreover&#44; several studies have clearly shown that serum levels of vitamin E are significantly reduced in patients with NAFLD &#40;<a class="elsevierStyleCrossRef" href="#bib42">42</a>&#41;&#46; Zamin Jr&#46; et al&#46; &#40;<a class="elsevierStyleCrossRef" href="#bib44">44</a>&#41; found that vitamin E supplementation significantly reduced OS in the liver and concluded that vitamin E plays an important therapeutic role in preventing diseases such as NASH&#46; In our study&#44; the reduced vitamin E concentrations in both groups of ovariectomized animals &#40;sedentary and trained&#41; can be explained by the low concentration of GSH&#44; which reduces dihydroascorbate into ascorbate and thereby ensures that vitamin E is recycled &#40;<a class="elsevierStyleCrossRef" href="#fig4">Figure 4</a>&#41;&#46; Furthermore&#44; our resistance training protocol did not reverse the effects of ovariectomy and actually reduced the vitamin E concentrations in the trained sham group &#40;Sham-Rt&#41;&#46; A possible reason for the reduced vitamin E levels is the replenishment process&#44; which involves the redistribution of vitamin E from the liver to the muscles during exercise &#40;<a class="elsevierStyleCrossRef" href="#bib45">45</a>&#41;&#46; This result is in agreement with the findings of Aikawa et al&#46; &#40;<a class="elsevierStyleCrossRef" href="#bib46">46</a>&#41;&#44; who provided clear evidence of vitamin E depletion in the liver of trained rats under normal and reduced vitamin E diets&#46;</p><p id="para410" class="elsevierStylePara elsevierViewall">Another physiological consequence of ovariectomy and menopause is an increase in CAT enzymatic activity and a decrease in the enzymatic activities of SOD and GSH-Px &#40;<a class="elsevierStyleCrossRef" href="#bib47">47</a>&#44;<a class="elsevierStyleCrossRef" href="#bib48">48</a>&#41;&#46; These phenomena can be explained by the fact that when OS increases&#44; SOD activity decreases&#44; due to the irreversible inactivation of its H<span class="elsevierStyleInf">2</span>O<span class="elsevierStyleInf">2</span> product&#46; Meanwhile&#44; the CAT activity increases to eliminate H<span class="elsevierStyleInf">2</span>O<span class="elsevierStyleInf">2</span>&#46;</p><p id="para420" class="elsevierStylePara elsevierViewall">Although the results of previous training studies are sometimes conflicting&#44; it is clear that regular physical activity leads to an increase in antioxidant enzyme activity&#44; especially in tissues such as skeletal muscle &#40;<a class="elsevierStyleCrossRef" href="#bib24">24</a>&#44;<a class="elsevierStyleCrossRef" href="#bib48">48</a>&#41;&#46; It has been established that strenuous physical exercise promotes oxidative stress due to the generation of oxygen free radicals&#44; which stimulate the up-regulation of antioxidant enzyme activity in skeletal muscle&#46; The effect of chronic exercise on OS in the liver and on antioxidative systems has been investigated in several studies&#44; the results of which have shown that exercise training leads to reduced CAT activity &#40;<a class="elsevierStyleCrossRef" href="#bib50">50</a>&#44;<a class="elsevierStyleCrossRef" href="#bib51">51</a>&#41; and increased SOD activity &#40;<a class="elsevierStyleCrossRef" href="#bib52">52</a>&#41;&#46; This imbalance between SOD and CAT activity could be attributed to higher levels of pro-inflammatory molecules &#40;<a class="elsevierStyleCrossRef" href="#bib53">53</a>&#41;&#44; which are known to increase SOD mRNA expression &#40;<a class="elsevierStyleCrossRef" href="#bib54">54</a>&#41;&#46; In addition&#44; CAT activity may be inhibited by the superoxide produced during exercise &#40;<a class="elsevierStyleCrossRef" href="#bib55">55</a>&#41;&#46; In our study&#44; no changes in SOD and CAT activity could be attributed to the physiological adaptation induced by OS &#40;<a class="elsevierStyleCrossRef" href="#fig5">Figures 5A&#44; B</a>&#41;&#46;</p><p id="para430" class="elsevierStylePara elsevierViewall">In summary&#44; the relevant findings and limitations reported in this paper are as follows&#46; The resistance