Muscle activation patterns and motor anatomy of Anna's hummingbirds Calypte anna and zebra finches Taeniopygia guttata.

Flying animals exhibit profound transformations in anatomy, physiology, and neural architecture. Although much is known about adaptations in the avian skeleton and musculature, less is known about neuroanatomy and motor unit integration for bird flight. Hummingbirds are among the most maneuverable and specialized of vertebrate fliers, and two unusual neuromuscular features have been previously reported: (1) the pectoralis major has a unique distribution pattern of motor end plates (MEPs) compared with all other birds and (2) electromyograms (EMGs) from the hummingbird's pectoral muscles, the pectoralis major and the supracoracoideus, show activation bursts composed of one or a few spikes that appear to have a very consistent pattern. Here, we place these findings in a broader context by comparing the MEPs, EMGs, and organization of the spinal motor neuron pools of flight muscles of Anna's hummingbird Calypte anna, zebra finches Taeniopygia guttata, and, for MEPs, several other species. The previously shown MEP pattern of the hummingbird pectoralis major is not shared with its closest taxonomic relative, the swift, and appears to be unique to hummingbirds. MEP arrangements in previously undocumented wing muscles show patterns that differ somewhat from other avian muscles. In the parallel-fibered strap muscles of the shoulder, MEP patterns appear to relate to muscle length, with the smallest muscles having fibers that span the entire muscle. MEP patterns in pennate distal wing muscles were the same regardless of size, with tightly clustered bands in the middle portion of the muscle, not evenly distributed bands over the muscle's entire length. Muscle activations were examined during slow forward flight in both species, during hovering in hummingbirds, and during slow ascents in zebra finches. The EMG bursts of a wing muscle, the pronator superficialis, were highly variable in peak number, size, and distribution across wingbeats for both species. In the pectoralis major, although the individual EMG bursts were much shorter in duration in hummingbirds relative to zebra finches, the variables describing the normalized amplitude and area of the activation bursts were otherwise indistinguishable between taxa during these flight modes. However, the degree of variation in the time intervals between EMG peaks was much lower in hummingbirds, which is a plausible explanation for the "patterned" EMG signals reported previously.

Sex differences in cannabinoid receptor-1 (CB1) pharmacology in mice selectively bred for high voluntary wheel-running behavior.

The endocannabinoid system (ECS) is involved in regulation of various physiological functions, including locomotion, antinociception, emotional states, and motivated behaviors. The ECS has been implicated in regulation of voluntary wheel running in mice via actions at the cannabinoid receptor-1 (CB1). Previously, we showed that four replicate lines of mice bred for high levels of voluntary wheel running (high-runner or HR lines) sex-specifically (females only) decreased running in response to antagonism of the CB1 receptor, as compared with four unselected Control lines. Here, we administered a CB1 receptor agonist, WIN 55,212-2 (WIN). We predicted that if CB1 activation is involved in the regulation of voluntary wheel running, then HR mice would show a greater response to CB1 agonism. Following our previous protocols, mice from generation 53 were acclimated to running wheels for 24 days, then received, in random order, either an intra-peritoneal injection of vehicle or a low (0.5 mg/kg), medium (1 mg/kg) or high dosage (3 mg/kg) of WIN. Each mouse received an injection and then experienced two nights without injections, for a total period of 12 days. Response to WIN was quantified as wheel revolutions, time spent running, and average running speed in the 10-120 min immediately following injection. Injection decreased wheel revolutions in all mice, but male HR mice decreased their running to a greater degree relative to Controls in response to the high dose of WIN over the entire period analyzed, whereas HR females showed a differential response relative to Controls only in the latter 70-120 min post-injection. These results, in conjunction with our previous study, show that (a) aspects of endocannabinoid signaling have diverged in four lines of mice bred for high levels of voluntary exercise and (b) male and female HR mice differ from one another in CB1 signaling as it relates to wheel running.

The biological control of voluntary exercise, spontaneous physical activity and daily energy expenditure in relation to obesity: human and rodent perspectives.

Mammals expend energy in many ways, including basic cellular maintenance and repair, digestion, thermoregulation, locomotion, growth and reproduction. These processes can vary tremendously among species and individuals, potentially leading to large variation in daily energy expenditure (DEE). Locomotor energy costs can be substantial for large-bodied species and those with high-activity lifestyles. For humans in industrialized societies, locomotion necessary for daily activities is often relatively low, so it has been presumed that activity energy expenditure and DEE are lower than in our ancestors. Whether this is true and has contributed to a rise in obesity is controversial. In humans, much attention has centered on spontaneous physical activity (SPA) or non-exercise activity thermogenesis (NEAT), the latter sometimes defined so broadly as to include all energy expended due to activity, exclusive of volitional exercise. Given that most people in Western societies engage in little voluntary exercise, increasing NEAT may be an effective way to maintain DEE and combat overweight and obesity. One way to promote NEAT is to decrease the amount of time spent on sedentary behaviours (e.g. watching television). The effects of voluntary exercise on other components of physical activity are highly variable in humans, partly as a function of age, and have rarely been studied in rodents. However, most rodent studies indicate that food consumption increases in the presence of wheels; therefore, other aspects of physical activity are not reduced enough to compensate for the energetic cost of wheel running. Most rodent studies also show negative effects of wheel access on body fat, especially in males. Sedentary behaviours per se have not been studied in rodents in relation to obesity. Several lines of evidence demonstrate the important role of dopamine, in addition to other neural signaling networks (e.g. the endocannabinoid system), in the control of voluntary exercise. A largely separate literature points to a key role for orexins in SPA and NEAT. Brain reward centers are involved in both types of physical activities and eating behaviours, likely leading to complex interactions. Moreover, voluntary exercise and, possibly, eating can be addictive. A growing body of research considers the relationships between personality traits and physical activity, appetite, obesity and other aspects of physical and mental health. Future studies should explore the neurobiology, endocrinology and genetics of physical activity and sedentary behaviour by examining key brain areas, neurotransmitters and hormones involved in motivation, reward and/or the regulation of energy balance.

How to run far: multiple solutions and sex-specific responses to selective breeding for high voluntary activity levels.

The response to uniform selection may occur in alternate ways that result in similar performance. We tested for multiple adaptive solutions during artificial selection for high voluntary wheel running in laboratory mice. At generation 43, the four replicate high runner (HR) lines averaged 2.85-fold more revolutions per day as compared with four non-selected control (C) lines, and females ran 1.11-fold more than males, with no sex-by-linetype interaction. Analysis of variance indicated significant differences among C lines but not among HR for revolutions per day. By contrast, average speed varied significantly among HR lines, but not among C, and showed a sex-by-linetype interaction, with the HR/C ratio being 2.02 for males and 2.45 for females. