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    "textoCompleto" => "<p class="elsevierStylePara"> </p><p class="elsevierStylePara"><span class="elsevierStyleBold">Introduction </span></p><p class="elsevierStylePara"> It is generally accepted that obesity is predominantly associated with an impaired insulin-stimulated glucose uptake rate in skeletal muscle&#44; which has been attributed to insulin resistance&#46; Many studies have focused on the glucose transporter system as part of the underlying mechanisms&#46; Glucose transport into the skeletal muscle cell mediated by the glucose transporter proteins GLUT1 and GLUT4&#46;<span class="elsevierStyleSup">1 </span>The GLUT1 glucose transporter isoform is thought to support basal glucose transport&#44;<span class="elsevierStyleSup">2&#44;3 </span>while the GLUT4 isoform increases glucose transport in response to insulin and contraction&#46; Insulin and contractions translocate the GLUT4 from the intracellular pool to the plasma membrane and to the T-tubules&#46;<span class="elsevierStyleSup">4&#44;5</span></p><p class="elsevierStylePara"> In rodents&#44; it is well known that the glucose uptake capacity is greater in red oxidative muscles than in white glycolytic muscles&#46;<span class="elsevierStyleSup">6---10 </span>One underlying mechanism seems to be a greater level of GLUT4 expression&#44; both intracellularly<span class="elsevierStyleSup">6&#44;7</span> and at the plasma membrane&#46;<span class="elsevierStyleSup">8 </span>In human skeletal muscle&#44; glucose uptake was positively associated with the proportion of type I fibers and negatively associated with the proportion of type IIb fibers&#46;<span class="elsevierStyleSup">11 </span>These results were supported in the in vitro study by Zierath et al&#46;<span class="elsevierStyleSup">12 </span>that reported the insulin-stimulated increase in glucose uptake over basal is strongly correlated&#44; both positively with the percentage of type I muscle fibers and negatively with the percentage of type IIa fibers&#46; However&#44; unconvincing results regarding the relationship between fiber type distribution and GLUT4 content in human muscle have been reported&#46;<span class="elsevierStyleSup">13---15 </span>Andersen et al&#46;<span class="elsevierStyleSup">13</span> found no correlation between fiber type and GLUT4 content&#44; whereas Houmard and colleagues<span class="elsevierStyleSup">15 </span>showed a weak correlation between fiber type composition and GLUT4 content&#46;</p><p class="elsevierStylePara"> It has been shown that different muscles exhibit large differences in their GLUT4 content&#44; and this variation is often associated with differences in insulin-stimulated glucose uptake&#46;<span class="elsevierStyleSup">16&#44;17 </span>As different muscles are composed of a mixture of several different muscle fiber types&#44;<span class="elsevierStyleSup">18 </span>it is possible that a significant difference exists in GLUT4 content between muscles&#46;</p><p class="elsevierStylePara"> Possibly&#44; the differences in GLUT4 content and insulin-stimulated glucose uptake are more related to training status&#46; Changes in the skeletal muscle activity level is a key regulator of GLUT4 content in rats&#46;<span class="elsevierStyleSup">19&#44;20 </span>In humans&#44; athletes have more GLUT4 than untrained age-matched control subjects&#44;<span class="elsevierStyleSup">21&#44;22 </span>and in both normal healthy control subjects and individuals with diminished insulin-stimulated glucose uptake&#44; exercise training has been shown to increase GLUT4 content&#46;<span class="elsevierStyleSup">20&#44;23&#44;24 </span>Additionally&#44; a decrease in activity level will decrease GLUT4 content&#46;<span class="elsevierStyleSup">20&#44;25 </span>Finally&#44; changes in physical activity and GLUT4 content have been shown to be connected with changes in insulin-stimulated glucose uptake&#46;<span class="elsevierStyleSup">20</span></p><p class="elsevierStylePara"> The main objective of our work was to investigate the GLUT4 mRNA expression in soleus &#40;a predominantly slow-twitch muscle&#41; and gastrocnemius &#40;a predominantly fast-twitch muscle&#41; in obese mice in response to endurance training&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Subjects and methods </span></p><p class="elsevierStylePara"><span class="elsevierStyleBold">Animals </span></p><p class="elsevierStylePara"> Forty male C57BL&#47;6 mice &#40;4 weeks age&#41; were used in this study&#46; Eight non-obese mice &#40;Normal Base &#91;NB&#93; group&#59; Standard diet fed ad libitum&#41; served as non-obese non-trained controls &#40;NB group was considered as a baseline value in calculating gene expression in Real-Time that is expressed as relative values&#41; and 32 mice were put on a high fat diet &#40;HFD&#41; regimen for 12 wk consisting of ad libitum access to a 60&#37; kcal fat diet &#40;High-Fat Diet&#44; Razi Vaccine &#38; Serum Research Institute&#44; Iran&#41;&#46; At week 16&#44; the obese mice were randomized into the following treatment groups &#40;<span class="elsevierStyleItalic">n </span>&#61; 8 each group&#41;&#58; &#40;1&#41; Obese Base &#91;OB&#93; group&#59; &#40;2&#41; Low Intensity &#91;LI&#93; group&#59; &#40;3&#41; High Intensity &#91;HI&#93; group&#59; or &#40;4&#41; Obese Control &#91;OC&#93; group&#46; OB mice were killed before the training program&#46; LI and HI trained for 5 days&#47;wk for 12 wk on a motorized treadmill&#46; OC served as non-trained controls&#46; All mice were housed binary in cages and the temperature of the animal room was maintained at 22 <span class="elsevierStyleSup">¿</span>C&#44; and an artificial 12&#58;12-h light---dark cycle was set&#46; A familiarization period of two weeks in which mice ran 7---10 m&#47;min for 10---15 min on a 5&#37; slope was applied&#46; Thereafter&#44; training was continued in next 12 wk for 60 min continuously at 15 m&#47;min on a 5&#37; slope &#40;for LI&#41; and&#47;or for 41 min continuously at 22 m&#47;min on a 5&#37; slope &#40;for HI&#41;&#46; Total daily work was matched for both groups and set at 900 m distance running&#46; OC mice were kept sedentarily in the cage during the training program&#46; Weight was recorded weekly&#46; LI&#44; HI and OC were killed at end of training program&#46; Food was withdrawn 12---14 h before killing&#46; Experiments were approved by the Research Ethics Committee&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">uscle preparation </span></p><p class="elsevierStylePara"> Mice were anesthetized intraperitoneally with a mixture of Ketamine &#40;30---50 mg&#47;kg bw&#41; and Xylazine &#40;3---5 mg&#47;kg bw&#41; and were killed via direct heart blood withdrawal for muscle sampling 48 h after last exercise session&#44; and the soleus and gastrocnemius muscles were dissected&#46; Muscle samples were quickly frozen into liquid nitrogen and stored at &#8722;80 <span class="elsevierStyleSup">¿</span>C&#46; Tissue preparation and total RNA extraction procedures are described elsewhere in details&#46;<span class="elsevierStyleSup">26 </span></p><p class="elsevierStylePara"><span class="elsevierStyleBold">Real-time PCR </span></p><p class="elsevierStylePara"> RNA was reverse-transcribed with reverse transcriptase and random hexamer primers according to the manufacturer&#8217;s instructions &#40;AccuPower Green Star qPCR PreMix&#44; BiONEER&#44; Daejeon&#44; Korea&#41;&#46; Then&#44; PCR-mastermix containing the specific primers&#44; Hot Star Taq DNA polymerase and SYBR-Green PCR buffer were added&#46; All the samples were determined as duplicates&#44; and for a negative control the same set-up was used except for the addition of reverse transcriptase&#46; No PCR product was detected under these latter conditions&#46;</p><p class="elsevierStylePara"> GLUT4 mRNA and &#946;-actin&#44; a house-keeping gene&#44; levels were determined by quantitative reverse transcription---PCR of total cell RNA by using the forward primer 5 CCG CGG CCT CCT ATG AGA TAC T3 and the reverse primer 5 AGG CAC CCC GAA GAT GAG T3 for amplification of GLUT4 mRNA and the forward primer 5 ACA ATG AGC TGC GTG TGG CC 3 and the reverse primer 5 CCT CGT AGA TGG GCA CAG TG 3 for amplification of &#946;-actin mRNA&#46; Amplification products were electrophoresed on 2&#37; agarose gels and stained with ethidium bromide&#46;</p><p class="elsevierStylePara"> Real-time quantitation of GLUT4 to &#946;-actin mRNA was performed using a SYBR-Green PCR assay &#40;Rotor-gene 6000&#44; Corbett&#41;&#46; GLUT4 mRNA and &#946;-actin mRNA were amplified in separate tubes and the thermal cycling protocol for 40 cycles was denaturation at 95 <span class="elsevierStyleSup">¿</span>C for 20 s&#44; annealing at 60 <span class="elsevierStyleSup">¿</span>C for 60 s&#44; extension at 72 <span class="elsevierStyleSup">¿</span>C for 30 s that started with initial denaturation at 95 <span class="elsevierStyleSup">¿</span>C for 15 min and completed with final extension at 72 <span class="elsevierStyleSup">¿</span>C for 10 min&#46; During the extension step&#44; increase in fluorescence was measured in real time&#46; Data were obtained as CT values &#40;threshold cycle&#41;&#46; Relative gene expression was calculated using the Pfaffl formula<span class="elsevierStyleSup">27</span>&#58;</p><p class="elsevierStylePara"><img src="276v51n191-90458595fig1.jpg"></img></p><p class="elsevierStylePara"><span class="elsevierStyleBold">Data analysis </span></p><p class="elsevierStylePara"> Values are presented as mean &#177; SD&#46; One-way analysis of variance &#40;ANOVA&#41; followed by Tukey&#8217;s Honestly Significant Differences test were used to assess the effects of training on relative mRNA expression of GLUT4 in different groups&#46; Paired <span class="elsevierStyleItalic">t</span>-test&#44; also&#44; was used to assess the differences between soleus and gastrocnemius muscle GLUT4 mRNA in each group&#46; For all tests SPSS 21&#46;0 software was used &#40;SPSS Inc&#46;&#44; Chicago&#44; IL&#44; USA&#41;&#44; and <span class="elsevierStyleItalic">p </span>&#8804; 0&#46;05 was considered statistically significant&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Results </span></p><p class="elsevierStylePara"> Mean weight values of mice in different groups during HFD feeding and training phases are presented in Fig&#46; 1&#46; Weight in those groups who consumed HFD &#40;i&#46;e&#46; OB&#44; OC&#44; LI and HI&#41; have significantly increased compared to NB from the 5th week during HFD feeding phase which stayed higher until the 12th week &#40;<span class="elsevierStyleItalic">p </span>&#8804; 0&#46;05&#41;&#46; The weight gain in all 4 treatment groups was 18&#37; in average compared to NB that reached 32&#37; at the 12th week &#40;Fig&#46; 1&#41;&#46;</p><p class="elsevierStylePara"><img alt="Figure 1 Weight changes in mice during HFD feeding and training phases&#46; &#42;&#44; significant difference between obese and NB groups&#59;n†&#44; significant difference between training groups and OB&#59; ‡&#44; significant difference between HI and LI groups&#46; NB &#61; Normal Base&#44; OB &#61; Obese Base&#44; OC &#61; Obese Control&#44; LI &#61; Low Intensity training&#44; HI &#61; High Intensity training&#46;" src="276v51n191-90458595fig2.jpg"></img></p><p class="elsevierStylePara"><span class="elsevierStyleBold">Figure 1 </span>Weight changes in mice during HFD feeding and training phases&#46; &#42;&#44; significant difference between obese and NB groups&#59;n&#8224;&#44; significant difference between training groups and OB&#59; <span class="elsevierStyleItalic">&#8225;</span>&#44; significant difference between HI and LI groups&#46; NB &#61; Normal Base&#44; OB &#61; Obese Base&#44; OC &#61; Obese Control&#44; LI &#61; Low Intensity training&#44; HI &#61; High Intensity training&#46;</p><p class="elsevierStylePara"> During the training phase&#44; only the training groups &#40;LI &#38; HI&#41; and OC were studied and weight loss in training groups started to differentiate from OC group from the 6th week which became significant from the 8th week &#40;<span class="elsevierStyleItalic">p </span>&#8804; 0&#46;05&#41;&#46; So that&#44; weight loss in training groups &#40;compared to OC group&#41; was 5&#37; and 7&#37; in the 6th and 12th weeks of training phase&#44; respectively&#46; Interestingly&#44; the difference of weight loss between LI and HI appeared from the 9th week and became statistically different in the 11th and 12th weeks &#40;<span class="elsevierStyleItalic">p </span>&#8804; 0&#46;05&#41;&#46; In fact&#44; the rate of weight loss was higher in HI than LI&#44; so that weight changes percentage at the end of 12 weeks training in HI group was 8&#37; vs&#46; 5&#46;5&#37; in LI group&#46;</p><p class="elsevierStylePara"> GLUT4 mRNA expression of soleus muscle in the mice who engaged in LI group increased about 2&#46;2 fold&#44; against &#8764;1&#46;6 fold for gastrocnemius&#44; relative to NB &#40;<span class="elsevierStyleItalic">p </span>&#8804; 0&#46;05&#41;&#59; similarly&#44; relative GLUT4 mRNA expression increased&#44; albeit statistically nonsignificant&#44; by high intensity exercise training in soleus and gastrocnemius muscles by &#8764;2&#46;1 and &#8764;1&#46;8 fold&#44; respectively &#40;<span class="elsevierStyleItalic">p </span>&#62; 0&#46;05&#41;&#46; In addition&#44; GLUT4 mRNA expression of soleus and gastrocnemius muscles in LI and HI groups were significantly higher than OB and OC groups &#40;<span class="elsevierStyleItalic">p </span>&#8804; 0&#46;05&#41;&#44; which have slightly down-regulated in latter groups &#40;Fig&#46; 2&#41;&#46;</p><p class="elsevierStylePara"><img alt="Figure 2 GLUT4 mRNA content of experimental groups’ gastrocnemius and soleus muscle relative to matched muscle in NB group&#46; &#42;&#44; significant increase relative to NB &#40;p &#8804; 0&#46;05&#41;&#59; †&#44; significant difference in compare to OB and OC &#40;p &#8804; 0&#46;05&#41;&#46;" src="276v51n191-90458595fig3.jpg"></img></p><p class="elsevierStylePara"><span class="elsevierStyleBold">Figure 2 </span>GLUT4 mRNA content of experimental groups&#8217; gastrocnemius and soleus muscle relative