training program used in this study failed to attenuate the hepatic oxidative damage caused by ovariectomy and increased the OS in the liver&#46; One limitation of the present study was the lack of a group of ovariectomized animals treated with hormone replacement therapy to confirm the negative effects of resistance training on oxidative stress biomarkers in the liver&#46; Further studies should be conducted to more accurately elucidate and characterize the psychological and metabolic stress imposed by this resistance training protocol in rats and humans&#46;</p></span><span id="cesec170" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle220">AUTHOR CONTRIBUTIONS</span><p id="para440" class="elsevierStylePara elsevierViewall">Rodrigues MF&#44; Stotzer US&#44; and Domingos MM designed the research project and performed the experiments&#46; Deminice R and Jord&#227;o-J&#250;nior AA supervised the analysis of experimental results&#46; Shiguemoto GE&#44; Selistre-de-Ara&#250;jo HS&#44; and Baldissera V supervised the research project&#46; Tomaz LM and Ferreira FC participated in the analysis of experimental results&#46; Sousa NM and Leite RD participated in the analysis of experimental results and statistical analysis&#46; Perez SE supervised the research project&#46;</p></span></span>"
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              "titulo" => "Ovariectomy and sham operation"
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              "titulo" => "Resistance training protocol"
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              "titulo" => "Euthanasia and preparation of tissue samples"
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              "titulo" => "Reduced and oxidized glutathione levels"
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              "titulo" => "Vitamin E"
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              "titulo" => "Catalase and superoxide dismutase activities"
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              "titulo" => "Lipid peroxidation"
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              "titulo" => "Protein determination"
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              "titulo" => "Isolation of RNA and quantitative real-time &#40;RT&#41; polymerase chain reaction &#40;PCR&#41;"
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              "titulo" => "Statistical analysis"
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          "titulo" => "RESULTS"
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          "titulo" => "DISCUSSION"
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          "titulo" => "ACKNOWLEDGMENTS"
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    "fechaRecibido" => "2013-01-31"
    "fechaAceptado" => "2013-04-28"
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            0 => "Resistance Training"
            1 => "Oxidative Stress"
            2 => "Ovariectomy"
            3 => "Hepatic Steatosis"
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        "resumen" => "<span id="ceabs10" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle10">OBJECTIVE</span><p id="spara110" class="elsevierStyleSimplePara elsevierViewall">The objective of this study was to assess the effects of resistance training on oxidative stress markers in the livers of ovariectomized rats&#46;</p></span> <span id="ceabs20" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle20">METHOD</span><p id="spara120" class="elsevierStyleSimplePara elsevierViewall">Adult Sprague-Dawley rats were divided into the following four groups &#40;n &#61; 8 per group&#41;&#58; sham-operated sedentary&#44; ovariectomized sedentary&#44; sham-operated resistance training&#44; and ovariectomized resistance training&#46; During the resistance training period&#44; the animals climbed a 1&#46;1-m vertical ladder with weights attached to their tails&#59; the sessions were conducted 3 times per week&#44; with 4-9 climbs and 8-12 dynamic movements per climb&#46; The oxidative stress was assessed by