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_:b419251025 , _:b419251026 , _:b419251027 , _:b419251036 , _:b419251037 , _:b419251038 , _:b419251039 , _:b419251032 , _:b419251033 , _:b419251034 , _:b419251035 , _:b419251012 , _:b419251013 , _:b419251014 , _:b419251015 , _:b419251008 , _:b419251009 , _:b419251010 , _:b419251011 , _:b419251020 , _:b419251021 , _:b419251022 , _:b419251023 , _:b419251016 , _:b419251017 , _:b419251018 , _:b419251019 , _:b419251060 , _:b419251061 , _:b419251062 , _:b419251063 , _:b419251056 , _:b419251057 , _:b419251058 , _:b419251059 , _:b419251068 , _:b419251069 , _:b419251070 , _:b419251071 , _:b419251064 , _:b419251065 , _:b419251066 , _:b419251067 , _:b419251044 , _:b419251045 , _:b419251046 , _:b419251047 , _:b419251040 , _:b419251041 , _:b419251042 , _:b419251043 , _:b419251052 , _:b419251053 , _:b419251054 , _:b419251055 , _:b419251048 , _:b419251049 , _:b419251050 , _:b419251051 , _:b419250964 , _:b419250965 , _:b419250966 , _:b419250967 , _:b419250960 , _:b419250961 , _:b419250962 , _:b419250963 , _:b419250972 , _:b419250973 , _:b419250974 , _:b419250975 , _:b419250968 , _:b419250969 , _:b419250970 , _:b419250971 , _:b419250948 , _:b419250949 , _:b419250950 , _:b419250951 , _:b419250944 , _:b419250945 , _:b419250946 , _:b419250947 , _:b419250956 , _:b419250957 , _:b419250958 , _:b419250959 , _:b419250952 , _:b419250953 , _:b419250954 , _:b419250955 , _:b419250996 , _:b419250997 , _:b419250998 , _:b419250999 , _:b419250992 , _:b419250993 , _:b419250994 , _:b419250995 , _:b419251004 , _:b419251005 , _:b419251006 , _:b419251007 , _:b419251000 , _:b419251001 , _:b419251002 , _:b419251003 , _:b419250980 , _:b419250981 , _:b419250982 , _:b419250983 , _:b419250976 , _:b419250977 , _:b419250978 , _:b419250979 , _:b419250988 , _:b419250989 , _:b419250990 , _:b419250991 , _:b419250984 , _:b419250985 , _:b419250986 , _:b419250987 , _:b419251216 , _:b419251217 , _:b419251204 , _:b419251205 , _:b419251206 , _:b419251207 , _:b419251200 , _:b419251201 , _:b419251202 , _:b419251203 , _:b419251212 , _:b419251213 , _:b419251214 , _:b419251215 , _:b419251208 , _:b419251209 , _:b419251210 , _:b419251211 ; ns1:pmcid "PMC0" ; bibo:doi "10.2337%2Fdb09-0482" . @prefix ns4: . ns4:contains _:b6461021 , _:b6461047 , _:b6461073 . _:b6461021 rdf:type ns4:Section . @prefix dc: . _:b6461021 dc:title "results" ; ns4:contains _:b6461022 , _:b6461023 , _:b6461040 , _:b6461041 , _:b6461042 , _:b6461043 , _:b6461044 , _:b6461045 , _:b6461046 , _:b6461024 , _:b6461025 , _:b6461026 , _:b6461027 , _:b6461028 , _:b6461029 , _:b6461030 , _:b6461031 , _:b6461032 , _:b6461033 , _:b6461034 , _:b6461035 , _:b6461036 , _:b6461037 , _:b6461038 , _:b6461039 . _:b6461022 rdf:type ns1:Context ; rdf:value "As reported previously (>>7<<), resveratrol increased physical endurance (309.2 \u00B1 37 vs. 212.3 \u00B1 19 m) (Fig. 3A) of wild-type mice." ; ns1:mentions . _:b6461023 rdf:type ns1:Context ; rdf:value "Prolonged activation of AMPK increases Glut4 expression (>>33<<). To determine whether increased expression of Glut4 expression was involved in the resveratrol-mediated improvement of physical endurance, we measured Glut4 mRNA levels in skeletal muscle." ; ns1:mentions . _:b6461024 rdf:type ns1:Context ; rdf:value "It is believed that the resveratrol effects are mediated by increased expression and function of PGC-1\u03B1, the master regulator of mitochondrial biogenesis and oxidative phosphorylation (>>34<<). We measured the expression of PGC-1\u03B1 and other resveratrol-induced genes critical for mitochondrial function in the skeletal muscle of resveratrol-treated AMPK-deficient mice. As shown in Fig. 3C, resveratrol induced the expression of" ; ns1:mentions . _:b6461025 rdf:type ns1:Context ; rdf:value "To determine whether the requirement of AMPK for the resveratrol effect is cell autonomous, we treated mefs developed from AMPK\u03B11/\u03B12 double knockout (>>32<<) and wild-type embryos with resveratrol for 6\u201324 h." ; ns1:mentions . _:b6461026 rdf:type ns1:Context ; rdf:value "Sirt1 overexpression (>>35<<\u201337), AMPK activation (21,22,38), and resveratrol treatment (5,7) all improve glucose tolerance and insulin sensitivity." ; ns1:mentions , , . _:b6461027 rdf:type ns1:Context ; rdf:value "Sirt1 overexpression (35\u201337), AMPK activation (>>21<<,22,38), and resveratrol treatment (5,7) all improve glucose tolerance and insulin sensitivity." ; ns1:mentions . _:b6461028 rdf:type ns1:Context ; rdf:value "Sirt1 overexpression (35\u201337), AMPK activation (21,>>22<<,38), and resveratrol treatment (5,7) all improve glucose tolerance and insulin sensitivity." ; ns1:mentions . _:b6461029 rdf:type ns1:Context ; rdf:value "Sirt1 overexpression (35\u201337), AMPK activation (21,22,>>38<<), and resveratrol treatment (5,7) all improve glucose tolerance and insulin sensitivity." ; ns1:mentions . _:b6461030 rdf:type ns1:Context ; rdf:value "Sirt1 overexpression (35\u201337), AMPK activation (21,22,38), and resveratrol treatment (>>5<<,7) all improve glucose tolerance and insulin sensitivity." ; ns1:mentions . _:b6461031 rdf:type ns1:Context ; rdf:value "Sirt1 overexpression (35\u201337), AMPK activation (21,22,38), and resveratrol treatment (5,>>7<<) all improve glucose tolerance and insulin sensitivity." ; ns1:mentions . _:b6461032 rdf:type ns1:Context ; rdf:value "C: Relative levels of ROS in skeletal muscle extracts from wild-type (WT) and AMPK\u03B11\u2212/\u2212 mice fed high-fat diet (\u25A1) alone or high-fat diet supplemented with resveratrol (\u25A0) as measured by DCF fluorescence (>>52<<) (n = 4\u20135 for each genotype)." ; ns1:mentions . _:b6461033 rdf:type ns1:Context ; rdf:value "Insulin-resistant states are associated with increased production of ROS, and reduction of ROS with antioxidants improves insulin sensitivity (>>39<<,40). Since resveratrol is thought to have antioxidant properties, it is possible that some insulin sensitizing of resveratrol effect may be related to its antioxidant effect." ; ns1:mentions . _:b6461034 rdf:type ns1:Context ; rdf:value "Insulin-resistant states are associated with increased production of ROS, and reduction of ROS with antioxidants improves insulin sensitivity (39,>>40<<). Since resveratrol is thought to have antioxidant properties, it is possible that some insulin sensitizing of resveratrol effect may be related to its antioxidant effect." ; ns1:mentions . _:b6461035 rdf:type ns1:Context ; rdf:value "Insulin-resistant tissues are often associated with elevated levels of intramyocellular lipids such as diacylglyceride (DAG) and ceramide (>>41<<). Mitochondrial dysfunction decreases fat oxidation, leading to accumulation of these lipids, which decrease insulin sensitivity (41,42)." ; ns1:mentions . _:b6461036 rdf:type ns1:Context ; rdf:value "Mitochondrial dysfunction decreases fat oxidation, leading to accumulation of these