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La insulina i les contraccions transporten el GLUT4 de la reserva intracel&#183;lular a la membrana plasm&#224;tica i als t&#250;buls T<span class="elsevierStyleSup">4&#44;5</span>&#46;</p><p class="elsevierStylePara"> Se sap que en els rosegadors&#44; la capacitat d&#8217;absorci&#243; de la glucosa &#233;s major en els m&#250;sculs oxidatius vermells que en els m&#250;sculs glucol&#237;tics blancs<span class="elsevierStyleSup">6-10</span>&#46; Sembla que un mecanisme subjacent t&#233; un major nivell d&#8217;expressi&#243; de GLUT4&#44; tant a nivell intracel&#183;lular<span class="elsevierStyleSup">6&#44;7</span> com a la membrana plasm&#224;tica8&#46; En el m&#250;scul esquel&#232;tic hum&#224;&#44; la captaci&#243; de glucosa s&#8217;associa positivament a la proporci&#243; de fibres de tipus i i negativament a la proporci&#243; de fibres de tipus iib<span class="elsevierStyleSup">11</span>&#46; Aquests resultats es basen en l&#8217;estudi in vitro de Zierath et al&#46;<span class="elsevierStyleSup">12</span>&#44; que reportaren que l&#8217;augment d&#8217;insulina estimulada en la captaci&#243; de la glucosa per damunt del nivell basal correlaciona estretament&#44; tant afirmativament amb el percentatge de fibres musculars de tipus i&#44; com negativament amb el percentatge de fibres tipus iia&#46; Tanmateix&#44; s&#8217;han reportat resultats poc convincents respecte a la relaci&#243; entre la distribuci&#243; del tipus de fibra i el contingut de GLUT4 en el m&#250;scul hum&#224;<span class="elsevierStyleSup">13-15</span>&#46;</p><p class="elsevierStylePara"> Andersen et al&#46;<span class="elsevierStyleSup">13</span> no trobaren cap correlaci&#243; entre tipus de fibra i contingut de GLUT4&#44; mentre que Houmard et al&#46;<span class="elsevierStyleSup">15</span> mostraren una baixa correlaci&#243; entre tipus de composici&#243; de la fibra i contingut de GLUT4&#46;</p><p class="elsevierStylePara"> Ha estat demostrat que els diferents m&#250;sculs presenten importants difer&#232;ncies de contingut de GLUT4&#44; i aquestes difer&#232;ncies sovint estan associades a les difer&#232;ncies en la captaci&#243; de la glucosa estimulada per la insulina<span class="elsevierStyleSup">16&#44;17</span>&#46; Com que el diferents m&#250;sculs es composen d&#8217;una combinaci&#243; de diferents tipus de fibres musculars<span class="elsevierStyleSup">18</span>&#44; &#233;s possible que hi hagi difer&#232;ncies significatives en el contingut de GLUT4 entre m&#250;sculs&#46;</p><p class="elsevierStylePara"> Probablement&#44; les difer&#232;ncies en el contingut de GLUT4 i la captaci&#243; de la glucosa estimulada per la insulina estan relacionades amb el nivell d&#8217;entrenament&#46; Els canvis de nivell d&#8217;activitat del m&#250;scul esquel&#232;tic s&#243;n la clau reguladora del contingut de GLUT4 en ratolins<span class="elsevierStyleSup">19&#44;20</span>&#46; Entre els humans&#44; els esportistes tenen m&#233;s GLUT4 que els subjectes control de la mateixa edat no entrenats<span class="elsevierStyleSup">21&#44;22</span>&#44; i en ambd&#243;s&#44; subjectes control normals i sans i individus amb la captaci&#243; disminu&#239;da de la glucosa estimulada per la insulina&#44; l&#8217;entrenament f&#237;sic ha demostrat que incrementa el contingut de GLUT4<span class="elsevierStyleSup">20&#44;23&#44;24</span>&#46; A m&#233;s&#44; una disminuci&#243; del nivell d&#8217;activitat pot minorar el contingut de GLUT4<span class="elsevierStyleSup">20&#44;25</span>&#46; Finalment&#44; s&#8217;ha demostrat que canvis en l&#8217;activitat f&#237;sica i el contingut de GLUT4 estan relacionats amb canvis en la captaci&#243; de la glucosa estimulada per la insulina<span class="elsevierStyleSup">20</span>&#46;</p><p class="elsevierStylePara"> El principal objectiu d&#8217;aquest treball fou investigar l&#8217;expressi&#243; de l&#8217;ARNm del GLUT4 en el soli &#40;m&#250;scul predominantment de contracci&#243; lenta&#41; i el gastrocnemi &#40;m&#250;scul predominantment de contracci&#243; r&#224;pida&#41; com a resposta a l&#8217;entrenament de resist&#232;ncia en ratolins obesos&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Subjectes i m&#232;todes </span></p><p class="elsevierStylePara"><span class="elsevierStyleBold">Animals </span></p><p class="elsevierStylePara"> En aquest estudi s&#8217;utilitzaren 40 ratolins mascles C57BL&#47;6 &#40;de 4 setmanes&#41;&#46; Vuit ratolins no obesos &#40;base normal &#91;BN&#93;&#59; dieta est&#224;ndard alimentats <span class="elsevierStyleItalic">ad libitum</span>&#41; van servir de control no obesos no entrenats &#40;el grup BN es consider&#224; com a valor de refer&#232;ncia per calcular l&#8217;expressi&#243; g&#232;nica en temps real que s&#8217;expressa com a valors relatius&#41; i 32 ratolins van ser sotmesos a una dieta aliment&#224;ria rica en greixos &#40;DAG&#41; durant 12 setmanes que consist&#237; en acc&#233;s lliure a una dieta rica en greixos de 60&#37; kcal &#40;<span class="elsevierStyleItalic">High-Fat Diet&#44; Razi Vaccine &#38; Serum Research Institute</span>&#44; Iran&#41;&#46; A la setmana 16&#44; els ratolins obesos foren assignats aleat&#242;riament als grups de tractament seg&#252;ents &#40;n &#61; 8 cada grup&#41;&#58; 1&#41; grup ob&#232;s base &#91;OB&#93;&#59; 2&#41; grup de baixa intensitat &#91;BI&#93;&#59; 3&#41; grup d&#8217;alta intensitat &#91;AI&#93;&#44; o 4&#41; grup control ob&#232;s &#91;CO&#93;&#46; Els ratolins OB moriren abans del programa d&#8217;entrenament&#46; BI i AI entrenaren 5 dies per setmana durant 12 setmanes en una cinta de c&#243;rrer motoritzada&#46; Els CO serviren de control no entrenats&#46; Tots els ratolins van ser ficats en g&#224;bies bin&#224;ries&#44; la temperatura de la sala dels animals es mantingu&#233; a 22 &#176;C i es cre&#224; un cicle artificial de 12 h de llum i 12 h de foscor&#46; S&#8217;aplic&#224; un per&#237;ode de familiaritzaci&#243; de 2 setmanes en qu&#232; els ratolins corrien 7-10 m durant 10-15 min en un pendent del 5&#37;&#46; Despr&#233;s&#44; es continu&#224; l&#8217;entrenament a les 12 setmanes seg&#252;ents&#44; durant 60 min&#44; de forma cont&#237;nua a 15 m&#47;min en un pendent del 5&#37; &#40;els BI&#41; i&#47;o 41 min cont&#237;nuament a 22 m&#47;min en un pendent del 5&#37; &#40;els AI&#41;&#46; El treball diari total es f&#233;u coincidir en ambd&#243;s grups i es fix&#224; en 900 m la dist&#224;ncia de carrera&#46; Els ratolins CO es mantingueren sedentaris a la g&#224;bia durant el programa d&#8217;entrenament&#46; El pes es registr&#224; setmanalment&#46; BI&#44; AI i CO van ser morts al final del programa d&#8217;entrenament&#46; Se&#8217;ls retir&#224; el menjar 12-14 h abans de morir&#46; Els experiments van ser aprovats pel Comit&#232; d&#8217;&#232;tica d&#8217;investigaci&#243;&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Preparaci&#243; del m&#250;scul </span></p><p class="elsevierStylePara"> Els ratolins van ser anestesiats per via intraperitoneal amb una preparaci&#243; de ketamina &#40;30-50 mg&#47;kg en perfusi&#243; cont&#237;nua &#91;pc&#93;&#41; i xilacina &#40;3-5 mg&#47;kg pc&#41; i moriren per extracci&#243; de sang del cor per prendre mostres del m&#250;scul 48 h despr&#233;s de la darrera sessi&#243; d&#8217;exercici&#44; i es van dissecar els m&#250;sculs soli i gastrocnemi&#46; Les mostres de m&#250;scul es congelaren r&#224;pidament en nitrogen l&#237;quid i es conservaren a &#8722;80 &#176;C&#46; Els procediments de preparaci&#243; dels teixits i l&#8217;extracci&#243; d&#8217;ARN total estan a bastament descrits en diversos estudis<span class="elsevierStyleSup">26</span>&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">PCR en temps real </span></p><p