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UMTS uses wideband code-division multiple access W-CDMA radio access technology to offer greater spectral efficiency and bandwidth to mobile network operators. These speeds are significantly faster than the 9. The first national consumer UMTS networks launched in with a heavy emphasis on telco-provided mobile applications such as mobile TV and video calling. While they suggest covering just a channel access method namely a variant of CDMA , they are actually the common names for the whole air interface standards. It supports conventional cellular voice, text and MMS services, but can also carry data at high speeds, allowing mobile operators to deliver higher bandwidth applications including streaming and broadband Internet access.
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The mitochondrial protein deacetylase sirtuin SIRT 3 may mediate exercise training-induced increases in mitochondrial biogenesis and improvements in reactive oxygen species ROS handling. Exercise training for 6. We propose an important role for AMPK in regulating mitochondrial function and ROS handling in skeletal muscle in response to exercise training. Mitochondrial density and capacity for oxidative ATP synthesis in skeletal muscle are tightly linked to cellular energetic demands Spina et al.
Protein deacetylases such as sirtuins SIRTs are important modulators of gene expression and protein activity and are involved in mitochondrial biogenesis.
SIRT3 modulates mitochondrial gene expression and function. SIRT3-driven processes occur via a general deacetylation of mitochondrial proteins Lombard et al.
AMPK functions as a major sensor of cellular energy status and can be activated pharmacologically and in response to muscle contraction and CR Koh et al. Exercise training increases ROS processing in skeletal muscle Parise et al. MnSOD and catalase govern the conversion of superoxide to water and oxygen in sequential steps Reid, Other regulatory acetylation sites have also been reported Qiu et al.
Whether this pathway is redundant or exclusively dependent on AMPK is unknown. Exercise training consisted of 15 sessions of one-legged knee extensor endurance training over the course of 3 weeks. The duration of the exercise sessions started at 1 h per session and was gradually increased to 2 h per session for the final 5 sessions.
Vastus lateralis muscle samples were obtained from healthy young men before and after an acute bout of one-legged knee-extensor exercise.
Selected data from this experiment have been published previously Treebak et al. Needle biopsies from the right and left vastus lateralis muscle were obtained before Pre and immediately after Post exercise cessation as described Treebak et al.
All animal experiments complied with the European Convention for the protection of Vertebrate Animals used for Experiments and Other Scientific Purposes Council of Europe , Strasbourg, France, and were approved by the Danish Animal Experimental Inspectorate and Selected data from these experiments have been published previously Brandauer et al.
The treadmill was horizontal the first week, and the incline was increased by 2. The morning following the final exercise bout, mice were anesthetized by an intraperitoneal i. Mice were anesthetized via i. In order to determine the acute effect of a single injection of AICAR, we analyzed samples originating from a previous study Leick et al.
Based on the recorded running distances of each mouse, mice were assigned to one of the three groups so that the average amount of training performed in the training period was similar between the three groups i. Mice then performed an acute treadmill exercise bout for 1 h or served as sedentary controls. Mice were euthanized by cervical dislocation immediately after the acute exercise bout. All muscle samples were processed using steel bead homogenization Tissue Lyser II, Qiagen in ice-cold lysis buffer pH 7.
Western blots were executed in a balanced design, with samples from all experimental conditions present on all gels, and identical internal control samples included on each blot. The internal control sample was prepared as a pool of all samples from a given experiment. All samples on individual gels were normalized to the internal control sample in order to permit comparison of samples resolved on separate gels.
David Gius. Complete alignment of the bands was confirmed Figure 7. The Tukey test was used post-hoc. If statistical interactions were borderline significant 0. As incomplete sample sets are excluded from statistical analyses in repeated-measures ANOVA, statistical analyses performed on data from the human training study were only performed on 6 sets of samples. We found no interaction effects in any of the data sets presented in Figures 1A—F , likely due to low statistical power.
Collectively, these data suggest a regulatory role for AMPK in mediating the effects of endurance exercise training on mitochondrial protein expression. Figure 1. Figure 2. Quadriceps muscles were obtained the day after the final bout of exercise. We employed a model of one-legged endurance exercise training to study SIRT3 and MnSOD protein abundance in trained and untrained human skeletal muscle.
Collectively, our data indicate that protein content of MnSOD is upregulated in response to exercise training in human skeletal muscle. Figure 3. One-legged knee-extensor endurance exercise training increases MnSOD protein levels in human vastus lateralis muscle.
Male individuals performed 15 sessions of 1—2 h of one-legged knee extensor endurance exercise training over a period of 3 weeks. We performed Western blot analyses for proteins involved in oxidative phosphorylation to determine the efficacy of the AICAR treatment.
However, no significant interaction effects were found for Complex I-V, most likely due to low statistical power. When individual t -tests were performed, significant increases in protein content were found in the WT group for Complexes I, IV, and V.
Figure 4. Figure 5. Figure 6. Quadriceps muscles were obtained the day after the last injection. Deacetylation of this residue would be expected to occur if superoxides are produced in mitochondria during exercise.
Figure 7. Unexpectedly, 60 min of high-intensity acute exercise caused further increases in SIRT3 protein abundance in both previously sedentary and trained mice Figure 8A.
This increase in protein content was not accompanied by a parallel increase in SIRT3 mRNA in these samples relative to samples from any of the other groups data not shown. No additional change occurred in response to acute exercise Figures 8B,C.
Figure 2C. Figure 8. Although the importance of mitochondria as a source of superoxides during muscle contraction has been questioned McArdle et al. We detected increased acetylation of MnSOD K in the samples from exercise-trained animals, whereas acetylation levels were unaffected by acute exercise Figure 8E.