lipids, which decrease insulin sensitivity (>>41<<,42). Since resveratrol increases mitochondrial function (7), we measured the DAG and ceramide levels in resveratrol-treated skeletal muscle. As shown in Fig. 4D and E, resveratrol decreased DAG and ceramide levels in wild-type mice but" ; ns1:mentions . _:b6461037 rdf:type ns1:Context ; rdf:value "Mitochondrial dysfunction decreases fat oxidation, leading to accumulation of these lipids, which decrease insulin sensitivity (41,>>42<<). Since resveratrol increases mitochondrial function (7), we measured the DAG and ceramide levels in resveratrol-treated skeletal muscle. As shown in Fig. 4D and E, resveratrol decreased DAG and ceramide levels in wild-type mice but not" ; ns1:mentions . _:b6461038 rdf:type ns1:Context ; rdf:value "Since resveratrol increases mitochondrial function (>>7<<), we measured the DAG and ceramide levels in resveratrol-treated skeletal muscle." ; ns1:mentions . _:b6461039 rdf:type ns1:Context ; rdf:value "Adiponectin, a cytokine that is produced by fat and is decreased in obese and diabetic subjects (>>43<<), increases insulin sensitivity by activating AMPK (38)." ; ns1:mentions . _:b6461040 rdf:type ns1:Context ; rdf:value "Adiponectin, a cytokine that is produced by fat and is decreased in obese and diabetic subjects (43), increases insulin sensitivity by activating AMPK (>>38<<). To determine whether resveratrol increased insulin sensitivity by increasing adiponectin levels, we measured serum adiponectin levels in resveratrol-treated mice. Adiponectin levels were similar for wild-type, AMPK\u03B11\u2212/\u2212, and AMPK\u03B12\u2212/\u2212" ; ns1:mentions . _:b6461041 rdf:type ns1:Context ; rdf:value "Although this can be explained, at least in part, by the observation that resveratrol inhibits insulin secretion by inhibiting the electrical activity in the \u03B2-cells (>>44<<), more studies are required to fully understand the disparity between the insulin levels and insulin sensitivity in resveratrol-treated AMPK-null mice." ; ns1:mentions . _:b6461042 rdf:type ns1:Context ; rdf:value "A previous report (>>7<<) has shown that resveratrol treatment leads to deacetylation of PGC-1\u03B1, suggesting that the catalytic activity of Sirt1 is increased by resveratrol." ; ns1:mentions . _:b6461043 rdf:type ns1:Context ; rdf:value "Since AMPK activity increases the NAD-to-NADH ratio and Sirt1 activity (>>45<<), we measured the NAD-to-NADH ratio in C2C12 myotubes treated with resveratrol for 6 and 16 h." ; ns1:mentions . _:b6461044 rdf:type ns1:Context ; rdf:value "PGC-1\u03B1 is a coactivator for its own transcription (>>46<<), and, as a result, deacetylation of PGC-1\u03B1 by Sirt1 should increase its ability to positively regulate PGC-1\u03B1 transcription." ; ns1:mentions . _:b6461045 rdf:type ns1:Context ; rdf:value "A previous report (>>7<<) showed that resveratrol did not induce PGC-1\u03B1 transcription in Sirt1\u2212/\u2212 mefs and concluded that Sirt1 is essential for resveratrol-induced transcription of PGC-1\u03B1." ; ns1:mentions . _:b6461046 rdf:type ns1:Context ; rdf:value "Consistent with previous reports (>>47<<), \u2212Sirt1 mefs had higher AMPK activity than +Sirt1 mefs (Fig." ; ns1:mentions . _:b6461047 rdf:type ns4:Section ; dc:title "discussion" ; ns4:contains _:b6461048 , _:b6461049 , _:b6461050 , _:b6461051 , _:b6461052 , _:b6461053 , _:b6461054 , _:b6461055 , _:b6461072 , _:b6461064 , _:b6461065 , _:b6461066 , _:b6461067 , _:b6461068 , _:b6461069 , _:b6461070 , _:b6461071 , _:b6461056 , _:b6461057 , _:b6461058 , _:b6461059 , _:b6461060 , _:b6461061 , _:b6461062 , _:b6461063 . _:b6461048 rdf:type ns1:Context ; rdf:value "One target for treating metabolic disorders is Sirt1 (>>8<<), an NAD+-dependent protein deacetylase (48)." ; ns1:mentions . _:b6461049 rdf:type ns1:Context ; rdf:value "One target for treating metabolic disorders is Sirt1 (8), an NAD+-dependent protein deacetylase (>>48<<). Resveratrol, which was discovered in a small-molecule screen to be a Sirt1 activator (1), has drawn a great deal of interest for its therapeutic potential in treating metabolic disorders such as type 2 diabetes. By increasing" ; ns1:mentions . _:b6461050 rdf:type ns1:Context ; rdf:value "Resveratrol, which was discovered in a small-molecule screen to be a Sirt1 activator (>>1<<), has drawn a great deal of interest for its therapeutic potential in treating metabolic disorders such as type 2 diabetes." ; ns1:mentions . _:b6461051 rdf:type ns1:Context ; rdf:value "In Sirt1 knockin mice (>>36<<), the Sirt1 transgene, which was expressed from the \u03B2-actin locus, is overexpressed in fat but not in muscle or liver, key organs affected in metabolic disorders." ; ns1:mentions . _:b6461052 rdf:type ns1:Context ; rdf:value "More recently, transgenic mice in which the Sirt1 transgene is expressed from its own promoter have been reported by two independent groups (>>35<<,37). These transgenic mice, which have a more physiological expression pattern of the Sirt1 transgene than the knockin mice (36), are more insulin sensitive partly due to increased adiponectin levels and decreased hepatic steatosis." ; ns1:mentions . _:b6461053 rdf:type ns1:Context ; rdf:value "More recently, transgenic mice in which the Sirt1 transgene is expressed from its own promoter have been reported by two independent groups (35,>>37<<). These transgenic mice, which have a more physiological expression pattern of the Sirt1 transgene than the knockin mice (36), are more insulin sensitive partly due to increased adiponectin levels and decreased hepatic steatosis." ; ns1:mentions . _:b6461054 rdf:type ns1:Context ; rdf:value "These transgenic mice, which have a more physiological expression pattern of the Sirt1 transgene than the knockin mice (>>36<<), are more insulin sensitive partly due to increased adiponectin levels and decreased hepatic steatosis." ; ns1:mentions . _:b6461055 rdf:type ns1:Context ; rdf:value "Surprisingly, the transgenic mice developed by Banks et al. (>>35<<), which are congenic in C57BL6/J background, have reduced metabolic rate and body temperature and have the same amount of fat despite eating less." ; ns1:mentions . _:b6461056 rdf:type ns1:Context ; rdf:value "Nevertheless, this study shows that the function of Sirt1 on energy balance may be opposite of what was previously thought (>>7<<) and would be predicted if the central target of resveratrol was Sirt1." ; ns1:mentions . _:b6461057 rdf:type ns1:Context ; rdf:value "One possible explanation for the reduced fat mass in the knockin mice (>>36<<) is that expression of the Sirt1 transgene from the \u03B2-actin locus impaired adipocyte differentiation during development (49)." ; ns1:mentions . _:b6461058 rdf:type ns1:Context ; rdf:value "One possible explanation for the reduced fat mass in the knockin mice (36) is that expression of the Sirt1 transgene from the \u03B2-actin