class="elsevierStylePara"> S&#8217;utilitz&#224; ARN transcriptasa inversa o retrotranscripci&#243; i encebadors d&#8217;hex&#224;mers aleatoris segons les instruccions del fabricant &#40;AccuPower Green Star qPCR PreMix&#44; BiONEER&#44; Daejeon&#44; Corea&#41;&#46; Despr&#233;s&#44; s&#8217;hi afegiren PCR Master Mix que contenien encebadors espec&#237;fics&#44; ADN Taq Hotstar polimerasa i tamp&#243; de SYBR-Green PCR&#46; Totes les mostres foren determinades per duplicat&#44; i per a un control negatiu s&#8217;utilitz&#224; la mateixa configuraci&#243;&#44; tret de l&#8217;addici&#243; de la transcriptasa inversa&#46; No es detect&#224; cap producte de PCR en aquestes darreres condicions&#46;</p><p class="elsevierStylePara"> Els nivells d&#8217;ARNm de GLUT4 i &#946;-actina&#44; un gen de manteniment&#44; es determinaren per transcripci&#243; inversa quantitativa-PCR de l&#8217;ARN cel&#183;lular total amb l&#8217;encebador directe 5&#180;CCG CGG CCT CCT ATG AGA TAC T3&#180; i l&#8217;encebador revers 5&#180;AGG CAC CCC GAA GAT GAG T3&#180; per a l&#8217;amplificaci&#243; de l&#8217;ARNm del GLUT4 i l&#8217;encebador directe 5&#180; ACA ATG AGC TGC GTG TGG CC 3&#180;i l&#8217;encebador revers 5&#180;CCT CGT AGA TGG GCA CAG TG 3&#180;per a l&#8217;amplificaci&#243; de &#946;-actina ARNm&#46; Els productes d&#8217;amplificaci&#243; van ser sotmesos a electroforesi en gels d&#8217;agarosa al 2&#37; i es van tenyir amb bromur d&#8217;etidi&#46;</p><p class="elsevierStylePara"> La quantificaci&#243; en temps real de GLUT4 a &#946;-actina d&#8217;ARNm es realitz&#224; mitjan&#231;ant un assaig de PCR SYBR-Green &#40;Rotor-gene 6000&#44; Corbett&#41;&#46; GLUT4 ARNm i &#946;-actina d&#8217;ARNm foren amplificats en tubs separats i el protocol de ciclat t&#232;rmic de 40 cicles consist&#237; en la desnaturalitzaci&#243; a 95 &#176;C durant 20 s&#44; hibridaci&#243; a 60 &#176;C durant 60 s&#44; extensi&#243; a 72 &#176;C durant 30 s que s&#8217;inici&#224; amb una desnaturalitzaci&#243; inicial a 95 &#176;C durant 15 min i es complet&#224; amb una extensi&#243; final a 72 &#176;C durant 10 min&#46; Durant l&#8217;etapa d&#8217;extensi&#243;&#44; l&#8217;augment de la fluoresc&#232;ncia fou mesurat en temps real&#46; Les dades van ser obtingudes com a valors CT &#40;valor del cicle llindar&#41;&#46; L&#8217;expressi&#243; g&#232;nica relativa fou calculada amb la f&#243;rmula de Pfaffl<span class="elsevierStyleSup">27</span>&#58;</p><p class="elsevierStylePara"><img src="278v51n191-90458592fig1.jpg"></img></p><p class="elsevierStylePara"><span class="elsevierStyleBold">An&#224;lisi de dades </span></p><p class="elsevierStylePara"> Els valors es presenten com a mitjana &#177; DE&#46; S&#8217;utilitz&#224; l&#8217;an&#224;lisi de vari&#224;ncia &#40;ANOVA&#41; d&#8217;una via seguit del test <span class="elsevierStyleItalic">Honestly-significant-difference</span> &#40;HSD&#41; de Tukey per a avaluar els efectes de l&#8217;entrenament sobre l&#8217;expressi&#243; relativa d&#8217;ARNm de GLUT4 en diferents grups&#46; Tamb&#233; s&#8217;utilitz&#224; la prova <span class="elsevierStyleItalic">t</span> aparellada&#44; per valorar la difer&#232;ncia entre l&#8217;ARNm del GLUT4 dels m&#250;sculs soli i gastrocnemi de cada grup&#46; En tots els tests s&#8217;utilitz&#224; el programari SPSS 21&#46;0 &#40;SPSS Inc&#46;&#44; Chicago&#44; IL&#44; EUA&#41;&#44; i el valor de significaci&#243; estad&#237;stica s&#8217;establ&#237; en p &#8804; 0&#44;05&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Resultats </span></p><p class="elsevierStylePara"> A la figura 1 es presenta la mitjana del valors del pes dels ratolins dels diferents grups durant la dieta alta en greixos &#40;DAG&#41; i les fases d&#8217;entrenament&#46; El pes dels grups que consumiren DAG &#40;&#233;s a dir&#44; OB&#44; CO&#44; BI i AI&#41; augment&#224; significativament en comparaci&#243; amb BN a partir de la cinquena setmana durant la fase DAG que es mantingu&#233; alta fins a la setmana 12 &#40;p &#8804; 0&#44;05&#41;&#46; L&#8217;augment de pes dels 4 grups tractats fou del 18&#37; de mitjana en comparaci&#243; amb BN&#44; que assol&#237; el 32&#37; a la dotzena setmana &#40;fig&#46; 1&#41;&#46;</p><p class="elsevierStylePara"><img alt="Figura 1 Canvi de pes de ratolins durant les fases d’alimentaci&#243; DAG i d’entrenament&#46; &#42;&#58; difer&#232;ncia significativa entre els grups obesos i normal&#59; †&#58; difer&#232;ncia significativa entre grups d’entrenament i OB&#59; ‡&#58; difer&#232;ncies significatives entre els grups AI&#47;BI&#46; AI&#58; entrenament a alta intensitat&#59; BI&#58; entrenament a baixa intensitat&#59; BN&#58; base normal&#59; CO&#58; control obesos&#59; OB&#58; obesos base&#46;" src="278v51n191-90458592fig2.jpg"></img></p><p class="elsevierStylePara"><span class="elsevierStyleBold">Figura 1 </span> Canvi de pes de ratolins durant les fases d&#8217;alimentaci&#243; DAG i d&#8217;entrenament&#46; &#42;&#58; difer&#232;ncia significativa entre els grups obesos i normal&#59; &#8224;&#58; difer&#232;ncia significativa entre grups d&#8217;entrenament i OB&#59; &#8225;&#58; difer&#232;ncies significatives entre els grups AI&#47;BI&#46; AI&#58; entrenament a alta intensitat&#59; BI&#58; entrenament a baixa intensitat&#59; BN&#58; base normal&#59; CO&#58; control obesos&#59; OB&#58; obesos base&#46;</p><p class="elsevierStylePara"> Durant la fase d&#8217;entrenament nom&#233;s s&#8217;estudiaren els grups d&#8217;entrenament &#40;BI i AI&#41; i CO&#44; i la p&#232;rdua de pes dels grups d&#8217;entrenament comen&#231;&#224; a diferenciar-se del grup CO des de la sisena setmana&#44; i esdevingu&#233; significativa a partir de la vuitena setmana &#40;p &#8804; 0&#44;05&#41;&#46; Per tant&#44; la p&#232;rdua de pes en els grups d&#8217;entrenament &#40;en comparaci&#243; amb el grup CO&#41; fou del 5 i del 7&#37; en la sisena i dotzena setmana de la fase d&#8217;entrenament&#44; respectivament&#46; &#201;s interessant assenyalar que la difer&#232;ncia de la p&#232;rdua de pes entre BI i AI aparegu&#233; a partir de la setmana 9 i arrib&#224; a ser estad&#237;sticament divergent a les setmanes 11 i 12 &#40;p &#8804; 0&#44;05&#41;&#46; De fet&#44; la r&#224;tio de la p&#232;rdua de pes fou major en AI que en BI&#44; de manera que els canvis en el percentatge de pes al final de l&#8217;entrenament de 12 setmanes del grup AI fou 8&#37;<span class="elsevierStyleItalic"> vs </span>5&#44;5&#37; del grup BI&#46;</p><p class="elsevierStylePara"> L&#8217;expressi&#243; de l&#8217;ARNm del GLUT4 del m&#250;scul soli en els ratolins que participaren en el grup BI augment&#224; aproximadament 2&#44;2 vegades&#44; enfront a ~ 1&#44;6 vegades en el gastrocnemi&#44; en relaci&#243; a BN &#40;p &#61; 0&#44;05&#41;&#59; de manera similar&#44; respecte a l&#8217;expressi&#243; de l&#8217;ARNm del GLUT4 increment&#224;&#44; tot i que&#44; estad&#237;sticament&#44; de manera no significativa&#44; per la pr&#224;ctica d&#8217;exercici d&#8217;alta intensitat en els m&#250;sculs soli i gastrocnemi ~ 2&#44;1 i ~ 1&#44;8 vegades&#44; respectivament &#40;p &#8804; 0&#44;05&#41;&#46; A m&#233;s&#44; l&#8217;expressi&#243; d&#8217;ARNm del GLUT4 dels m&#250;sculs soli i gastrocnemi en els grups BI i AI fou significativament major que en els grups OB i CO &#40;p &#8804; 0&#44;05&#41;&#44; amb una lleugera regulaci&#243; en descens <span class="elsevierStyleItalic">&#40;down regulation&#41;</span> en els darrers grups &#40;fig&#46; 2&#41;&#46;</p><p class="elsevierStylePara"><img alt="Figura 2 Contingut d’ARNm de GLUT4 dels grups experimentals gastrocnemi i soli en relaci&#243; amb el m&#250;scul emparellat en el grup BN &#40;p &#8804; 0&#44;05&#41;&#46; &#42;&#58; augment significatiu amb relaci&#243; a BN&#59; †&#58; difer&#232;ncia