to matched muscle in NB group&#46; &#42;&#44; significant increase relative to NB &#40;<span class="elsevierStyleItalic">p </span>&#8804; 0&#46;05&#41;&#59; &#8224;&#44; significant difference in compare to OB and OC &#40;<span class="elsevierStyleItalic">p </span>&#8804; 0&#46;05&#41;&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Discussion </span></p><p class="elsevierStylePara"> Ploug et al&#46;<span class="elsevierStyleSup">28 </span>found that GLUT4 in nonstimulated &#40;by insulin&#41; fibers is distributed along all the muscle fibers&#44; and is present both at the surface &#40;68&#37; of total GLUT4&#41; and in the core &#40;32&#37; of total GLUT4&#41; of the fibers&#46; They also found that the nuclei are displaced by and aligned with the blood vessels that course along the fiber surface&#44; thereby placing a large fraction of GLUT4 close to the source of glucose and obviating the need for diffusion over long distances&#46;<span class="elsevierStyleSup">28</span></p><p class="elsevierStylePara"> It is well documented that in the normal state&#44; GLUT4 cycles slowly between the plasma membrane and one or more intracellular compartments&#44; with the vast majority of the transporter residing in vesicular compartments within the cell interior&#46;<span class="elsevierStyleSup">28&#44;29 </span>Insulin stimulated accumulation of GLUT4 protein at the cell surface can be caused by 10---20 fold increase in the rate of exocytosis with a smaller decrease &#40;2---3 fold&#41; in the rate of GLUT4 endocytosis&#46;<span class="elsevierStyleSup">5&#44;10&#44;30</span></p><p class="elsevierStylePara"> Insulin stimulates glucose transport through GLUT4 translocation from the intracellular storage pool to the plasma membrane&#46; Insulin resistance caused by insulin deficiency or abnormal insulin signaling results in a decrease in GLUT4 expression and translocation&#44; and then causes hyperglycemia and diabetes&#46;<span class="elsevierStyleSup">30---32 </span>Muscle contraction has also been shown to increase GLUT4 content in the cell membrane&#46;<span class="elsevierStyleSup">29&#44;30</span></p><p class="elsevierStylePara"> Many studies suggest that exercise training cause GLUT4 protein expression and translocation to plasma membrane by a distinct mechanism from insulin signaling&#46;<span class="elsevierStyleSup">5&#44;33 </span>Most studies on rat skeletal muscle have indicated that more GLUT4 is present in the type I fibers compared with the type II fibers&#44;<span class="elsevierStyleSup">10&#44;34 </span>as we found in mice&#44; too&#46; In this study&#44; we used only the soleus muscle&#44; generally known as a type I muscle&#46;</p><p class="elsevierStylePara"> Our study is in agreement with other studies<span class="elsevierStyleSup">35---39 </span>that indicate exercise trainings increase GLUT4 protein expression in diabetic subjects like as what occurred in our obese subjects&#46;</p><p class="elsevierStylePara"> We found that GLUT-4 mRNA expression is increased by 12 weeks endurance training&#44; both in soleus &#40;a predominantly slow-twitch muscle&#41; and gastrocnemius &#40;a predominantly fast-twitch muscle&#41; in obese mice&#44; with an intensity-depended pattern&#46; This means that exercise intensity can affect the GLUT-4 gene expression &#40;and&#44; probably&#44; protein&#41; response to training which may result in significant metabolic changes especially in obese cases&#46; In agreement with our findings&#44; it has been shown that&#44; GLUT-4 protein and glucose transport are markedly higher in red-oxidative &#40;type I and IIa&#41; muscle fibers than in white-glycolytic fibers &#40;type IIb&#41;&#46;<span class="elsevierStyleSup">10&#44;40 </span>This might be an important factor in the adaptation to exercise&#44; because endurance training results in a shift from type IIb to type IIa fibers&#46;<span class="elsevierStyleSup">41 </span>However&#44; in human skeletal muscles&#44; there is a much smaller difference in the GLUT-4 expression in different muscle fiber types&#46;<span class="elsevierStyleSup">34&#44;42</span></p><p class="elsevierStylePara"> Daugaard et al&#46;<span class="elsevierStyleSup">34 </span>isolated individual muscle fibers and typed them according to myosin isoform&#46; GLUT-4 was &#8764;20&#37; higher in fibers expressing myosin heavy chain I than in those expressing myosin heavy chain IIA or IIB&#46; No difference could be detected between IIa and IIb fibers&#46; After 2 weeks of exercise training&#44; GLUT-4 was increased by &#8764;23&#37; in type I muscles&#44; but there were no changes in type IIa or IIb&#46; However&#44; the low-intensity exercise that was used is primarily known to recruit type I fibers&#46;</p><p class="elsevierStylePara"> According to our findings&#44; low intensity exercise training resulted in increased expression of GLUT4 mRNA in the oxidative muscle &#40;i&#46;e&#46; soleus&#41;&#44; probably due to cell requirements related to cellular oxidative capacity&#46; On the other hand&#44; high intensity exercise training did not result in increased expression of GLUT4 mRNA in the soleus&#44; nor in the gastrocnemius muscles&#46; As gastrocnemius muscle consists the mixture of red and white fibers&#44; another explanation would be that exercise training with low intensity was not enough to exercise or even activate red fibers of the gastrocnemius muscle&#44; and a nonsignificant increase in this muscle may be come from its white fibers&#46; This scenario can be extended to what observed findings related to high intensity&#59; i&#46;e&#46; a nonsignificant increase in this muscle by high intensity exercise may be come from its red fibers only&#46;</p><p class="elsevierStylePara"> Hormones also regulate muscle glucose transporter protein concentration&#46; GLUT-4 expression was increased by insulin and thyroid hormones<span class="elsevierStyleSup">20&#44;43 </span>and decreased by elevated cAMP&#46;<span class="elsevierStyleSup">44 </span>Regulation of GLUT-4 expression by contractile activity is independent of hormonal regulation&#44; because treadmill running increased GLUT-4 in diabetic rats&#46;<span class="elsevierStyleSup">45 </span>The effects of insulin-deficiency and denervation on GLUT-4 concentrations were additive&#46;<span class="elsevierStyleSup">45 </span>Physical training also increases muscle GLUT-4 protein and mRNA in patients with type 2 diabetes&#46;<span class="elsevierStyleSup">46 </span>These results suggest that muscle contractile activity directly modulates muscle GLUT-4 expression&#44; independent of insulin action&#46; Histone deacetylase &#40;HDAC5&#41; is a critical mediator of changes to GLUT4 mRNA levels in response to exercise&#46;<span class="elsevierStyleSup">51 </span>A series of histone modifications mediated by histone deacetylases and histone methyltransferases have been shown to climax in a metabolic knockdown of the GLUT4 gene in the skeletal muscle of rats&#46;<span class="elsevierStyleSup">52</span></p><p class="elsevierStylePara"> There is considerable evidence that GLUT4 mRNA levels in adipose and muscle