locus impaired adipocyte differentiation during development (>>49<<). Therefore, the Sirt1 knockin model (36) cannot distinguish between a direct effect of Sirt1 and an indirect one caused by reduced fat." ; ns1:mentions . _:b6461059 rdf:type ns1:Context ; rdf:value "Therefore, the Sirt1 knockin model (>>36<<) cannot distinguish between a direct effect of Sirt1 and an indirect one caused by reduced fat." ; ns1:mentions . _:b6461060 rdf:type ns1:Context ; rdf:value "Resveratrol has been reported to affect the activities of many enzymes (>>11<<) including AMPK (12). To evaluate the possibility that the effects of resveratrol are mediated by AMPK, we studied the effects of resveratrol in mice deficient in AMPK\u03B11 or -\u03B12." ; ns1:mentions . _:b6461061 rdf:type ns1:Context ; rdf:value "Resveratrol has been reported to affect the activities of many enzymes (11) including AMPK (>>12<<). To evaluate the possibility that the effects of resveratrol are mediated by AMPK, we studied the effects of resveratrol in mice deficient in AMPK\u03B11 or -\u03B12." ; ns1:mentions . _:b6461062 rdf:type ns1:Context ; rdf:value "The ability of AMPK to increase NAD and the NAD-to-NADH ratio (>>45<<) may also explain how resveratrol treatment can lead to Sirt1 activation without directly activating it." ; ns1:mentions . _:b6461063 rdf:type ns1:Context ; rdf:value "Moreover, AMPK can activate PGC-1\u03B1, one of Sirt1 substrates, by directly phosphorylating it (>>50<<,51), indicating that activation of AMPK can affect the Sirt1-dependent pathways in multiple ways." ; ns1:mentions . _:b6461064 rdf:type ns1:Context ; rdf:value "Moreover, AMPK can activate PGC-1\u03B1, one of Sirt1 substrates, by directly phosphorylating it (50,>>51<<), indicating that activation of AMPK can affect the Sirt1-dependent pathways in multiple ways." ; ns1:mentions . _:b6461065 rdf:type ns1:Context ; rdf:value "Whether it is the direct target of resveratrol or not, it appears that Sirt1 can also function upstream of AMPK in HepG2 hepatocytes and HEK293T cell line (>>26<<,27). Since Sirt1 is not required for resveratrol-mediated activation of AMPK activation in mefs (Fig. 6A), it is possible that Sirt1 is upstream of AMPK only in certain cell types." ; ns1:mentions . _:b6461066 rdf:type ns1:Context ; rdf:value "Whether it is the direct target of resveratrol or not, it appears that Sirt1 can also function upstream of AMPK in HepG2 hepatocytes and HEK293T cell line (26,>>27<<). Since Sirt1 is not required for resveratrol-mediated activation of AMPK activation in mefs (Fig. 6A), it is possible that Sirt1 is upstream of AMPK only in certain cell types." ; ns1:mentions . _:b6461067 rdf:type ns1:Context ; rdf:value "Resveratrol-induced physical endurance has been largely attributed to increased mitochondrial function (>>7<<). Certainly, converting muscle fiber to mitochondria-rich, slow-twitch fibers increases physical endurance (52)." ; ns1:mentions . _:b6461068 rdf:type ns1:Context ; rdf:value "Certainly, converting muscle fiber to mitochondria-rich, slow-twitch fibers increases physical endurance (>>52<<). However, it is also likely that the glycogen content in skeletal muscle, which is known to be increased by AMPK activity (34), also plays a role in resveratrol-induced physical endurance." ; ns1:mentions . _:b6461069 rdf:type ns1:Context ; rdf:value "However, it is also likely that the glycogen content in skeletal muscle, which is known