significativa en comparaci&#243; amb OB i CO &#40;p &#8804; 0&#44;05&#41;&#46; " src="278v51n191-90458592fig3.jpg"></img></p><p class="elsevierStylePara"><span class="elsevierStyleBold">Figura 2 </span>Contingut d&#8217;ARNm de GLUT4 dels grups experimentals gastrocnemi i soli en relaci&#243; amb el m&#250;scul emparellat en el grup BN &#40;p &#8804; 0&#44;05&#41;&#46; &#42;&#58; augment significatiu amb relaci&#243; a BN&#59; &#8224;&#58; difer&#232;ncia significativa en comparaci&#243; amb OB i CO &#40;p &#8804; 0&#44;05&#41;&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Discussi&#243; </span></p><p class="elsevierStylePara"> Ploug et al&#46;<span class="elsevierStyleSup">28</span> establiren que GLUT4 en fibres no estimulades &#40;per la insulina&#41; es distribu&#239;a al llarg de totes les fibres musculars&#44; i era present tant en la superf&#237;cie &#40;68&#37; del GLUT4 total&#41; com en el nucli &#40;32&#37; del GLUT4 total&#41; de les fibres&#46; Tamb&#233; trobaren que els nuclis eren despla&#231;ats i alineats amb els vasos sanguinis que cursen al llarg de la superf&#237;cie de la fibra&#44; que ocasionen una gran fracci&#243; de GLUT4 prop de la font de la glucosa evitant la necessitat de difusi&#243; a trav&#233;s de llargues dist&#224;ncies&#46; Est&#224; ben documentat que&#44; en estat normal&#44; el GLUT4 circula lentament entre la membrana plasm&#224;tica i un o m&#233;s compartiments intracel&#183;lulars&#44; amb la gran majoria de transportadors que resideixen en els compartiments vesiculars dins de l&#8217;interior de la c&#232;l&#183;lula<span class="elsevierStyleSup">28&#44;29</span>&#46;</p><p class="elsevierStylePara"> L&#8217;acumulaci&#243; d&#8217;insulina estimulada de la prote&#239;na GLUT4 en la superf&#237;cie de la c&#232;l&#183;lula pot ser causada per un increment de 10 a 20 vegades de la r&#224;tio d&#8217;exocitosi amb una petita disminuci&#243; &#40;2-3 vegades&#41; de la r&#224;tio d&#8217;endocitosi del GLUT4<span class="elsevierStyleSup">10&#44;45&#44;30</span>&#46;</p><p class="elsevierStylePara"> La insulina estimula el transport de glucosa a trav&#233;s de la translocaci&#243; de GLUT4 a partir de l&#8217;agrupaci&#243; d&#8217;emmagatzemament intracel&#183;lular cap a la membrana plasm&#224;tica&#46; La resist&#232;ncia a la insulina causada per la manca d&#8217;insulina o uns resultats anormals de senyalitzaci&#243; de la insulina amb un descens en l&#8217;expressi&#243; y translocaci&#243; de GLUT4 causen la hipergluc&#232;mia i la diabetis<span class="elsevierStyleSup">30-32</span>&#46; Tamb&#233; s&#8217;ha demostrat que la contracci&#243; muscular augmenta el contingut de GLUT4 de la membrana cel&#183;lular<span class="elsevierStyleSup">29&#44;30</span>&#46; Diversos estudis suggereixen que l&#8217;exercici f&#237;sic causa l&#8217;expressi&#243; de la prote&#239;na GLUT4 i la translocaci&#243; en membrana plasm&#224;tica mitjan&#231;ant un mecanisme diferent de senyalitzaci&#243; de la insulina<span class="elsevierStyleSup">5&#44;33</span>&#46; La majoria d&#8217;estudis sobre el m&#250;scul esquel&#232;tic dels ratolins indiquen que hi ha m&#233;s GLUT4 en les fibres de tipus i que en les del II<span class="elsevierStyleSup">10&#44;34</span>&#44; la qual cosa coincideix amb els resultats que vam trobar en el ratolins&#46; En aquest estudi nom&#233;s vam utilitzar el m&#250;scul soli&#44; generalment conegut com a m&#250;scul de fibres de tipus I&#46;</p><p class="elsevierStylePara"> El nostre estudi concorda amb altres<span class="elsevierStyleSup">35-39</span> que han indicat que l&#8217;exercici f&#237;sic incrementa l&#8217;expressi&#243; de la prote&#239;na GLUT4 en subjectes diab&#232;tics&#44; tal com succe&#237; en els nostres subjectes obesos&#46;</p><p class="elsevierStylePara"> Vam veure que l&#8217;expressi&#243; d&#8217;ARNm de GLUT4 s&#8217;incrementa amb 12 setmanes d&#8217;entrenament de resist&#232;ncia&#44; en ambd&#243;s m&#250;sculs&#44; en el soli &#40;m&#250;scul en qu&#232; predominen les fibres de contracci&#243; lenta&#41; i en el gastrocnemi &#40;en qu&#232; predominen les fibres de contracci&#243; r&#224;pida&#41; en ratolins obesos&#44; amb patr&#243; intens de depend&#232;ncia&#46; Aix&#242; significa que la intensitat de l&#8217;exercici pot afectar l&#8217;expressi&#243; del gen GLUT4 &#40;i&#44; probablement&#44; la prote&#239;na&#41; en resposta a l&#8217;entrenament&#44; la qual cosa pot provocar canvis metab&#242;lics significatius especialment en els casos obesos&#46; En conson&#224;ncia amb els nostres resultats&#44; s&#8217;ha demostrat que la prote&#239;na GLUT4 i el transport de glucosa s&#243;n immensament superiors en les fibres musculars oxidatives vermelles &#40;tipus I i IIa&#41; que en les fibres glucol&#237;tiques blanques &#40;tipus IIb&#41;<span class="elsevierStyleSup">10&#44;40</span>&#46; Aix&#242; podria ser un factor important en l&#8217;adaptaci&#243; a l&#8217;exercici&#44; degut a qu&#232; els resultats de l&#8217;entrenament de resist&#232;ncia faciliten un canvi de fibres de tipus iib a fibres tipus IIa<span class="elsevierStyleSup">41</span>&#46; Tanmateix&#44; en els m&#250;sculs esquel&#232;tics humans hi ha una difer&#232;ncia molt menor en l&#8217;expressi&#243; de GLUT4 en diferents tipus de fibres musculars<span class="elsevierStyleSup">34&#44;42</span>&#46; Daugaard et al&#46;<span class="elsevierStyleSup">34</span> van a&#239;llar fibres musculars individuals i les van classificar segons la isoforma de la miosina&#46; GLUT4 fou &#8764; 20&#37; m&#233;s elevada en les fibres que expressen la cadena pesada de la miosina i que en las que expressen la cadena pesada de la miosina IIa o IIb&#46; No es van trobar difer&#232;ncies entre les fibres IIa i IIb&#46; Despr&#233;s de 2 setmanes d&#8217;entrenament&#44; GLUT4 increment&#224; ~ 23&#37; de fibres musculars de tipus i&#44; per&#242; no hi hagu&#233; canvis en els tipus IIa o IIb&#46; Tanmateix&#44; l&#8217;exercici de baixa intensitat utilitzat &#233;s conegut principalment per reclutar fibres de tipus I&#46;</p><p class="elsevierStylePara"> D&#8217;acord amb els nostres resultats&#44; l&#8217;entrenament de baixa intensitat origin&#224; una expressi&#243; major d&#8217;ARNm de GLUT4 en el m&#250;scul oxidatiu &#40;&#233;s a dir&#44; el soli&#41;&#44; probablement degut a les necessitats relacionades amb la capacitat oxidativa de les c&#232;l&#183;lules&#46; D&#8217;altra banda&#44; la pr&#224;ctica d&#8217;exercici d&#8217;alta intensitat no produ&#237; una expressi&#243; major d&#8217;ARNm de GLUT4 en el soli ni en el gastrocnemi&#46; Com que el m&#250;scul gastrocnemi consisteix en una barreja de fibres vermelles i blanques&#44; una altra explicaci&#243; podria ser que l&#8217;entrenament amb exercicis de baixa intensitat no fou suficient per exercir o fins i tot per activar les fibres vermelles del m&#250;scul gastrocnemi&#44; i un augment no significatiu d&#8217;aquest m&#250;scul podria procedir de les fibres blanques&#46; Aquest escenari pot ampliar-se al que s&#8217;ha observat en les troballes relacionades amb l&#8217;alta intensitat&#59; &#233;s a dir&#44; un augment no significatiu d&#8217;aquest m&#250;scul amb l&#8217;exercici d&#8217;alta intensitat podria procedir nom&#233;s de les fibres vermelles&#46;</p><p class="elsevierStylePara"> Les hormones tamb&#233; regulen la concentraci&#243; de la prote&#239;na del m&#250;scul transportador de la glucosa&#46; L&#8217;expressi&#243; de GLUT4 s&#8217;increment&#224; degut a la insulina i a les hormones tiroidees<span class="elsevierStyleSup">20&#44;43</span> i disminu&#237; per un AMPc elevat<span class="elsevierStyleSup">44</span>&#46; La regulaci&#243; de l&#8217;expressi&#243; de GLUT4 per l&#8217;activitat contr&#224;ctil &#233;s independent de la