tissue decrease with obesity&#44;<span class="elsevierStyleSup">47&#44;48 </span>and that increases in GLUT4 mRNA can alleviate or compensate insulin resistance&#46;<span class="elsevierStyleSup">49&#44;50 </span>Indeed&#44; our finding that GLUT4 mRNA levels were significantly increased by exercise training&#44; and this increase was associated with weight loss&#44; supports this idea&#46; Therefore&#44; increased transcription of GLUT4 in muscle during weight loss may be a certain event in reversing insulin resistance&#46;</p><p class="elsevierStylePara"> Collectively&#44; previous studies and our findings suggest that transcriptional regulation of the GLUT4 gene is responsive to body energy balance changes&#46; Additionally&#44; we found an intensity- and fiber type-dependent response of GLUT4 to exercise training that is not considered in previous studies&#46;</p><p class="elsevierStylePara"> It can be concluded that any disturbance in body energy balance especially by exercise training and&#47;or high fat diet can influence such molecular and cellular mechanisms which act to establish a stable homeostasis&#46; GLUT4 would decrease if energy intake is increased and&#44; reversely&#44; would increase if energy expenditure is increased&#46; These changes are almost similar in slow- and fast-twitch muscles with slightly more responsiveness in slow-twitch muscles that are more engaged in glucose metabolism&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Conflicts of interest </span></p><p class="elsevierStylePara"> The authors have no conflicts of interest to declare&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Acknowledgment </span></p><p class="elsevierStylePara"> Authors would like to thank University of Guilan which financially supported this research project&#46;</p><hr></hr><p class="elsevierStylePara"> Received 10 October 2015&#59; <br></br> accepted 14 December 2015<br></br> Available online 10 March 2016</p><p class="elsevierStylePara"><a href="http&#58;&#47;&#47;dx&#46;doi&#46;org&#47;10&#46;1016&#47;j&#46;apunts&#46;2015&#46;12&#46;001" class="elsevierStyleCrossRefs">http&#58;&#47;&#47;dx&#46;doi&#46;org&#47;10&#46;1016&#47;j&#46;apunts&#46;2015&#46;12&#46;001</a></p><p class="elsevierStylePara"> &#8727; Corresponding author&#46;<br></br><span class="elsevierStyleItalic">E-mail addresses&#58; </span><a href="mailto&#58;Mohebbih&#64;yahoo&#46;com" class="elsevierStyleCrossRefs">Mohebbih&#64;yahoo&#46;com</a>&#44; <a href="mailto&#58;mohebbi&#64;guilan&#46;ac&#46;ir" class="elsevierStyleCrossRefs">mohebbi&#64;guilan&#46;ac&#46;ir</a> &#40;H&#46; Mohebbi&#41;&#46;</p>"
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        "resumen" => "<p class="elsevierStylePara"> The aim of this study was to investigate the expression of GLUT4 mRNA in soleus and gastrocnemius muscles in obese mice in response to endurance training&#46; Forty male C57BL&#47;6 mice were used in this study&#46; Eight mice &#40;Normal Base &#91;NB&#93;&#41; served as non-obese non-trained controls&#44; and 32 mice were put on a high fat diet &#40;HFD&#41; regimen &#40;60&#37; kcal fat&#41; for 12 weeks&#46; At week 16&#44; the obese mice were randomized into the following treatment groups &#40;<span class="elsevierStyleItalic">n </span>&#61; 8 each group&#41;&#58; Obese Base &#91;OB&#93;&#59; Low Intensity &#91;LI&#93;&#59; High Intensity &#91;HI&#93;&#59; or Obese Control &#91;OC&#93; groups&#46; LI and HI trained for 5 days&#47;week for 12 weeks on a motorized treadmill at 15 m&#47;min on a 5&#37; slope &#40;for LI&#41;&#44; and&#47;or at 22 m&#47;min on a 5&#37; slope &#40;for HI&#41;&#46; OC mice were kept sedentarily in the cage during the training program&#46; GLUT4 mRNA expression was measured in gastrocnemius and soleus muscles using a Real Time-PCR method&#46; GLUT4 mRNA expression of soleus muscle in LI group increased about 2&#46;2 fold&#44; against about 1&#46;6 fold for gastrocnemius &#40;<span class="elsevierStyleItalic">p </span>&#60; &#46;05&#41;&#46; In addition&#44; GLUT4 mRNA expression of soleus and gastrocnemius muscles in LI and HI groups were significantly higher than OB and OC groups &#40;<span class="elsevierStyleItalic">p </span>&#60; &#46;05&#41;&#46; It can be concluded that any disturbance in body energy balance&#44; especially by exercise training and&#47;or high fat diet can influence these molecular and cellular mechanisms that act to establish a stable homeostasis&#46;</p>"
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        "resumen" => "<p class="elsevierStylePara"> El objetivo de este estudio fue investigar la expresi&#243;n de mARN de GLUT4 en los m&#250;sculos s&#243;leo y gastrocnemio en ratones obesos en respuesta al entrenamiento de resistencia&#46; Cuarenta machos C57BL&#47;6 ratones fueron utilizados en este estudio&#46; Ocho ratones &#40;normal base &#91;NB&#93;&#41; sirvieron como no obesos controles no entrenados&#44; y 32 ratones fueron puestos en una dieta alta en grasa &#40;HFD&#41;&#44; r&#233;gimen que siguieron &#40;60&#37; de grasa&#44; kcal&#41; durante 12 semanas&#46; En la semana 16 los ratones obesos fueron distribuidos aleatoriamente en los siguientes grupos de tratamiento &#40;n &#61; 8 cada grupo&#41;&#58; base obesos &#40;OB&#41;&#59; baja intensidad &#40;LI&#41;&#59; alta intensidad &#40;HI&#41;&#59; o grupos de control &#40;OC&#41; obesos&#46; LI y HI fueron entrenados durante 5 d&#237;as&#47;semana durante 12 semanas en una cinta rodante motorizada a 15 m&#47;min en una pendiente del 5&#37; &#40;para LI&#41; y&#47;o en 22 m&#47;min en una pendiente 5&#37; &#40;para HI&#41;&#46; Los ratones OCse mantuvieron sedentariamente en la jaula durante el programa de formaci&#243;n&#46; GLUT4 expresi&#243;n de mRNA se midi&#243; en los m&#250;sculos gastrocnemio y s&#243;leo&#44; utilizando el m&#233;todo en <span class="elsevierStyleItalic">real time</span>-PCR&#46; La expresi&#243;n de GLUT4 del mARN del m&#250;sculo s&#243;leo en el grupo LI aument&#243; aproximadamente &#8764;2&#44;2 veces&#44; frente a &#8764;1&#44;6 veces para los gemelos &#40;p &#8804; 0&#44;05&#41;&#46; Adem&#225;s&#44; la expresi&#243;n de GLUT4 mARN en los m&#250;sculos s&#243;leo y gastrocnemio en los grupos LI y HI fue significativamente mayor que en OB y en los grupos OC &#40;p &#8804; 0&#44;05&#41;&#46; Se puede concluir que cualquier alteraci&#243;n en el equilibrio energ&#233;tico del cuerpo&#44; especialmente por la pr&#225;ctica de ejercicio y&#47;o dieta alta en grasas puede influir en esos mecanismos moleculares y celulares que act&#250;an para establecer una homeostasis estable&#46;</p>"
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The effect of 12 weeks endurance training at 2 different intensities on GLUT4 mRNA expression of soleus and gastrocnemius muscles in obese mice
El efecto de 12 semanas de entrenamiento de resistencia con 2 intensidades diferentes en la expresión de GLUT4 mARN en los músculos sóleo y gastrocnemio en ratones obesos
Hamid Mohebbia, Hadi Rohanib, Sadegh Hassan-Niac
a Department of Exercise Physiology, Faculty of Physical Education and Sport Sciences, University of Guilan, Rasht, Iran