measuring the levels of reduced glutathione and oxidized glutathione&#44; the enzymatic activity of catalase and superoxide dismutase&#44; lipid peroxidation&#44; vitamin E concentrations&#44; and the gene expression of glutathione peroxidase&#46;</p></span> <span id="ceabs30" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle30">RESULTS</span><p id="spara130" class="elsevierStyleSimplePara elsevierViewall">The results showed significant reductions in the reduced glutathione&#47;oxidized glutathione ratio &#40;4&#46;11&#177;0&#46;65 nmol&#47;g tec&#41;&#44; vitamin E concentration &#40;55&#46;36&#177;11&#46;11 nmol&#47;g&#41;&#44; and gene expression of glutathione peroxidase &#40;0&#46;49&#177;0&#46;16 arbitrary units&#41; in the livers of ovariectomized rats compared with the livers of unovariectomized animals &#40;5&#46;71&#177;0&#46;71 nmol&#47;g tec&#44; 100&#46;14&#177;10&#46;99 nmol&#47;g&#44; and 1&#46;09&#177;0&#46;54 arbitrary units&#44; respectively&#41;&#46; Moreover&#44; resistance training for 10 weeks was not able to reduce the oxidative stress in the livers of ovariectomized rats and induced negative changes in the hepatic anti-oxidative&#47;oxidative balance&#46;</p></span> <span id="ceabs40" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="cestitle40">CONCLUSION</span><p id="spara140" class="elsevierStyleSimplePara elsevierViewall">Our findings indicate that the resistance training program used in this study was not able to attenuate the hepatic oxidative damage caused by ovariectomy and increased the hepatic oxidative stress&#46;</p></span>"
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          "en" => "<p id="spara10" class="elsevierStyleSimplePara elsevierViewall">The sham-operated resistance training &#40;Sham-Rt&#41; and ovariectomized resistance training &#40;Ovx-Rt&#41; groups&#39; maximal workload at weeks 1&#44; 5&#44; and 10&#46; Values are presented as mean &#177; SDM<span class="elsevierStyleItalic">&#59; p &#8804; 0&#46;05</span>&#46; <span class="elsevierStyleItalic">a</span> statistically significant difference compared to week 1&#44; <span class="elsevierStyleItalic">b</span> statistically significant difference between weeks 5 and 10&#46;</p>"
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          "en" => "<p id="spara20" class="elsevierStyleSimplePara elsevierViewall"><span class="elsevierStyleBold">Figure 2a -</span> Levels of GSH &#40;a&#41; in liver homogenates 48h after the last session of the resistance training program in sham-operated sedentary &#40;Sham-Sed&#41;&#44; ovariectomized sedentary &#40;Ovx-Sed&#41;&#44; sham-operated resistance training&#44; &#40;Sham-Rt&#41; and ovariectomized resistance training &#40;Ovx-Rt&#41;&#46; Values are presented as mean &#177; SDM&#59; <span class="elsevierStyleItalic">p &#8804; 0&#46;05&#46; a</span> statistically significant difference compared to Sham-Sed&#44; <span class="elsevierStyleItalic">c</span> compared to Sham-Rt&#46;</p> <p id="spara30" class="elsevierStyleSimplePara elsevierViewall"><span class="elsevierStyleBold">Figure 2b -</span> Levels of GSSG &#40;b&#41; in liver homogenates 48h after the last session of the resistance training program in sham-operated sedentary &#40;Sham-Sed&#41;&#44; ovariectomized sedentary &#40;Ovx-Sed&#41;&#44; sham-operated resistance training&#44; &#40;Sham-Rt&#41; and ovariectomized resistance training &#40;Ovx-Rt&#41;&#46; Values are presented as mean &#177; SDM&#59; <span class="elsevierStyleItalic">p &#8804; 0&#46;05&#46; a</span> statistically significant difference compared to Sham-Sed&#44; <span class="elsevierStyleItalic">c</span> compared to Sham-Rt&#46;</p> <p id="spara40" class="elsevierStyleSimplePara elsevierViewall"><span class="elsevierStyleBold">Figure 2c -</span> Levels of GSH&#47;GSSG &#40;c&#41; in liver homogenates 48h after the last session of the resistance training program in sham-operated sedentary &#40;Sham-Sed&#41;&#44; ovariectomized sedentary &#40;Ovx-Sed&#41;&#44; sham-operated resistance training&#44; &#40;Sham-Rt&#41; and ovariectomized resistance training &#40;Ovx-Rt&#41;&#46; Values are