regulaci&#243; hormonal&#44; perqu&#232; la carrera en la cinta de c&#243;rrer augment&#224; el GLUT4 en els ratolins diab&#232;tics<span class="elsevierStyleSup">45</span>&#46; S&#8217;hi afegiren els efectes de la defici&#232;ncia de la insulina i denervaci&#243; en les concentracions de GLUT4<span class="elsevierStyleSup">45</span>&#46;</p><p class="elsevierStylePara"> L&#8217;entrenament f&#237;sic tamb&#233; augmenta la prote&#239;na del GLUT4 i l&#8217;ARNm muscular en pacients amb diabetis tipus 2<span class="elsevierStyleSup">46</span>&#46; Aquests resultats suggereixen que l&#8217;activitat contr&#224;ctil del m&#250;scul modula directament l&#8217;expressi&#243; de GLUT4 del m&#250;scul&#44; independentment de l&#8217;acci&#243; de la insulina&#46; La histona deacetilasa 5 &#40;HDAC5&#41; &#233;s un mediador cr&#237;tic dels canvis del nivell d&#8217;ARNm de GLUT4 en resposta a l&#8217;entrenament<span class="elsevierStyleSup">47</span>&#46; S&#8217;ha demostrat que una s&#232;rie de modificacions de les histones causades per la histona deacetilasa i la histona metiltransferasa culminaran en una caiguda metab&#242;lica del gen GLUT4 en el m&#250;scul esquel&#232;tic dels ratolins<span class="elsevierStyleSup">48</span>&#46;</p><p class="elsevierStylePara"> Hi ha evid&#232;ncies significatives que els nivells d&#8217;ARNm de GLUT4 disminueixen en el m&#250;scul del teixit adip&#243;s amb l&#8217;obesitat<span class="elsevierStyleSup">49&#44;50</span>&#44; y que l&#8217;augment d&#8217;ARNm de GLUT4 pot alleujar o compensar la resist&#232;ncia a la insulina<span class="elsevierStyleSup">51&#44;52</span>&#46; Tal com era d&#8217;esperar&#44; les nostres troballes sobre els nivells d&#8217;ARNm del GLUT4 augmentaren significativament amb l&#8217;entrenament&#44; i aquest increment s&#8217;associ&#224; a una p&#232;rdua de pes&#44; la qual cosa recolza aquesta idea&#46; Per tant&#44; l&#8217;augment de la transcripci&#243; de GLUT4 en el m&#250;scul durant la p&#232;rdua de pes pot ser una prova definitiva en la reversi&#243; de la resist&#232;ncia a la insulina&#46;</p><p class="elsevierStylePara"> Estudis anteriors&#44; juntament amb els nostres resultats&#44; suggereixen que la regulaci&#243; transcripcional del gen GLUT4 &#233;s sensible a canvis en l&#8217;equilibri d&#8217;energia del cos&#46; A m&#233;s&#44; nosaltres vam trobar una resposta de GLUT4 a l&#8217;entrenament depenent de la intensitat i del tipus de fibra&#44; que no es plantejava en estudis anteriors&#46;</p><p class="elsevierStylePara"> Podem concloure que tot tipus d&#8217;alteraci&#243; de l&#8217;equilibri energ&#232;tic del cos&#44; especialment per la pr&#224;ctica d&#8217;exercici i&#47;o dieta alta en greixos&#44; pot influir en els mecanismes moleculars i cel&#183;lulars que actuen per establir una home&#242;stasi estable&#46; GLUT4 podria disminuir si s&#8217;augment&#233;s la ingesta d&#8217;energia&#44; i a la inversa&#44; augmentaria si s&#8217;augment&#233;s la despesa energ&#232;tica&#46; Aquests canvis s&#243;n gaireb&#233; similars en els m&#250;sculs de contracci&#243; lenta i r&#224;pida&#44; amb una mica m&#233;s de capacitat de resposta en els m&#250;sculs de contracci&#243; lenta&#44; perqu&#232; estan m&#233;s compromesos amb el metabolisme de la glucosa&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Conflicte d&#8217;interessos </span></p><p class="elsevierStylePara"> Els autors declaren que no tenen cap conflicte d&#8217;interessos</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Agra&#239;ment </span></p><p class="elsevierStylePara"> Els autors agraeixen a la Universitat de Guilan el suport econ&#242;mic atorgat a aquest projecte d&#8217;investigaci&#243;&#46;</p><hr></hr><p class="elsevierStylePara"> Rebut el 10 d&#8217;octubre de 2015&#59; <br></br> acceptat el 14 de desembre de 2015</p><p class="elsevierStylePara"> &#42; Autor per a correspond&#232;ncia&#46;<br></br><span class="elsevierStyleItalic">Correu electr&#242;nic&#58;</span><a href="mailto&#58;mohebbi-h&#64;yahoo&#46;com" class="elsevierStyleCrossRefs">mohebbi-h&#64;yahoo&#46;com</a> &#40;H&#46; Mohebbi&#41;&#46;</p>"
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            0 => "Entrenament de resist&#232;ncia"
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            0 => "Endurance training"
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        "resumen" => "<p class="elsevierStylePara"> L&#8217;objectiu d&#8217;aquest estudi fou investigar la resposta a l&#8217;entrenament de resist&#232;ncia de l&#8217;expressi&#243; de l&#8217;ARNm del GLUT4 dels m&#250;sculs soli i gastrocnemi en ratolins obesos&#46; En aquest estudi es van utilitzar 40 ratolins mascles C57BL&#47;6&#46; Vuit ratolins &#40;base normal &#91;BN&#93;&#41; serviren com a controls no obesos i no entrenats&#44; i 32 ratolins foren sotmesos a una dieta alta en greixos &#40;DAG&#41; &#40;augment del 60&#37; de kcal&#41; durant 12 setmanes&#46; A la setmana 16 els ratolins obesos foren distribu&#239;ts aleat&#242;riament en els grups de tractament seg&#252;ents &#40;n &#61; 8 cada grup&#41;&#58; obesos base &#40;OB&#41;&#59; baixa intensitat &#40;BI&#41;&#59; alta intensitat &#40;AI&#41;&#44; o grups de control d&#8217;obesitat &#40;CO&#41;&#46; BI i AI foren entrenats 5 dies per setmana durant 12 setmanes en una cinta de c&#243;rrer motoritzada a 15 m&#47;min amb un pendent del 5&#37; &#40;en BI&#41; i&#47;o en 22 m&#47; min en un pendent del 5&#37; &#40;en AI&#41;&#46; Els ratolins CO es mantingueren sedentaris a la g&#224;bia durant el programa d&#8217;entrenament&#46; Es mesur&#224; l&#8217;expressi&#243; de l&#8217;ARNm del GLUT4 dels m&#250;sculs gastrocnemi i soli&#44; utilitzant el m&#232;tode PCR en temps real&#46; L&#8217;expressi&#243; de l&#8217;ARNm del GLUT4 del m&#250;scul soli en el grup BI augment&#224; aproximadament 2&#44;2 vegades&#44; enfront a 1&#44;6 vegades en el gastrocnemi &#40;p &#60; 0&#44;05&#41;&#46; A m&#233;s&#44; l&#8217;expressi&#243; de l&#8217;ARNm del GLUT4 en els m&#250;sculs soli i gastrocnemi dels grups BI i AI fou significativament major que en OB i en els grups CO &#40;p &#60; 0&#44;05&#41;&#46; Podem concloure que tota alteraci&#243; de l&#8217;equilibri energ&#232;tic del cos&#44; especialment per l&#8217;entrenament f&#237;sic i&#47;o una dieta alta en greixos&#44; pot influir en els mecanismes moleculars i cel&#183;lulars que actuen per establir una home&#242;stasi estable&#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;n &#61; 8 each group&#41;&#58; obese base &#91;OB&#93;&#59; low intensity &#91;LI&#93;&#59; high intensity &#91;HI&#93;&#44; 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&#44; can influence these molecular and cellular mechanisms that act to establish a stable homeostasis&#46;</p>"
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Efectes de 12 setmanes d’entrenament de resistència a 2 intensitats diferents en l’expressió de l’ARNm del GLUT4 dels músculs soli i gastrocnemi en ratolins obesos
The effect of 12 weeks endurance training at 2 different intensities on GLUT4 mRNA expression of soleus and gastrocnemius muscles in obese mice
Hamid Mohebbia, Hadi Rohanib, Sadegh Hassan-Niac
a Department of Exercice Physiology, Faculty of Physical Education and Sport Sciences, University of Guilan, Rasht, Iran
b Department of Sport Medicine, Sport Sciences Research Institute of Iran, Teheran, Iran
c Faculty of Biological Science, Tarbiat Modares University, Teheran, Iran