b Department of Sport Medicine, Sport Sciences Research Institute of Iran, Tehran, Iran
c Faculty of Biological Science, Tarbiat Modares University, Tehran, Iran
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        "titulo" => "El efecto de 12 semanas de entrenamiento de resistencia con 2 intensidades diferentes en la expresi&#243;n de GLUT4 mARN en los m&#250;sculos s&#243;leo y gastrocnemio en ratones obesos"
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    "textoCompleto" => "<p class="elsevierStylePara"> </p><p class="elsevierStylePara"><span class="elsevierStyleBold">Introduction </span></p><p class="elsevierStylePara"> It is generally accepted that obesity is predominantly associated with an impaired insulin-stimulated glucose uptake rate in skeletal muscle&#44; which has been attributed to insulin resistance&#46; Many studies have focused on the glucose transporter system as part of the underlying mechanisms&#46; Glucose transport into the skeletal muscle cell mediated by the glucose transporter proteins GLUT1 and GLUT4&#46;<span class="elsevierStyleSup">1 </span>The GLUT1 glucose transporter isoform is thought to support basal glucose transport&#44;<span class="elsevierStyleSup">2&#44;3 </span>while the GLUT4 isoform increases glucose transport in response to insulin and contraction&#46; Insulin and contractions translocate the GLUT4 from the intracellular pool to the plasma membrane and to the T-tubules&#46;<span class="elsevierStyleSup">4&#44;5</span></p><p class="elsevierStylePara"> In rodents&#44; it is well known that the glucose uptake capacity is greater in red oxidative muscles than in white glycolytic muscles&#46;<span class="elsevierStyleSup">6---10 </span>One underlying mechanism seems to be a greater level of GLUT4 expression&#44; both intracellularly<span class="elsevierStyleSup">6&#44;7</span> and at the plasma membrane&#46;<span class="elsevierStyleSup">8 </span>In human skeletal muscle&#44; glucose uptake was positively associated with the proportion of type I fibers and negatively associated with the proportion of type IIb fibers&#46;<span class="elsevierStyleSup">11 </span>These results were supported in the in vitro study by Zierath et al&#46;<span class="elsevierStyleSup">12 </span>that reported the insulin-stimulated increase in glucose uptake over basal is strongly correlated&#44; both positively with the percentage of type I muscle fibers and negatively with the percentage of type IIa fibers&#46; However&#44; unconvincing results regarding the relationship between fiber type distribution and GLUT4 content in human muscle have been reported&#46;<span class="elsevierStyleSup">13---15 </span>Andersen et al&#46;<span class="elsevierStyleSup">13</span> found no correlation between fiber type and GLUT4 content&#44; whereas Houmard and colleagues<span class="elsevierStyleSup">15 </span>showed a weak correlation between fiber type composition and GLUT4 content&#46;</p><p class="elsevierStylePara"> It has been shown that different muscles exhibit large differences in their GLUT4 content&#44; and this variation is often associated with differences in insulin-stimulated glucose uptake&#46;<span class="elsevierStyleSup">16&#44;17 </span>As different muscles are composed of a mixture of several different muscle fiber types&#44;<span class="elsevierStyleSup">18 </span>it is possible that a significant difference exists in GLUT4 content between muscles&#46;</p><p class="elsevierStylePara"> Possibly&#44; the differences in GLUT4 content and insulin-stimulated glucose uptake are more related to training status&#46; Changes in the skeletal muscle activity level is a key regulator of GLUT4 content in rats&#46;<span class="elsevierStyleSup">19&#44;20 </span>In humans&#44; athletes have more GLUT4 than untrained age-matched control subjects&#44;<span class="elsevierStyleSup">21&#44;22 </span>and in both normal healthy control subjects and individuals with diminished insulin-stimulated glucose uptake&#44; exercise training has been shown to increase GLUT4 content&#46;<span class="elsevierStyleSup">20&#44;23&#44;24 </span>Additionally&#44; a decrease in activity level will decrease GLUT4 content&#46;<span class="elsevierStyleSup">20&#44;25 </span>Finally&#44; changes in physical activity and GLUT4 content have been shown to be connected with changes in insulin-stimulated glucose uptake&#46;<span class="elsevierStyleSup">20</span></p><p class="elsevierStylePara"> The main objective of our work was to investigate the GLUT4 mRNA expression in soleus &#40;a predominantly slow-twitch muscle&#41; and gastrocnemius &#40;a predominantly fast-twitch muscle&#41; in obese mice in response to endurance training&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Subjects and methods </span></p><p class="elsevierStylePara"><span class="elsevierStyleBold">Animals </span></p><p class="elsevierStylePara"> Forty male C57BL&#47;6 mice &#40;4 weeks age&#41; were used in this study&#46; Eight non-obese mice &#40;Normal Base &#91;NB&#93; group&#59; Standard diet fed ad libitum&#41; served as non-obese non-trained controls &#40;NB group was considered as a baseline value in calculating gene expression in Real-Time that is expressed as relative values&#41; and 32 mice were put on a high fat diet &#40;HFD&#41; regimen for 12 wk consisting of ad libitum access to a 60&#37; kcal fat diet &#40;High-Fat Diet&#44; Razi Vaccine &#38; Serum Research Institute&#44; Iran&#41;&#46; At week 16&#44; the obese mice were randomized into the following treatment groups &#40;<span class="elsevierStyleItalic">n </span>&#61; 8 each group&#41;&#58; &#40;1&#41; Obese Base &#91;OB&#93; group&#59; &#40;2&#41; Low Intensity &#91;LI&#93; group&#59; &#40;3&#41; High Intensity &#91;HI&#93; group&#59; or &#40;4&#41; Obese Control &#91;OC&#93; group&#46; OB mice were killed before the training program&#46; LI and HI trained for 5 days&#47;wk for 12 wk on a motorized treadmill&#46; OC served as non-trained controls&#46; All mice were housed binary in cages and the temperature of the animal room was maintained at 22 <span class="elsevierStyleSup">¿</span>C&#44; and an artificial 12&#58;12-h light---dark cycle was set&#46; A familiarization period of two weeks in which mice ran 7---10 m&#47;min for 10---15 min on a 5&#37; slope was applied&#46; Thereafter&#44; training was continued in next 12 wk for 60 min continuously at 15 m&#47;min on a 5&#37; slope &#40;for LI&#41; and&#47;or for 41 min continuously at 22 m&#47;min on a 5&#37; slope &#40;for HI&#41;&#46; Total daily work was matched for both groups and set at 900 m distance running&#46; OC mice were kept sedentarily in the cage during the training program&#46; Weight was recorded weekly&#46; LI&#44; HI and OC were killed at end of training program&#46; Food was withdrawn 12---14 h before killing&#46; Experiments were approved by the Research Ethics Committee&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">uscle preparation </span></p><p class="elsevierStylePara"> Mice were anesthetized intraperitoneally with a mixture of Ketamine &#40;30---50 mg&#47;kg bw&#41; and