to be increased by AMPK activity (>>34<<), also plays a role in resveratrol-induced physical endurance." ; ns1:mentions . _:b6461070 rdf:type ns1:Context ; rdf:value "We do not have a clear explanation for this, but one possibility is that the antiadipogenic function of Sirt1 (>>49<<) is being driven by the resveratrol-AMPK\u03B11-Sirt1 pathway in AMPK\u03B12\u2212/\u2212 mice." ; ns1:mentions . _:b6461071 rdf:type ns1:Context ; rdf:value "In skeletal muscle, the AMPK\u03B11 isoform makes up only \u223C20% of the total AMPK activity (>>17<<), and, yet, resveratrol-induced mitochondrial biogenesis (Fig." ; ns1:mentions . _:b6461072 rdf:type ns1:Context ; rdf:value "One possible explanation is that in addition to the AMPK\u03B12 activity, a crosstalk between the skeletal muscle and either the nervous system or fat, where the AMPK\u03B11 isoform is predominant (>>17<<), is required for resveratrol-induced mitochondrial biogenesis or reduction of ROS in skeletal muscle." ; ns1:mentions . _:b6461073 rdf:type ns4:Section ; dc:title "research design and methods" ; ns4:contains _:b6461074 , _:b6461075 , _:b6461076 , _:b6461077 , _:b6461078 , _:b6461079 , _:b6461080 . _:b6461074 rdf:type ns1:Context ; rdf:value "AMPK\u03B12\u2212/\u2212 (>>22<<) and AMPK\u03B11\u2212/\u2212 (28) mice were backcrossed to C57BL/6J for at least six generations before this study." ; ns1:mentions . _:b6461075 rdf:type ns1:Context ; rdf:value "AMPK\u03B12\u2212/\u2212 (22) and AMPK\u03B11\u2212/\u2212 (>>28<<) mice were backcrossed to C57BL/6J for at least six generations before this study." ; ns1:mentions . _:b6461076 rdf:type ns1:Context ; rdf:value "and fed a high-fat diet (40% calories from fat; Bio-serv) or a high-fat diet supplemented with resveratrol (400 mg \u00B7 kg\u22121 \u00B7 day\u22121; Orchid Chemicals and Pharmaceuticals) for 12 weeks as previously described (>>7<<). All experiments were approved by the National Heart, Lung, and Blood Institute Animal Care and Use Committee." ; ns1:mentions . _:b6461077 rdf:type ns1:Context ; rdf:value "Reactive oxygen species (ROS) levels were determined in muscle extracts using the ROS-sensitive fluorescent dye dichlorodihydrofluorescein (DCF), as previously described (>>29<<,30). Briefly, oxidation-insensitive dye (carboxy-DCFDA) was used as a control to ensure that changes in the fluorescence seen with the oxidation-sensitive dye (H2DCFDA) were due to changes in ROS production." ; ns1:mentions . _:b6461078 rdf:type ns1:Context ; rdf:value "Reactive oxygen species (ROS) levels were determined in muscle extracts using the ROS-sensitive fluorescent dye dichlorodihydrofluorescein (DCF), as previously described (29,>>30<<). Briefly, oxidation-insensitive dye (carboxy-DCFDA) was used as a control to ensure that changes in the fluorescence seen with the oxidation-sensitive dye (H2DCFDA) were due to changes in ROS production." ; ns1:mentions . _:b6461079 rdf:type ns1:Context ; rdf:value "Sirt1\u2212/\u2212 mouse embryonic fibroblasts (mefs) (>>31<<) were maintained in DMEM supplemented with 10% fetal bovine serum." ; ns1:mentions . _:b6461080 rdf:type ns1:Context ; rdf:value "AMPK\u03B11/\u03B12 double knockout and wild-type mefs were derived as previously described (>>32<<). Cells were treated with 50 \u03BCmol/l resveratrol (Sigma) or DMSO for the indicated time. C2C12 myoblast cells (ATCC) were maintained in DMEM and 10% FBS. 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