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    "textoCompleto" => "<p class="elsevierStylePara"> </p><span class="elsevierStyleBold">Introducci&#243; </span><p class="elsevierStylePara"> Generalment s&#8217;admet que l&#8217;obesitat est&#224; fonamentalment associada a una alteraci&#243; de la taxa d&#8217;absorci&#243; de glucosa estimulada per la insulina en el m&#250;scul esquel&#232;tic&#44; que s&#8217;ha atribu&#239;t a la resist&#232;ncia a la insulina&#46; Diversos estudis s&#8217;han centrat en el sistema de transport de la glucosa com a part dels mecanismes subjacents&#46; La glucosa transporta les c&#232;llules del m&#250;scul esquel&#232;tic a trav&#233;s de les prote&#239;nes transportadores de glucosa GLUT1 i GLUT4<span class="elsevierStyleSup">1</span>&#46; Sembla que la isoforma GLUT1 &#233;s la transportadora de la glucosa que suporta el transport de la glucosa basal<span class="elsevierStyleSup">2&#44;3</span>&#44; mentre que la isoforma GLUT4 augmenta el transport de glucosa en resposta a la insulina i a la contracci&#243;&#46; La insulina i les contraccions transporten el GLUT4 de la reserva intracel&#183;lular a la membrana plasm&#224;tica i als t&#250;buls T<span class="elsevierStyleSup">4&#44;5</span>&#46;</p><p class="elsevierStylePara"> Se sap que en els rosegadors&#44; la capacitat d&#8217;absorci&#243; de la glucosa &#233;s major en els m&#250;sculs oxidatius vermells que en els m&#250;sculs glucol&#237;tics blancs<span class="elsevierStyleSup">6-10</span>&#46; Sembla que un mecanisme subjacent t&#233; un major nivell d&#8217;expressi&#243; de GLUT4&#44; tant a nivell intracel&#183;lular<span class="elsevierStyleSup">6&#44;7</span> com a la membrana plasm&#224;tica8&#46; En el m&#250;scul esquel&#232;tic hum&#224;&#44; la captaci&#243; de glucosa s&#8217;associa positivament a la proporci&#243; de fibres de tipus i i negativament a la proporci&#243; de fibres de tipus iib<span class="elsevierStyleSup">11</span>&#46; Aquests resultats es basen en l&#8217;estudi in vitro de Zierath et al&#46;<span class="elsevierStyleSup">12</span>&#44; que reportaren que l&#8217;augment d&#8217;insulina estimulada en la captaci&#243; de la glucosa per damunt del nivell basal correlaciona estretament&#44; tant afirmativament amb el percentatge de fibres musculars de tipus i&#44; com negativament amb el percentatge de fibres tipus iia&#46; Tanmateix&#44; s&#8217;han reportat resultats poc convincents respecte a la relaci&#243; entre la distribuci&#243; del tipus de fibra i el contingut de GLUT4 en el m&#250;scul hum&#224;<span class="elsevierStyleSup">13-15</span>&#46;</p><p class="elsevierStylePara"> Andersen et al&#46;<span class="elsevierStyleSup">13</span> no trobaren cap correlaci&#243; entre tipus de fibra i contingut de GLUT4&#44; mentre que Houmard et al&#46;<span class="elsevierStyleSup">15</span> mostraren una baixa correlaci&#243; entre tipus de composici&#243; de la fibra i contingut de GLUT4&#46;</p><p class="elsevierStylePara"> Ha estat demostrat que els diferents m&#250;sculs presenten importants difer&#232;ncies de contingut de GLUT4&#44; i aquestes difer&#232;ncies sovint estan associades a les difer&#232;ncies en la captaci&#243; de la glucosa estimulada per la insulina<span class="elsevierStyleSup">16&#44;17</span>&#46; Com que el diferents m&#250;sculs es composen d&#8217;una combinaci&#243; de diferents tipus de fibres musculars<span class="elsevierStyleSup">18</span>&#44; &#233;s possible que hi hagi difer&#232;ncies significatives en el contingut de GLUT4 entre m&#250;sculs&#46;</p><p class="elsevierStylePara"> Probablement&#44; les difer&#232;ncies en el contingut de GLUT4 i la captaci&#243; de la glucosa estimulada per la insulina estan relacionades amb el nivell d&#8217;entrenament&#46; Els canvis de nivell d&#8217;activitat del m&#250;scul esquel&#232;tic s&#243;n la clau reguladora del contingut de GLUT4 en ratolins<span class="elsevierStyleSup">19&#44;20</span>&#46; Entre els humans&#44; els esportistes tenen m&#233;s GLUT4 que els subjectes control de la mateixa edat no entrenats<span class="elsevierStyleSup">21&#44;22</span>&#44; i en ambd&#243;s&#44; subjectes control normals i sans i individus amb la captaci&#243; disminu&#239;da de la glucosa estimulada per la insulina&#44; l&#8217;entrenament f&#237;sic ha demostrat que incrementa el contingut de GLUT4<span class="elsevierStyleSup">20&#44;23&#44;24</span>&#46; A m&#233;s&#44; una disminuci&#243; del nivell d&#8217;activitat pot minorar el contingut de GLUT4<span class="elsevierStyleSup">20&#44;25</span>&#46; Finalment&#44; s&#8217;ha demostrat que canvis en l&#8217;activitat f&#237;sica i el contingut de GLUT4 estan relacionats amb canvis en la captaci&#243; de la glucosa estimulada per la insulina<span class="elsevierStyleSup">20</span>&#46;</p><p class="elsevierStylePara"> El principal objectiu d&#8217;aquest treball fou investigar l&#8217;expressi&#243; de l&#8217;ARNm del GLUT4 en el soli &#40;m&#250;scul predominantment de contracci&#243; lenta&#41; i el gastrocnemi &#40;m&#250;scul predominantment de contracci&#243; r&#224;pida&#41; com a resposta a l&#8217;entrenament de resist&#232;ncia en ratolins obesos&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Subjectes i m&#232;todes </span></p><p class="elsevierStylePara"><span class="elsevierStyleBold">Animals </span></p><p class="elsevierStylePara"> En aquest estudi s&#8217;utilitzaren 40 ratolins mascles C57BL&#47;6 &#40;de 4 setmanes&#41;&#46; Vuit ratolins no obesos &#40;base normal &#91;BN&#93;&#59; dieta est&#224;ndard alimentats <span class="elsevierStyleItalic">ad libitum</span>&#41; van servir de control no obesos no entrenats &#40;el grup BN es consider&#224; com a valor de refer&#232;ncia per calcular l&#8217;expressi&#243; g&#232;nica en temps real que s&#8217;expressa com a valors relatius&#41; i 32 ratolins van ser sotmesos a una dieta aliment&#224;ria rica en greixos &#40;DAG&#41; durant 12 setmanes que consist&#237; en acc&#233;s lliure a una dieta rica en greixos de 60&#37; kcal &#40;<span class="elsevierStyleItalic">High-Fat Diet&#44; Razi Vaccine &#38; Serum Research Institute</span>&#44; Iran&#41;&#46; A la setmana 16&#44; els ratolins obesos foren assignats aleat&#242;riament als grups de tractament seg&#252;ents &#40;n &#61; 8 cada grup&#41;&#58; 1&#41; grup ob&#232;s base &#91;OB&#93;&#59; 2&#41; grup de baixa intensitat &#91;BI&#93;&#59; 3&#41; grup d&#8217;alta intensitat &#91;AI&#93;&#44; o 4&#41; grup control ob&#232;s &#91;CO&#93;&#46; Els ratolins OB moriren abans del programa d&#8217;entrenament&#46; BI i AI entrenaren 5 dies per setmana durant 12 setmanes en una cinta de c&#243;rrer motoritzada&#46; Els CO serviren de control no entrenats&#46; Tots els ratolins van ser ficats en g&#224;bies bin&#224;ries&#44; la temperatura de la sala dels animals es mantingu&#233; a 22 &#176;C i es cre&#224; un cicle artificial de 12 h de llum i 12 h de foscor&#46; S&#8217;aplic&#224; un per&#237;ode de familiaritzaci&#243; de 2 setmanes en qu&#232; els ratolins corrien 7-10 m durant 10-15 min en un pendent del 5&#37;&#46; Despr&#233;s&#44; es continu&#224; l&#8217;entrenament a les 12 setmanes seg&#252;ents&#44; durant 60 min&#44; de forma cont&#237;nua a 15 m&#47;min en un pendent del 5&#37; &#40;els BI&#41; i&#47;o 41 min cont&#237;nuament a 22 m&#47;min en un pendent del 5&#37; &#40;els AI&#41;&#46; El treball diari total es f&#233;u coincidir en ambd&#243;s grups i es fix&#224; en 900 m la dist&#224;ncia de carrera&#46; Els ratolins CO es mantingueren sedentaris a la g&#224;bia durant el programa d&#8217;entrenament&#46; El pes es registr&#224; setmanalment&#46; BI&#44; AI i CO van ser morts al final del programa d&#8217;entrenament&#46; Se&#8217;ls retir&#224; el menjar 12-14 h abans de morir&#46; Els experiments van ser aprovats pel Comit&#232; d&#8217;&#232;tica d&#8217;investigaci&#243;&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Preparaci&#243; del m&#250;scul </span></p><p class="elsevierStylePara"> Els ratolins van ser anestesiats per via intraperitoneal amb una preparaci&#243; de