Based Torin 1 biological activity molecular homogeneity, the diversity in geometry and adaptation of muscular motors leads to variations in tension equivalent to those resulting from the diversity of molecules and their arrangements in molecular motors. It is remarkable that so many different mechanisms lead to the same final distributions of force per cross-sectional area at the microscopic and macroscopic levels.4.4. Variability of tensions in whole musclesThe variability of tension in RWJ 64809 solubility muscles has been the subject of thorough research. An important adaptive factor is sarcomere length. As predicted by the sliding filament model of muscle contraction, long filaments and long overlap between thick and thin filaments should occur in fibres with long sarcomeres. As in long overlap zones more actin yosin cross-bridges should be formed, the maximum tension which a fibre can produce should be correlated with sarcomere length [207,208]. The resting sarcomere length exhibits little variation in insect and vertebrate muscles (2? ), but much greater variations in crustacean muscles (7?7 ). Overall, tension scales isometrically with the resting sarcomere length [157]. In particular, the claw closer muscles of cancer crabs exhibit both the longest sarcomere lengths and extreme mean crushing forces (525?030 kPa; table 4 and figure 3c). This is a special adaptation of shell-crushing non-locomotory motors which is not found in locomotors (figure 3d). Many other factors have been invoked to explain the variations in muscle tension, such as the density of the myosin filaments, the non-uniformity of sarcomere length along the fibres, the diameter of myofibrillar bundles, the actin : myosin filament ratios and the cross-bridge duty factors. For example, the slightly higher tension than in other groups found in amphibians and molluscs (except crustaceans; figure 3c) may be explained by their higher proportion of fast oxidative fibres and their higher relative myofibrillar volume [4,206]. However, these various factors apparently play a minor role in arthropod and vertebrate muscles as more than 80 of the variation in muscle tension in a series of muscles from these groups can be explained by the resting sarcomere length ([157] and references therein). Two characteristics other than tension contribute to muscle performance: speed of contraction (and relaxation) and endurance. They influence tension because high tension requires that most of the crosssectional area of a fibre be myofibrils, whereas high endurance requires a large mitochondrial volume and short twitch duration requires an extended sarcoplasmic reticulum. Therefore, trade-offs are inherent in the functional design of muscles so that a muscle cannot be simultaneously strong, enduring and rapid. This is the reason why rapid muscles are weak (either enduring, e.g. katytid singing muscles, or not, e.g. lobster sound-producing muscles with their hypertrophied SR) [208]. However, special adaptations in the oscillatory (asynchronous) flight muscles of insects result in high contraction frequencies without a large volume of SR, which leaves room for more mitochondria, but their strength is nevertheless limited by the endurance requirements of flight [208]. They are built optimally for maximum output of energy in their narrow contraction range, whereas most vertebrate sarcomeres are optimized for optimal mechanical conversion of chemical energy across a wider contraction range [209]. These different adaptions contribute to the.Based molecular homogeneity, the diversity in geometry and adaptation of muscular motors leads to variations in tension equivalent to those resulting from the diversity of molecules and their arrangements in molecular motors. It is remarkable that so many different mechanisms lead to the same final distributions of force per cross-sectional area at the microscopic and macroscopic levels.4.4. Variability of tensions in whole musclesThe variability of tension in muscles has been the subject of thorough research. An important adaptive factor is sarcomere length. As predicted by the sliding filament model of muscle contraction, long filaments and long overlap between thick and thin filaments should occur in fibres with long sarcomeres. As in long overlap zones more actin yosin cross-bridges should be formed, the maximum tension which a fibre can produce should be correlated with sarcomere length [207,208]. The resting sarcomere length exhibits little variation in insect and vertebrate muscles (2? ), but much greater variations in crustacean muscles (7?7 ). Overall, tension scales isometrically with the resting sarcomere length [157]. In particular, the claw closer muscles of cancer crabs exhibit both the longest sarcomere lengths and extreme mean crushing forces (525?030 kPa; table 4 and figure 3c). This is a special adaptation of shell-crushing non-locomotory motors which is not found in locomotors (figure 3d). Many other factors have been invoked to explain the variations in muscle tension, such as the density of the myosin filaments, the non-uniformity of sarcomere length along the fibres, the diameter of myofibrillar bundles, the actin : myosin filament ratios and the cross-bridge duty factors. For example, the slightly higher tension than in other groups found in amphibians and molluscs (except crustaceans; figure 3c) may be explained by their higher proportion of fast oxidative fibres and their higher relative myofibrillar volume [4,206]. However, these various factors apparently play a minor role in arthropod and vertebrate muscles as more than 80 of the variation in muscle tension in a series of muscles from these groups can be explained by the resting sarcomere length ([157] and references therein). Two characteristics other than tension contribute to muscle performance: speed of contraction (and relaxation) and endurance. They influence tension because high tension requires that most of the crosssectional area of a fibre be myofibrils, whereas high endurance requires a large mitochondrial volume and short twitch duration requires an extended sarcoplasmic reticulum. Therefore, trade-offs are inherent in the functional design of muscles so that a muscle cannot be simultaneously strong, enduring and rapid. This is the reason why rapid muscles are weak (either enduring, e.g. katytid singing muscles, or not, e.g. lobster sound-producing muscles with their hypertrophied SR) [208]. However, special adaptations in the oscillatory (asynchronous) flight muscles of insects result in high contraction frequencies without a large volume of SR, which leaves room for more mitochondria, but their strength is nevertheless limited by the endurance requirements of flight [208]. They are built optimally for maximum output of energy in their narrow contraction range, whereas most vertebrate sarcomeres are optimized for optimal mechanical conversion of chemical energy across a wider contraction range [209]. These different adaptions contribute to the.