Xylazine &#40;3---5 mg&#47;kg bw&#41; and were killed via direct heart blood withdrawal for muscle sampling 48 h after last exercise session&#44; and the soleus and gastrocnemius muscles were dissected&#46; Muscle samples were quickly frozen into liquid nitrogen and stored at &#8722;80 <span class="elsevierStyleSup">¿</span>C&#46; Tissue preparation and total RNA extraction procedures are described elsewhere in details&#46;<span class="elsevierStyleSup">26 </span></p><p class="elsevierStylePara"><span class="elsevierStyleBold">Real-time PCR </span></p><p class="elsevierStylePara"> RNA was reverse-transcribed with reverse transcriptase and random hexamer primers according to the manufacturer&#8217;s instructions &#40;AccuPower Green Star qPCR PreMix&#44; BiONEER&#44; Daejeon&#44; Korea&#41;&#46; Then&#44; PCR-mastermix containing the specific primers&#44; Hot Star Taq DNA polymerase and SYBR-Green PCR buffer were added&#46; All the samples were determined as duplicates&#44; and for a negative control the same set-up was used except for the addition of reverse transcriptase&#46; No PCR product was detected under these latter conditions&#46;</p><p class="elsevierStylePara"> GLUT4 mRNA and &#946;-actin&#44; a house-keeping gene&#44; levels were determined by quantitative reverse transcription---PCR of total cell RNA by using the forward primer 5 CCG CGG CCT CCT ATG AGA TAC T3 and the reverse primer 5 AGG CAC CCC GAA GAT GAG T3 for amplification of GLUT4 mRNA and the forward primer 5 ACA ATG AGC TGC GTG TGG CC 3 and the reverse primer 5 CCT CGT AGA TGG GCA CAG TG 3 for amplification of &#946;-actin mRNA&#46; Amplification products were electrophoresed on 2&#37; agarose gels and stained with ethidium bromide&#46;</p><p class="elsevierStylePara"> Real-time quantitation of GLUT4 to &#946;-actin mRNA was performed using a SYBR-Green PCR assay &#40;Rotor-gene 6000&#44; Corbett&#41;&#46; GLUT4 mRNA and &#946;-actin mRNA were amplified in separate tubes and the thermal cycling protocol for 40 cycles was denaturation at 95 <span class="elsevierStyleSup">¿</span>C for 20 s&#44; annealing at 60 <span class="elsevierStyleSup">¿</span>C for 60 s&#44; extension at 72 <span class="elsevierStyleSup">¿</span>C for 30 s that started with initial denaturation at 95 <span class="elsevierStyleSup">¿</span>C for 15 min and completed with final extension at 72 <span class="elsevierStyleSup">¿</span>C for 10 min&#46; During the extension step&#44; increase in fluorescence was measured in real time&#46; Data were obtained as CT values &#40;threshold cycle&#41;&#46; Relative gene expression was calculated using the Pfaffl formula<span class="elsevierStyleSup">27</span>&#58;</p><p class="elsevierStylePara"><img src="276v51n191-90458595fig1.jpg"></img></p><p class="elsevierStylePara"><span class="elsevierStyleBold">Data analysis </span></p><p class="elsevierStylePara"> Values are presented as mean &#177; SD&#46; One-way analysis of variance &#40;ANOVA&#41; followed by Tukey&#8217;s Honestly Significant Differences test were used to assess the effects of training on relative mRNA expression of GLUT4 in different groups&#46; Paired <span class="elsevierStyleItalic">t</span>-test&#44; also&#44; was used to assess the differences between soleus and gastrocnemius muscle GLUT4 mRNA in each group&#46; For all tests SPSS 21&#46;0 software was used &#40;SPSS Inc&#46;&#44; Chicago&#44; IL&#44; USA&#41;&#44; and <span class="elsevierStyleItalic">p </span>&#8804; 0&#46;05 was considered statistically significant&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Results </span></p><p class="elsevierStylePara"> Mean weight values of mice in different groups during HFD feeding and training phases are presented in Fig&#46; 1&#46; Weight in those groups who consumed HFD &#40;i&#46;e&#46; OB&#44; OC&#44; LI and HI&#41; have significantly increased compared to NB from the 5th week during HFD feeding phase which stayed higher until the 12th week &#40;<span class="elsevierStyleItalic">p </span>&#8804; 0&#46;05&#41;&#46; The weight gain in all 4 treatment groups was 18&#37; in average compared to NB that reached 32&#37; at the 12th week &#40;Fig&#46; 1&#41;&#46;</p><p class="elsevierStylePara"><img alt="Figure 1 Weight changes in mice during HFD feeding and training phases&#46; &#42;&#44; significant difference between obese and NB groups&#59;n†&#44; significant difference between training groups and OB&#59; ‡&#44; significant difference between HI and LI groups&#46; NB &#61; Normal Base&#44; OB &#61; Obese Base&#44; OC &#61; Obese Control&#44; LI &#61; Low Intensity training&#44; HI &#61; High Intensity training&#46;" src="276v51n191-90458595fig2.jpg"></img></p><p class="elsevierStylePara"><span class="elsevierStyleBold">Figure 1 </span>Weight changes in mice during HFD feeding and training phases&#46; &#42;&#44; significant difference between obese and NB groups&#59;n&#8224;&#44; significant difference between training groups and OB&#59; <span class="elsevierStyleItalic">&#8225;</span>&#44; significant difference between HI and LI groups&#46; NB &#61; Normal Base&#44; OB &#61; Obese Base&#44; OC &#61; Obese Control&#44; LI &#61; Low Intensity training&#44; HI &#61; High Intensity training&#46;</p><p class="elsevierStylePara"> During the training phase&#44; only the training groups &#40;LI &#38; HI&#41; and OC were studied and weight loss in training groups started to differentiate from OC group from the 6th week which became significant from the 8th week &#40;<span class="elsevierStyleItalic">p </span>&#8804; 0&#46;05&#41;&#46; So that&#44; weight loss in training groups &#40;compared to OC group&#41; was 5&#37; and 7&#37; in the 6th and 12th weeks of training phase&#44; respectively&#46; Interestingly&#44; the difference of weight loss between LI and HI appeared from the 9th week and became statistically different in the 11th and 12th weeks &#40;<span class="elsevierStyleItalic">p </span>&#8804; 0&#46;05&#41;&#46; In fact&#44; the rate of weight loss was higher in HI than LI&#44; so that weight changes percentage at the end of 12 weeks training in HI group was 8&#37; vs&#46; 5&#46;5&#37; in LI group&#46;</p><p class="elsevierStylePara"> GLUT4 mRNA expression of soleus muscle in the mice who engaged in LI group increased about 2&#46;2 fold&#44; against &#8764;1&#46;6 fold for gastrocnemius&#44; relative to NB &#40;<span class="elsevierStyleItalic">p </span>&#8804; 0&#46;05&#41;&#59; similarly&#44; relative GLUT4 mRNA expression increased&#44; albeit statistically nonsignificant&#44; by high intensity exercise training in soleus and gastrocnemius muscles by &#8764;2&#46;1 and &#8764;1&#46;8 fold&#44; respectively &#40;<span class="elsevierStyleItalic">p </span>&#62; 0&#46;05&#41;&#46; In addition&#44; GLUT4 mRNA expression of soleus and gastrocnemius muscles in LI and HI groups were significantly higher than OB and OC groups &#40;<span class="elsevierStyleItalic">p </span>&#8804; 0&#46;05&#41;&#44; which have slightly down-regulated in latter groups &#40;Fig&#46; 2&#41;&#46;</p><p class="elsevierStylePara"><img alt="Figure 2 GLUT4 mRNA content of experimental groups’ gastrocnemius and soleus muscle relative to matched muscle in NB group&#46; &#42;&#44; significant increase relative to NB &#40;p &#8804; 0&#46;05&#41;&#59; †&#44; significant difference in compare to OB and OC &#40;p &#8804; 0&#46;05&#41;&#46;" src="276v51n191-90458595fig3.jpg"></img></p><p class="elsevierStylePara"><span class="elsevierStyleBold">Figure 2 </span>GLUT4 mRNA content of experimental groups&#8217; gastrocnemius and soleus muscle relative to matched muscle in NB group&#46; &#42;&#44; significant increase relative to NB &#40;<span class="elsevierStyleItalic">p </span>&#8804; 0&#46;05&#41;&#59; &#8224;&#44; significant difference in compare to OB and OC &#40;<span class="elsevierStyleItalic">p </span>&#8804; 0&#46;05&#41;&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Discussion </span></p><p class="elsevierStylePara"> Ploug et al&#46;<span class="elsevierStyleSup">28 </span>found that GLUT4 in nonstimulated &#40;by insulin&#41; fibers is distributed along all the muscle fibers&#44; and is present both at the surface &#40;68&#37; of total GLUT4&#41; and in the core &#40;32&#37; of total GLUT4&#41; of the fibers&#46; They also found that the nuclei are displaced by and aligned with the blood vessels that course along the fiber surface&#44; thereby placing a large fraction of GLUT4 close to the source of glucose and obviating the need for diffusion over long distances&#46;<span class="elsevierStyleSup">28</span></p><p class="elsevierStylePara"> It is well documented that in the normal state&#44; GLUT4 cycles slowly between the plasma membrane and one or more intracellular compartments&#44; with the vast majority of the transporter residing in vesicular compartments within the cell interior&#46;<span class="elsevierStyleSup">28&#44;29 </span>Insulin stimulated accumulation of GLUT4 protein at the cell surface can be caused by 10---20 fold increase in the rate of exocytosis with a smaller decrease &#40;2---3 fold&#41; in the rate of GLUT4 endocytosis&#46;<span class="elsevierStyleSup">5&#44;10&#44;30</span></p><p class="elsevierStylePara"> Insulin stimulates glucose transport through GLUT4 translocation from the intracellular storage pool to the plasma membrane&#46; Insulin resistance caused by insulin deficiency or abnormal insulin signaling results in a decrease in GLUT4 expression and translocation&#44; and then causes hyperglycemia and diabetes&#46;<span class="elsevierStyleSup">30---32 </span>Muscle contraction has also been shown to increase GLUT4 content in the cell membrane&#46;<span class="elsevierStyleSup">29&#44;30</span></p><p class="elsevierStylePara"> Many studies suggest that exercise training cause GLUT4 protein expression and translocation to plasma membrane by a distinct mechanism from insulin signaling&#46;<span class="elsevierStyleSup">5&#44;33 </span>Most studies on rat skeletal muscle have indicated that more GLUT4 is present in the type I fibers compared with the type II fibers&#44;<span class="elsevierStyleSup">10&#44;34 </span>as we found in mice&#44; too&#46; In this study&#44; we used only the soleus muscle&#44; generally known as a type I muscle&#46;</p><p class="elsevierStylePara"> Our study is in agreement with other studies<span class="elsevierStyleSup">35---39 </span>that indicate exercise trainings increase GLUT4 protein expression in diabetic subjects like as what occurred in our obese subjects&#46;</p><p class="elsevierStylePara"> We found that GLUT-4 mRNA expression is increased by 12 weeks endurance training&#44; both in soleus &#40;a predominantly slow-twitch muscle&#41; and gastrocnemius &#40;a predominantly fast-twitch muscle&#41; in obese mice&#44; with an intensity-depended pattern&#46; This means that exercise intensity can affect the GLUT-4 gene expression &#40;and&#44; probably&#44; protein&#41; response to training which may result in significant metabolic changes especially in obese cases&#46; In agreement with our findings&#44; it has been shown that&#44; GLUT-4 protein and glucose transport are markedly higher in red-oxidative &#40;type I and IIa&#41; muscle fibers than in white-glycolytic fibers &#40;type IIb&#41;&#46;<span class="elsevierStyleSup">10&#44;40 </span>This might be an important factor in the adaptation to exercise&#44; because endurance training results in a shift from type IIb to type IIa fibers&#46;<span class="elsevierStyleSup">41 </span>However&#44; in human skeletal muscles&#44; there is a much smaller difference in the GLUT-4 expression in different muscle fiber types&#46;<span class="elsevierStyleSup">34&#44;42</span></p><p class="elsevierStylePara"> Daugaard et al&#46;<span class="elsevierStyleSup">34 </span>isolated individual muscle fibers and typed them according to myosin isoform&#46; GLUT-4 was &#8764;20&#37; higher in fibers expressing myosin heavy chain I than in those expressing myosin heavy chain IIA or IIB&#46; No difference could be detected between IIa and IIb fibers&#46; After 2 weeks of exercise training&#44; GLUT-4 was increased by &#8764;23&#37; in type I muscles&#44; but there were no changes in type IIa or IIb&#46; However&#44; the low-intensity exercise that was used is primarily known to recruit type I fibers&#46;</p><p class="elsevierStylePara"> According to our findings&#44; low intensity exercise training resulted in increased expression of GLUT4 mRNA in the oxidative muscle &#40;i&#46;e&#46; soleus&#41;&#44; probably due to cell requirements related to cellular oxidative capacity&#46; On the other hand&#44; high intensity exercise training did not result in increased expression of GLUT4 mRNA in the soleus&#44; nor in the gastrocnemius muscles&#46; As gastrocnemius muscle consists the mixture of red and white fibers&#44; another explanation would be that exercise training with low intensity was not enough to exercise or even activate red fibers of the gastrocnemius muscle&#44; and a nonsignificant increase in this muscle may be come from its white fibers&#46; This scenario can be extended to what observed findings related to high intensity&#59; i&#46;e&#46; a nonsignificant increase in this muscle by high intensity exercise may be come from its red fibers only&#46;</p><p class="elsevierStylePara"> Hormones also regulate muscle glucose transporter protein concentration&#46; GLUT-4 expression was increased by insulin and thyroid hormones<span class="elsevierStyleSup">20&#44;43 </span>and decreased by elevated cAMP&#46;<span class="elsevierStyleSup">44 </span>Regulation of GLUT-4 expression by contractile activity is independent of hormonal regulation&#44; because treadmill running increased GLUT-4 in diabetic rats&#46;<span class="elsevierStyleSup">45 </span>The effects of insulin-deficiency and denervation on GLUT-4 concentrations were additive&#46;<span class="elsevierStyleSup">45 </span>Physical training also increases muscle GLUT-4 protein and mRNA in patients with type 2 diabetes&#46;<span class="elsevierStyleSup">46 </span>These results suggest that muscle contractile activity directly modulates muscle GLUT-4 expression&#44; independent of insulin action&#46; Histone deacetylase &#40;HDAC5&#41; is a critical mediator