ketamina &#40;30-50 mg&#47;kg en perfusi&#243; cont&#237;nua &#91;pc&#93;&#41; i xilacina &#40;3-5 mg&#47;kg pc&#41; i moriren per extracci&#243; de sang del cor per prendre mostres del m&#250;scul 48 h despr&#233;s de la darrera sessi&#243; d&#8217;exercici&#44; i es van dissecar els m&#250;sculs soli i gastrocnemi&#46; Les mostres de m&#250;scul es congelaren r&#224;pidament en nitrogen l&#237;quid i es conservaren a &#8722;80 &#176;C&#46; Els procediments de preparaci&#243; dels teixits i l&#8217;extracci&#243; d&#8217;ARN total estan a bastament descrits en diversos estudis<span class="elsevierStyleSup">26</span>&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">PCR en temps real </span></p><p class="elsevierStylePara"> S&#8217;utilitz&#224; ARN transcriptasa inversa o retrotranscripci&#243; i encebadors d&#8217;hex&#224;mers aleatoris segons les instruccions del fabricant &#40;AccuPower Green Star qPCR PreMix&#44; BiONEER&#44; Daejeon&#44; Corea&#41;&#46; Despr&#233;s&#44; s&#8217;hi afegiren PCR Master Mix que contenien encebadors espec&#237;fics&#44; ADN Taq Hotstar polimerasa i tamp&#243; de SYBR-Green PCR&#46; Totes les mostres foren determinades per duplicat&#44; i per a un control negatiu s&#8217;utilitz&#224; la mateixa configuraci&#243;&#44; tret de l&#8217;addici&#243; de la transcriptasa inversa&#46; No es detect&#224; cap producte de PCR en aquestes darreres condicions&#46;</p><p class="elsevierStylePara"> Els nivells d&#8217;ARNm de GLUT4 i &#946;-actina&#44; un gen de manteniment&#44; es determinaren per transcripci&#243; inversa quantitativa-PCR de l&#8217;ARN cel&#183;lular total amb l&#8217;encebador directe 5&#180;CCG CGG CCT CCT ATG AGA TAC T3&#180; i l&#8217;encebador revers 5&#180;AGG CAC CCC GAA GAT GAG T3&#180; per a l&#8217;amplificaci&#243; de l&#8217;ARNm del GLUT4 i l&#8217;encebador directe 5&#180; ACA ATG AGC TGC GTG TGG CC 3&#180;i l&#8217;encebador revers 5&#180;CCT CGT AGA TGG GCA CAG TG 3&#180;per a l&#8217;amplificaci&#243; de &#946;-actina ARNm&#46; Els productes d&#8217;amplificaci&#243; van ser sotmesos a electroforesi en gels d&#8217;agarosa al 2&#37; i es van tenyir amb bromur d&#8217;etidi&#46;</p><p class="elsevierStylePara"> La quantificaci&#243; en temps real de GLUT4 a &#946;-actina d&#8217;ARNm es realitz&#224; mitjan&#231;ant un assaig de PCR SYBR-Green &#40;Rotor-gene 6000&#44; Corbett&#41;&#46; GLUT4 ARNm i &#946;-actina d&#8217;ARNm foren amplificats en tubs separats i el protocol de ciclat t&#232;rmic de 40 cicles consist&#237; en la desnaturalitzaci&#243; a 95 &#176;C durant 20 s&#44; hibridaci&#243; a 60 &#176;C durant 60 s&#44; extensi&#243; a 72 &#176;C durant 30 s que s&#8217;inici&#224; amb una desnaturalitzaci&#243; inicial a 95 &#176;C durant 15 min i es complet&#224; amb una extensi&#243; final a 72 &#176;C durant 10 min&#46; Durant l&#8217;etapa d&#8217;extensi&#243;&#44; l&#8217;augment de la fluoresc&#232;ncia fou mesurat en temps real&#46; Les dades van ser obtingudes com a valors CT &#40;valor del cicle llindar&#41;&#46; L&#8217;expressi&#243; g&#232;nica relativa fou calculada amb la f&#243;rmula de Pfaffl<span class="elsevierStyleSup">27</span>&#58;</p><p class="elsevierStylePara"><img src="278v51n191-90458592fig1.jpg"></img></p><p class="elsevierStylePara"><span class="elsevierStyleBold">An&#224;lisi de dades </span></p><p class="elsevierStylePara"> Els valors es presenten com a mitjana &#177; DE&#46; S&#8217;utilitz&#224; l&#8217;an&#224;lisi de vari&#224;ncia &#40;ANOVA&#41; d&#8217;una via seguit del test <span class="elsevierStyleItalic">Honestly-significant-difference</span> &#40;HSD&#41; de Tukey per a avaluar els efectes de l&#8217;entrenament sobre l&#8217;expressi&#243; relativa d&#8217;ARNm de GLUT4 en diferents grups&#46; Tamb&#233; s&#8217;utilitz&#224; la prova <span class="elsevierStyleItalic">t</span> aparellada&#44; per valorar la difer&#232;ncia entre l&#8217;ARNm del GLUT4 dels m&#250;sculs soli i gastrocnemi de cada grup&#46; En tots els tests s&#8217;utilitz&#224; el programari SPSS 21&#46;0 &#40;SPSS Inc&#46;&#44; Chicago&#44; IL&#44; EUA&#41;&#44; i el valor de significaci&#243; estad&#237;stica s&#8217;establ&#237; en p &#8804; 0&#44;05&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Resultats </span></p><p class="elsevierStylePara"> A la figura 1 es presenta la mitjana del valors del pes dels ratolins dels diferents grups durant la dieta alta en greixos &#40;DAG&#41; i les fases d&#8217;entrenament&#46; El pes dels grups que consumiren DAG &#40;&#233;s a dir&#44; OB&#44; CO&#44; BI i AI&#41; augment&#224; significativament en comparaci&#243; amb BN a partir de la cinquena setmana durant la fase DAG que es mantingu&#233; alta fins a la setmana 12 &#40;p &#8804; 0&#44;05&#41;&#46; L&#8217;augment de pes dels 4 grups tractats fou del 18&#37; de mitjana en comparaci&#243; amb BN&#44; que assol&#237; el 32&#37; a la dotzena setmana &#40;fig&#46; 1&#41;&#46;</p><p class="elsevierStylePara"><img alt="Figura 1 Canvi de pes de ratolins durant les fases d’alimentaci&#243; DAG i d’entrenament&#46; &#42;&#58; difer&#232;ncia significativa entre els grups obesos i normal&#59; †&#58; difer&#232;ncia significativa entre grups d’entrenament i OB&#59; ‡&#58; difer&#232;ncies significatives entre els grups AI&#47;BI&#46; AI&#58; entrenament a alta intensitat&#59; BI&#58; entrenament a baixa intensitat&#59; BN&#58; base normal&#59; CO&#58; control obesos&#59; OB&#58; obesos base&#46;" src="278v51n191-90458592fig2.jpg"></img></p><p class="elsevierStylePara"><span class="elsevierStyleBold">Figura 1 </span> Canvi de pes de ratolins durant les fases d&#8217;alimentaci&#243; DAG i d&#8217;entrenament&#46; &#42;&#58; difer&#232;ncia significativa entre els grups obesos i normal&#59; &#8224;&#58; difer&#232;ncia significativa entre grups d&#8217;entrenament i OB&#59; &#8225;&#58; difer&#232;ncies significatives entre els grups AI&#47;BI&#46; AI&#58; entrenament a alta intensitat&#59; BI&#58; entrenament a baixa intensitat&#59; BN&#58; base normal&#59; CO&#58; control obesos&#59; OB&#58; obesos base&#46;</p><p class="elsevierStylePara"> Durant la fase d&#8217;entrenament nom&#233;s s&#8217;estudiaren els grups d&#8217;entrenament &#40;BI i AI&#41; i CO&#44; i la p&#232;rdua de pes dels grups d&#8217;entrenament comen&#231;&#224; a diferenciar-se del grup CO des de la sisena setmana&#44; i esdevingu&#233; significativa a partir de la vuitena setmana &#40;p &#8804; 0&#44;05&#41;&#46; Per tant&#44; la p&#232;rdua de pes en els grups d&#8217;entrenament &#40;en comparaci&#243; amb el grup CO&#41; fou del 5 i del 7&#37; en la sisena i dotzena setmana de la fase d&#8217;entrenament&#44; respectivament&#46; &#201;s interessant assenyalar que la difer&#232;ncia de la p&#232;rdua de pes entre BI i AI aparegu&#233; a partir de la setmana 9 i arrib&#224; a ser estad&#237;sticament divergent a les setmanes 11 i 12 &#40;p &#8804; 0&#44;05&#41;&#46; De fet&#44; la r&#224;tio de la p&#232;rdua de pes fou major en AI que en BI&#44; de manera que els canvis en el percentatge de pes al final de l&#8217;entrenament de 12 setmanes del grup AI fou 8&#37;<span class="elsevierStyleItalic"> vs </span>5&#44;5&#37; del grup BI&#46;</p><p class="elsevierStylePara"> L&#8217;expressi&#243; de l&#8217;ARNm del GLUT4 del m&#250;scul soli en els ratolins que participaren en el grup BI augment&#224; aproximadament 2&#44;2 vegades&#44; enfront a ~ 1&#44;6 vegades en el gastrocnemi&#44; en relaci&#243; a BN &#40;p &#61; 0&#44;05&#41;&#59; de manera similar&#44; respecte a l&#8217;expressi&#243; de l&#8217;ARNm del GLUT4 increment&#224;&#44; tot i que&#44; estad&#237;sticament&#44; de manera no significativa&#44; per la pr&#224;ctica d&#8217;exercici d&#8217;alta intensitat en els m&#250;sculs soli i gastrocnemi ~ 2&#44;1 i ~ 1&#44;8 