We also tested the hypothesis that acetylation of OSCP on the K residue is a marker of cellular energy stress, and that exercised trained muscle would be less susceptible to exercise-induced changes in K acetylation.
To compare the MnSOD and OSCP acetylation response obtained in exercised mouse quadriceps muscle with acutely exercised human skeletal muscle, we determined the acetylation level of these proteins in samples obtained from strenuously exercised vastus lateralis muscle of healthy young men. Figure 9. No significant effects were observed. The present study confirms these findings and provides additional evidence that AMPK is required to fully beget exercise training-induced increases in mitochondrial oxidative phosphorylation complexes.
This observation is in line with recent data showing that exercise training-induced adaptations in skeletal muscle on mitochondrial Complexes I-V are dependent on the upstream AMPK kinase, LKB1 Tanner et al. However, these studies only assessed citrate synthase activity and did not report the abundance of electron transport chain proteins. The one-legged endurance exercise training model represents a well-controlled method to study contraction-mediated adaptations in vastus lateralis muscle in humans Andersen et al.
Despite a relatively small sample size, we found near-significant increases in skeletal muscle MnSOD protein level in the trained, but not untrained leg of healthy volunteers. Conversely, SIRT3 protein levels were not increased. This is in conflict with emerging evidence that SIRT3 expression is increased in exercise-trained human and rodent skeletal muscle Lanza et al.
While an early cross-sectional study reported higher protein activity of MnSOD in skeletal muscle of individuals with high aerobic fitness Jenkins et al. Apart from obvious species and protocol differences between the two studies, these findings could also suggest additional, possibly time-sensitive, regulatory mechanisms of SIRT3 protein levels.
In the present study, SIRT1 and catalase protein abundance was not consistently increased with exercise training, despite a substantive increase in skeletal muscle SIRT3 protein levels in WT mice. These findings corroborate earlier reports showing that SIRT1 expression is not strongly correlated with contraction-induced changes in mitochondrial protein abundance Chabi et al.
These data suggest that the differential response in mitochondrial protein abundance following exercise training is unrelated to SIRT1 activation via NAD. Our data on mitochondrial protein abundance should be interpreted in context with previously published literature on fiber type switching following exercise training or AICAR administration. However, the finding that AICAR treatment increases mitochondrial biogenesis, but may not necessarily change fiber type suggests that these processes may commonly occur in tandem but not be regulated by the same physiological stimuli.
SIRT3 is known to have deacetylase activity in the mitochondrion Lombard et al. MnSOD has multiple acetylation sites Rardin et al. Rather, when normalized to total protein levels, MnSOD K acetylation was unaltered between exercise trained and untrained mice, whereas acetylation status of K on OSCP was increased. However, our data on MnSOD and OSCP acetylation following acute exercise are not in obvious agreement with reported exercise-induced reductions in acetylation of these sites Vassilopoulos et al.
The lack of exercise-induced changes in MnSOD and OSCP acetylation was compared with human skeletal muscle samples obtained in conjunction with a one-legged exercise protocol. Further studies are warranted to determine the exact factors regulating SIRT3-regulated protein deacetylation with acute exercise.
The substantial increase in K acetylation we found after exercise training in mice may seem counterintuitive. One explanation for this observed increased OSCP acetylation could be that total cellular mitochondrial content is enhanced.
Another possibility is that lysine residues other than K on OSCP or lysines on other subunits of the ATP synthase are important for ATP synthase activity, and that acetylation status of these is changed in response to training. The fact that concentrations of the mitochondrial SIRT3 protein were more than 3-fold higher in exercise-trained compared with untrained mice presents an intriguing observation.
Why increased deacetylase protein concentration would result in unchanged or even increased acetylation levels in MnSOD or OSCP, respectively, is not immediately clear. In short, an acute increase in SIRT3 protein or activity is consistent with an inhibition of protein synthesis and other energetically costly processes by AMPK activation Jensen et al.
These proteins are important for the regulation of mitochondrial adaptation and the handling of ROS, respectively. The interactive effects of acute and chronic exercise on MnSOD and OSCP acetylation status constitute an unexplored avenue, with implications for intensity- and duration-dependent mitochondrial adaptations that warrant further investigation.
Edited, revised and approved the final version of the manuscript: All. The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
The authors wish to acknowledge the following people from Section of Molecular Physiology, Department of Nutrition, Exercise and Sports, University of Copenhagen for generous access to human skeletal muscle samples: Prof. Wojtaszewski, Prof. Erik A. Richter, Prof.
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We find that depenalization policy caused the police to reallocate effort toward nondrug crime. Despite the overall fall in crime attributable to the policy, we find that the total welfare of local residents likely fell, as measured by house prices. We shed light on what would be the impacts on crime of a citywide depenalization policy by developing and calibrating a structural model of the market for cannabis and crime. Introduction In nearly every country the market for illicit drugs remains pervasive, despite long-running attempts to restrict such activities. All rights reserved. We contribute to this policy debate by evaluating an increasingly common policy intervention in the illicit drug market: the depenalization of cannabis possession, so that the possession of small quantities of cannabis is no longer a criminally prosecutable offense. We present evidence from a localized UK policing experiment that introduced such a policy and focus attention on measuring its impact on crime, considered to be a major social cost of illicit drug markets.
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Semisynthetic biosensors for mapping cellular concentrations of nicotinamide adenine dinucleotides
Thank you for visiting nature. You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser or turn off compatibility mode in Internet Explorer. In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript. The mechanistic target of rapamycin complex 1 mTORC1 kinase regulates cell growth by setting the balance between anabolic and catabolic processes. To be active, mTORC1 requires the environmental presence of amino acids and glucose.
Dihydroxyacetone phosphate signals glucose availability to mTORC1
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