presented as mean &#177; SDM&#59; <span class="elsevierStyleItalic">p &#8804; 0&#46;05&#46; a</span> statistically significant difference compared to Sham-Sed&#44; <span class="elsevierStyleItalic">c</span> compared to Sham-Rt&#46;</p> <p id="spara50" class="elsevierStyleSimplePara elsevierViewall"><span class="elsevierStyleBold">Figure 2d -</span> mRNA expression of glutathione peroxidase &#40;GSH-Px&#41; in the liver 48 h after the last session of the resistance training program in sham-operated sedentary &#40;Sham-Sed&#41;&#44; ovariectomized sedentary &#40;Ovx-Sed&#41;&#44; sham-operated resistance training&#44; &#40;Sham-Rt&#41; and ovariectomized resistance training &#40;Ovx-Rt&#41;&#46; Values are presented as mean &#177; SDM&#59; <span class="elsevierStyleItalic">p &#8804; 0&#46;05&#46; a</span> statistically significant difference compared to Sham-Sed&#44; <span class="elsevierStyleItalic">b</span> compared to Ovx-Sed&#46;</p>"
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          "en" => "<p id="spara80" class="elsevierStyleSimplePara elsevierViewall"><span class="elsevierStyleBold">Figure 5a -</span> Superoxide dismutase &#40;SOD&#41; &#40;a&#41; activity in the liver 48 h after the last session of the resistance training program in sham-operated sedentary &#40;Sham-Sed&#41;&#44; ovariectomized sedentary &#40;Ovx-Sed&#41;&#44; sham-operated resistance training&#44; &#40;Sham-Rt&#41; and ovariectomized resistance training &#40;Ovx-Rt&#41;&#46; Values are presented as mean &#177; SDM&#59; <span class="elsevierStyleItalic">p &#8804; 0&#46;05</span>&#46;</p> <p id="spara90" class="elsevierStyleSimplePara elsevierViewall"><span class="elsevierStyleBold">Figure 5b -</span> Catalase &#40;CAT&#41; &#40;b&#41; activity in the liver 48 h after the last session of the resistance training program in sham-operated sedentary &#40;Sham-Sed&#41;&#44; ovariectomized sedentary &#40;Ovx-Sed&#41;&#44; sham-operated resistance training&#44; &#40;Sham-Rt&#41; and ovariectomized resistance training &#40;Ovx-Rt&#41;&#46; Values are presented as mean &#177; SDM&#59; <span class="elsevierStyleItalic">p &#8804; 0&#46;05</span>&#46;</p>"
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          "leyenda" => "<p id="TFN01t01" class="elsevierStyleSimplePara elsevierViewall">Values are presented as the means&#177;SDM&#59; n <span class="elsevierStyleItalic">&#61;</span> 8 rats per group&#46; BW&#58; body weight&#44; <span class="elsevierStyleSup">&#8727;</span>significantly different from the Sham-Operated Sedentary &#40;Sham-Sed&#41; group &#40;<span class="elsevierStyleItalic">p&#8804;</span>0&#46;05<span class="elsevierStyleItalic">&#41;&#44;</span><span class="elsevierStyleSup">&#8224;</span> significantly different from the Ovariectomized Sedentary &#40;Ovx-Sed&#41; group &#40;<span class="elsevierStyleItalic">p</span>&#8804;0&#46;05<span class="elsevierStyleItalic">&#41;&#44;</span><span class="elsevierStyleSup">&#8225;</span> significantly different from the Sham-Operated Resistance Training &#40;Sham-Rt&#41; group &#40;<span class="elsevierStyleItalic">p</span>&#8804;0&#46;05<span class="elsevierStyleItalic">&#41;&#46;</span></p>"
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                  <table border="0" frame="\n
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      "titulo" => "REFERENCES"
      "seccion" => array:1 [
        0 => array:2 [
          "identificador" => "cebibsec10"
          "bibliografiaReferencia" => array:58 [
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        "texto" => "<p id="para450" class="elsevierStylePara elsevierViewall">The authors thank Mr&#46; Jos&#233; Carlos Lopes and Paula Pay&#227;o for laboratory-related technical assistance&#46; Financial support was provided by the Coordination for the Improvement of Higher Level &#40;CAPES&#41; and the Laboratory of Exercise Physiology Federal University of S&#227;o Carlos &#40;S&#227;o Carlos&#44; Brazil&#41;&#46;</p>"
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ISSN: 18075932
Original language: English
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