vegades&#44; respectivament &#40;p &#8804; 0&#44;05&#41;&#46; A m&#233;s&#44; l&#8217;expressi&#243; d&#8217;ARNm del GLUT4 dels m&#250;sculs soli i gastrocnemi en els grups BI i AI fou significativament major que en els grups OB i CO &#40;p &#8804; 0&#44;05&#41;&#44; amb una lleugera regulaci&#243; en descens <span class="elsevierStyleItalic">&#40;down regulation&#41;</span> en els darrers grups &#40;fig&#46; 2&#41;&#46;</p><p class="elsevierStylePara"><img alt="Figura 2 Contingut d’ARNm de GLUT4 dels grups experimentals gastrocnemi i soli en relaci&#243; amb el m&#250;scul emparellat en el grup BN &#40;p &#8804; 0&#44;05&#41;&#46; &#42;&#58; augment significatiu amb relaci&#243; a BN&#59; †&#58; difer&#232;ncia significativa en comparaci&#243; amb OB i CO &#40;p &#8804; 0&#44;05&#41;&#46; " src="278v51n191-90458592fig3.jpg"></img></p><p class="elsevierStylePara"><span class="elsevierStyleBold">Figura 2 </span>Contingut d&#8217;ARNm de GLUT4 dels grups experimentals gastrocnemi i soli en relaci&#243; amb el m&#250;scul emparellat en el grup BN &#40;p &#8804; 0&#44;05&#41;&#46; &#42;&#58; augment significatiu amb relaci&#243; a BN&#59; &#8224;&#58; difer&#232;ncia significativa en comparaci&#243; amb OB i CO &#40;p &#8804; 0&#44;05&#41;&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Discussi&#243; </span></p><p class="elsevierStylePara"> Ploug et al&#46;<span class="elsevierStyleSup">28</span> establiren que GLUT4 en fibres no estimulades &#40;per la insulina&#41; es distribu&#239;a al llarg de totes les fibres musculars&#44; i era present tant en la superf&#237;cie &#40;68&#37; del GLUT4 total&#41; com en el nucli &#40;32&#37; del GLUT4 total&#41; de les fibres&#46; Tamb&#233; trobaren que els nuclis eren despla&#231;ats i alineats amb els vasos sanguinis que cursen al llarg de la superf&#237;cie de la fibra&#44; que ocasionen una gran fracci&#243; de GLUT4 prop de la font de la glucosa evitant la necessitat de difusi&#243; a trav&#233;s de llargues dist&#224;ncies&#46; Est&#224; ben documentat que&#44; en estat normal&#44; el GLUT4 circula lentament entre la membrana plasm&#224;tica i un o m&#233;s compartiments intracel&#183;lulars&#44; amb la gran majoria de transportadors que resideixen en els compartiments vesiculars dins de l&#8217;interior de la c&#232;l&#183;lula<span class="elsevierStyleSup">28&#44;29</span>&#46;</p><p class="elsevierStylePara"> L&#8217;acumulaci&#243; d&#8217;insulina estimulada de la prote&#239;na GLUT4 en la superf&#237;cie de la c&#232;l&#183;lula pot ser causada per un increment de 10 a 20 vegades de la r&#224;tio d&#8217;exocitosi amb una petita disminuci&#243; &#40;2-3 vegades&#41; de la r&#224;tio d&#8217;endocitosi del GLUT4<span class="elsevierStyleSup">10&#44;45&#44;30</span>&#46;</p><p class="elsevierStylePara"> La insulina estimula el transport de glucosa a trav&#233;s de la translocaci&#243; de GLUT4 a partir de l&#8217;agrupaci&#243; d&#8217;emmagatzemament intracel&#183;lular cap a la membrana plasm&#224;tica&#46; La resist&#232;ncia a la insulina causada per la manca d&#8217;insulina o uns resultats anormals de senyalitzaci&#243; de la insulina amb un descens en l&#8217;expressi&#243; y translocaci&#243; de GLUT4 causen la hipergluc&#232;mia i la diabetis<span class="elsevierStyleSup">30-32</span>&#46; Tamb&#233; s&#8217;ha demostrat que la contracci&#243; muscular augmenta el contingut de GLUT4 de la membrana cel&#183;lular<span class="elsevierStyleSup">29&#44;30</span>&#46; Diversos estudis suggereixen que l&#8217;exercici f&#237;sic causa l&#8217;expressi&#243; de la prote&#239;na GLUT4 i la translocaci&#243; en membrana plasm&#224;tica mitjan&#231;ant un mecanisme diferent de senyalitzaci&#243; de la insulina<span class="elsevierStyleSup">5&#44;33</span>&#46; La majoria d&#8217;estudis sobre el m&#250;scul esquel&#232;tic dels ratolins indiquen que hi ha m&#233;s GLUT4 en les fibres de tipus i que en les del II<span class="elsevierStyleSup">10&#44;34</span>&#44; la qual cosa coincideix amb els resultats que vam trobar en el ratolins&#46; En aquest estudi nom&#233;s vam utilitzar el m&#250;scul soli&#44; generalment conegut com a m&#250;scul de fibres de tipus I&#46;</p><p class="elsevierStylePara"> El nostre estudi concorda amb altres<span class="elsevierStyleSup">35-39</span> que han indicat que l&#8217;exercici f&#237;sic incrementa l&#8217;expressi&#243; de la prote&#239;na GLUT4 en subjectes diab&#232;tics&#44; tal com succe&#237; en els nostres subjectes obesos&#46;</p><p class="elsevierStylePara"> Vam veure que l&#8217;expressi&#243; d&#8217;ARNm de GLUT4 s&#8217;incrementa amb 12 setmanes d&#8217;entrenament de resist&#232;ncia&#44; en ambd&#243;s m&#250;sculs&#44; en el soli &#40;m&#250;scul en qu&#232; predominen les fibres de contracci&#243; lenta&#41; i en el gastrocnemi &#40;en qu&#232; predominen les fibres de contracci&#243; r&#224;pida&#41; en ratolins obesos&#44; amb patr&#243; intens de depend&#232;ncia&#46; Aix&#242; significa que la intensitat de l&#8217;exercici pot afectar l&#8217;expressi&#243; del gen GLUT4 &#40;i&#44; probablement&#44; la prote&#239;na&#41; en resposta a l&#8217;entrenament&#44; la qual cosa pot provocar canvis metab&#242;lics significatius especialment en els casos obesos&#46; En conson&#224;ncia amb els nostres resultats&#44; s&#8217;ha demostrat que la prote&#239;na GLUT4 i el transport de glucosa s&#243;n immensament superiors en les fibres musculars oxidatives vermelles &#40;tipus I i IIa&#41; que en les fibres glucol&#237;tiques blanques &#40;tipus IIb&#41;<span class="elsevierStyleSup">10&#44;40</span>&#46; Aix&#242; podria ser un factor important en l&#8217;adaptaci&#243; a l&#8217;exercici&#44; degut a qu&#232; els resultats de l&#8217;entrenament de resist&#232;ncia faciliten un canvi de fibres de tipus iib a fibres tipus IIa<span class="elsevierStyleSup">41</span>&#46; Tanmateix&#44; en els m&#250;sculs esquel&#232;tics humans hi ha una difer&#232;ncia molt menor en l&#8217;expressi&#243; de GLUT4 en diferents tipus de fibres musculars<span class="elsevierStyleSup">34&#44;42</span>&#46; Daugaard et al&#46;<span class="elsevierStyleSup">34</span> van a&#239;llar fibres musculars individuals i les van classificar segons la isoforma de la miosina&#46; GLUT4 fou &#8764; 20&#37; m&#233;s elevada en les fibres que expressen la cadena pesada de la miosina i que en las que expressen la cadena pesada de la miosina IIa o IIb&#46; No es van trobar difer&#232;ncies entre les fibres IIa i IIb&#46; Despr&#233;s de 2 setmanes d&#8217;entrenament&#44; GLUT4 increment&#224; ~ 23&#37; de fibres musculars de tipus i&#44; per&#242; no hi hagu&#233; canvis en els tipus IIa o IIb&#46; Tanmateix&#44; l&#8217;exercici de baixa intensitat utilitzat &#233;s conegut principalment per reclutar fibres de tipus I&#46;</p><p class="elsevierStylePara"> D&#8217;acord amb els nostres resultats&#44; l&#8217;entrenament de baixa intensitat origin&#224; una expressi&#243; major d&#8217;ARNm de GLUT4 en el m&#250;scul oxidatiu &#40;&#233;s a dir&#44; el soli&#41;&#44; probablement degut a les necessitats relacionades amb la capacitat oxidativa de les c&#232;l&#183;lules&#46; D&#8217;altra banda&#44; la pr&#224;ctica d&#8217;exercici d&#8217;alta intensitat no produ&#237; una expressi&#243; major d&#8217;ARNm de GLUT4 en el soli ni en el gastrocnemi&#46; Com que el m&#250;scul gastrocnemi consisteix en una barreja de fibres vermelles i blanques&#44; una altra explicaci&#243; podria ser que l&#8217;entrenament amb exercicis de baixa intensitat no fou suficient per exercir o fins i tot per activar les fibres vermelles del m&#250;scul gastrocnemi&#44; i un augment no significatiu d&#8217;aquest m&#250;scul podria procedir de les fibres blanques&#46; Aquest escenari pot ampliar-se al que s&#8217;ha observat en les