of changes to GLUT4 mRNA levels in response to exercise&#46;<span class="elsevierStyleSup">51 </span>A series of histone modifications mediated by histone deacetylases and histone methyltransferases have been shown to climax in a metabolic knockdown of the GLUT4 gene in the skeletal muscle of rats&#46;<span class="elsevierStyleSup">52</span></p><p class="elsevierStylePara"> There is considerable evidence that GLUT4 mRNA levels in adipose and muscle tissue decrease with obesity&#44;<span class="elsevierStyleSup">47&#44;48 </span>and that increases in GLUT4 mRNA can alleviate or compensate insulin resistance&#46;<span class="elsevierStyleSup">49&#44;50 </span>Indeed&#44; our finding that GLUT4 mRNA levels were significantly increased by exercise training&#44; and this increase was associated with weight loss&#44; supports this idea&#46; Therefore&#44; increased transcription of GLUT4 in muscle during weight loss may be a certain event in reversing insulin resistance&#46;</p><p class="elsevierStylePara"> Collectively&#44; previous studies and our findings suggest that transcriptional regulation of the GLUT4 gene is responsive to body energy balance changes&#46; Additionally&#44; we found an intensity- and fiber type-dependent response of GLUT4 to exercise training that is not considered in previous studies&#46;</p><p class="elsevierStylePara"> It can be concluded that any disturbance in body energy balance especially by exercise training and&#47;or high fat diet can influence such molecular and cellular mechanisms which act to establish a stable homeostasis&#46; GLUT4 would decrease if energy intake is increased and&#44; reversely&#44; would increase if energy expenditure is increased&#46; These changes are almost similar in slow- and fast-twitch muscles with slightly more responsiveness in slow-twitch muscles that are more engaged in glucose metabolism&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Conflicts of interest </span></p><p class="elsevierStylePara"> The authors have no conflicts of interest to declare&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Acknowledgment </span></p><p class="elsevierStylePara"> Authors would like to thank University of Guilan which financially supported this research project&#46;</p><hr></hr><p class="elsevierStylePara"> Received 10 October 2015&#59; <br></br> accepted 14 December 2015<br></br> Available online 10 March 2016</p><p class="elsevierStylePara"><a href="http&#58;&#47;&#47;dx&#46;doi&#46;org&#47;10&#46;1016&#47;j&#46;apunts&#46;2015&#46;12&#46;001" class="elsevierStyleCrossRefs">http&#58;&#47;&#47;dx&#46;doi&#46;org&#47;10&#46;1016&#47;j&#46;apunts&#46;2015&#46;12&#46;001</a></p><p class="elsevierStylePara"> &#8727; Corresponding author&#46;<br></br><span class="elsevierStyleItalic">E-mail addresses&#58; </span><a href="mailto&#58;Mohebbih&#64;yahoo&#46;com" class="elsevierStyleCrossRefs">Mohebbih&#64;yahoo&#46;com</a>&#44; <a href="mailto&#58;mohebbi&#64;guilan&#46;ac&#46;ir" class="elsevierStyleCrossRefs">mohebbi&#64;guilan&#46;ac&#46;ir</a> &#40;H&#46; Mohebbi&#41;&#46;</p>"
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        "resumen" => "<p class="elsevierStylePara"> The aim of this study was to investigate the expression of GLUT4 mRNA in soleus and gastrocnemius muscles in obese mice in response to endurance training&#46; Forty male C57BL&#47;6 mice were used in this study&#46; Eight mice &#40;Normal Base &#91;NB&#93;&#41; served as non-obese non-trained controls&#44; and 32 mice were put on a high fat diet &#40;HFD&#41; regimen &#40;60&#37; kcal fat&#41; for 12 weeks&#46; At week 16&#44; the obese mice were randomized into the following treatment groups &#40;<span class="elsevierStyleItalic">n </span>&#61; 8 each group&#41;&#58; Obese Base &#91;OB&#93;&#59; Low Intensity &#91;LI&#93;&#59; High Intensity &#91;HI&#93;&#59; or Obese Control &#91;OC&#93; groups&#46; LI and HI trained for 5 days&#47;week for 12 weeks on a motorized treadmill at 15 m&#47;min on a 5&#37; slope &#40;for LI&#41;&#44; and&#47;or at 22 m&#47;min on a 5&#37; slope &#40;for HI&#41;&#46; OC mice were kept sedentarily in the cage during the training program&#46; GLUT4 mRNA expression was measured in gastrocnemius and soleus muscles using a Real Time-PCR method&#46; GLUT4 mRNA expression of soleus muscle in LI group increased about 2&#46;2 fold&#44; against about 1&#46;6 fold for gastrocnemius &#40;<span class="elsevierStyleItalic">p </span>&#60; &#46;05&#41;&#46; In addition&#44; GLUT4 mRNA expression of soleus and gastrocnemius muscles in LI and HI groups were significantly higher than OB and OC groups &#40;<span class="elsevierStyleItalic">p </span>&#60; &#46;05&#41;&#46; It can be concluded that any disturbance in body energy balance&#44; especially by exercise training and&#47;or high fat diet can influence these molecular and cellular mechanisms that act to establish a stable homeostasis&#46;</p>"
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        "resumen" => "<p class="elsevierStylePara"> El objetivo de este estudio fue investigar la expresi&#243;n de mARN de GLUT4 en los m&#250;sculos s&#243;leo y gastrocnemio en ratones obesos en respuesta al entrenamiento de resistencia&#46; Cuarenta machos C57BL&#47;6 ratones fueron utilizados en este estudio&#46; Ocho ratones &#40;normal base &#91;NB&#93;&#41; sirvieron como no obesos controles no entrenados&#44; y 32 ratones fueron puestos en una dieta alta en grasa &#40;HFD&#41;&#44; r&#233;gimen que siguieron &#40;60&#37; de grasa&#44; kcal&#41; durante 12 semanas&#46; En la semana 16 los ratones obesos fueron distribuidos aleatoriamente en los siguientes grupos de tratamiento &#40;n &#61; 8 cada grupo&#41;&#58; base obesos &#40;OB&#41;&#59; baja intensidad &#40;LI&#41;&#59; alta intensidad &#40;HI&#41;&#59; o grupos de control &#40;OC&#41; obesos&#46; LI y HI fueron entrenados durante 5 d&#237;as&#47;semana durante 12 semanas en una cinta rodante motorizada a 15 m&#47;min en una pendiente del 5&#37; &#40;para LI&#41; y&#47;o en 22 m&#47;min en una pendiente 5&#37; &#40;para HI&#41;&#46; Los ratones OCse mantuvieron sedentariamente en la jaula durante el programa de formaci&#243;n&#46; GLUT4 expresi&#243;n de mRNA se midi&#243; en los m&#250;sculos gastrocnemio y s&#243;leo&#44; utilizando el m&#233;todo en <span class="elsevierStyleItalic">real time</span>-PCR&#46; La expresi&#243;n de GLUT4 del mARN del m&#250;sculo s&#243;leo en el grupo LI aument&#243; aproximadamente &#8764;2&#44;2 veces&#44; frente a &#8764;1&#44;6 veces para los gemelos &#40;p &#8804; 0&#44;05&#41;&#46; Adem&#225;s&#44; la expresi&#243;n de GLUT4 mARN en los m&#250;sculos s&#243;leo y gastrocnemio en los grupos LI y HI fue significativamente mayor que en OB y en los grupos OC &#40;p &#8804; 0&#44;05&#41;&#46; Se puede concluir que cualquier alteraci&#243;n en el equilibrio energ&#233;tico del cuerpo&#44; especialmente por la pr&#225;ctica de ejercicio y&#47;o dieta alta en grasas puede influir en esos mecanismos moleculares y celulares que act&#250;an para establecer una homeostasis estable&#46;</p>"
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