troballes relacionades amb l&#8217;alta intensitat&#59; &#233;s a dir&#44; un augment no significatiu d&#8217;aquest m&#250;scul amb l&#8217;exercici d&#8217;alta intensitat podria procedir nom&#233;s de les fibres vermelles&#46;</p><p class="elsevierStylePara"> Les hormones tamb&#233; regulen la concentraci&#243; de la prote&#239;na del m&#250;scul transportador de la glucosa&#46; L&#8217;expressi&#243; de GLUT4 s&#8217;increment&#224; degut a la insulina i a les hormones tiroidees<span class="elsevierStyleSup">20&#44;43</span> i disminu&#237; per un AMPc elevat<span class="elsevierStyleSup">44</span>&#46; La regulaci&#243; de l&#8217;expressi&#243; de GLUT4 per l&#8217;activitat contr&#224;ctil &#233;s independent de la regulaci&#243; hormonal&#44; perqu&#232; la carrera en la cinta de c&#243;rrer augment&#224; el GLUT4 en els ratolins diab&#232;tics<span class="elsevierStyleSup">45</span>&#46; S&#8217;hi afegiren els efectes de la defici&#232;ncia de la insulina i denervaci&#243; en les concentracions de GLUT4<span class="elsevierStyleSup">45</span>&#46;</p><p class="elsevierStylePara"> L&#8217;entrenament f&#237;sic tamb&#233; augmenta la prote&#239;na del GLUT4 i l&#8217;ARNm muscular en pacients amb diabetis tipus 2<span class="elsevierStyleSup">46</span>&#46; Aquests resultats suggereixen que l&#8217;activitat contr&#224;ctil del m&#250;scul modula directament l&#8217;expressi&#243; de GLUT4 del m&#250;scul&#44; independentment de l&#8217;acci&#243; de la insulina&#46; La histona deacetilasa 5 &#40;HDAC5&#41; &#233;s un mediador cr&#237;tic dels canvis del nivell d&#8217;ARNm de GLUT4 en resposta a l&#8217;entrenament<span class="elsevierStyleSup">47</span>&#46; S&#8217;ha demostrat que una s&#232;rie de modificacions de les histones causades per la histona deacetilasa i la histona metiltransferasa culminaran en una caiguda metab&#242;lica del gen GLUT4 en el m&#250;scul esquel&#232;tic dels ratolins<span class="elsevierStyleSup">48</span>&#46;</p><p class="elsevierStylePara"> Hi ha evid&#232;ncies significatives que els nivells d&#8217;ARNm de GLUT4 disminueixen en el m&#250;scul del teixit adip&#243;s amb l&#8217;obesitat<span class="elsevierStyleSup">49&#44;50</span>&#44; y que l&#8217;augment d&#8217;ARNm de GLUT4 pot alleujar o compensar la resist&#232;ncia a la insulina<span class="elsevierStyleSup">51&#44;52</span>&#46; Tal com era d&#8217;esperar&#44; les nostres troballes sobre els nivells d&#8217;ARNm del GLUT4 augmentaren significativament amb l&#8217;entrenament&#44; i aquest increment s&#8217;associ&#224; a una p&#232;rdua de pes&#44; la qual cosa recolza aquesta idea&#46; Per tant&#44; l&#8217;augment de la transcripci&#243; de GLUT4 en el m&#250;scul durant la p&#232;rdua de pes pot ser una prova definitiva en la reversi&#243; de la resist&#232;ncia a la insulina&#46;</p><p class="elsevierStylePara"> Estudis anteriors&#44; juntament amb els nostres resultats&#44; suggereixen que la regulaci&#243; transcripcional del gen GLUT4 &#233;s sensible a canvis en l&#8217;equilibri d&#8217;energia del cos&#46; A m&#233;s&#44; nosaltres vam trobar una resposta de GLUT4 a l&#8217;entrenament depenent de la intensitat i del tipus de fibra&#44; que no es plantejava en estudis anteriors&#46;</p><p class="elsevierStylePara"> Podem concloure que tot tipus d&#8217;alteraci&#243; de l&#8217;equilibri energ&#232;tic del cos&#44; especialment per la pr&#224;ctica d&#8217;exercici i&#47;o dieta alta en greixos&#44; pot influir en els mecanismes moleculars i cel&#183;lulars que actuen per establir una home&#242;stasi estable&#46; GLUT4 podria disminuir si s&#8217;augment&#233;s la ingesta d&#8217;energia&#44; i a la inversa&#44; augmentaria si s&#8217;augment&#233;s la despesa energ&#232;tica&#46; Aquests canvis s&#243;n gaireb&#233; similars en els m&#250;sculs de contracci&#243; lenta i r&#224;pida&#44; amb una mica m&#233;s de capacitat de resposta en els m&#250;sculs de contracci&#243; lenta&#44; perqu&#232; estan m&#233;s compromesos amb el metabolisme de la glucosa&#46;</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Conflicte d&#8217;interessos </span></p><p class="elsevierStylePara"> Els autors declaren que no tenen cap conflicte d&#8217;interessos</p><p class="elsevierStylePara"><span class="elsevierStyleBold">Agra&#239;ment </span></p><p class="elsevierStylePara"> Els autors agraeixen a la Universitat de Guilan el suport econ&#242;mic atorgat a aquest projecte d&#8217;investigaci&#243;&#46;</p><hr></hr><p class="elsevierStylePara"> Rebut el 10 d&#8217;octubre de 2015&#59; <br></br> acceptat el 14 de desembre de 2015</p><p class="elsevierStylePara"> &#42; Autor per a correspond&#232;ncia&#46;<br></br><span class="elsevierStyleItalic">Correu electr&#242;nic&#58;</span><a href="mailto&#58;mohebbi-h&#64;yahoo&#46;com" class="elsevierStyleCrossRefs">mohebbi-h&#64;yahoo&#46;com</a> &#40;H&#46; Mohebbi&#41;&#46;</p>"
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        "resumen" => "<p class="elsevierStylePara"> L&#8217;objectiu d&#8217;aquest estudi fou investigar la resposta a l&#8217;entrenament de resist&#232;ncia de l&#8217;expressi&#243; de l&#8217;ARNm del GLUT4 dels m&#250;sculs soli i gastrocnemi en ratolins obesos&#46; En aquest estudi es van utilitzar 40 ratolins mascles C57BL&#47;6&#46; Vuit ratolins &#40;base normal &#91;BN&#93;&#41; serviren com a controls no obesos i no entrenats&#44; i 32 ratolins foren sotmesos a una dieta alta en greixos &#40;DAG&#41; &#40;augment del 60&#37; de kcal&#41; durant 12 setmanes&#46; A la setmana 16 els ratolins obesos foren distribu&#239;ts aleat&#242;riament en els grups de tractament seg&#252;ents &#40;n &#61; 8 cada grup&#41;&#58; obesos base &#40;OB&#41;&#59; baixa intensitat &#40;BI&#41;&#59; alta intensitat &#40;AI&#41;&#44; o grups de control d&#8217;obesitat &#40;CO&#41;&#46; BI i AI foren entrenats 5 dies per setmana durant 12 setmanes en una cinta de c&#243;rrer motoritzada a 15 m&#47;min amb un pendent del 5&#37; &#40;en BI&#41; i&#47;o en 22 m&#47; min en un pendent del 5&#37; &#40;en AI&#41;&#46; Els ratolins CO es mantingueren sedentaris a la g&#224;bia durant el programa d&#8217;entrenament&#46; Es mesur&#224; l&#8217;expressi&#243; de l&#8217;ARNm del GLUT4 dels m&#250;sculs gastrocnemi i soli&#44; utilitzant el m&#232;tode PCR en temps real&#46; L&#8217;expressi&#243; de l&#8217;ARNm del GLUT4 del m&#250;scul soli en el grup BI augment&#224; aproximadament 2&#44;2 vegades&#44; enfront a 1&#44;6 vegades en el gastrocnemi &#40;p &#60; 0&#44;05&#41;&#46; A m&#233;s&#44; l&#8217;expressi&#243; de l&#8217;ARNm del GLUT4 en els m&#250;sculs soli i gastrocnemi dels grups BI i AI fou significativament major que en OB i en els grups CO &#40;p &#60; 0&#44;05&#41;&#46; Podem concloure que tota alteraci&#243; de l&#8217;equilibri energ&#232;tic del cos&#44; especialment per l&#8217;entrenament f&#237;sic i&#47;o una dieta alta en greixos&#44; pot influir en els mecanismes moleculars i cel&#183;lulars que actuen per establir una home&#242;stasi estable&#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;n &#61; 8 each group&#41;&#58; obese base &#91;OB&#93;&#59; low intensity &#91;LI&#93;&#59; high intensity &#91;HI&#93;&#44; 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&#44; can influence these molecular and cellular mechanisms that act to establish a stable homeostasis&#46;</p>"
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                            0 => "Ren JM"
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                            0 => "Hirshman MF"
                            1 => "Goodyear LJ"
                            2 => "Wardzala LJ"
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ISSN: X8866581
Original language: Català
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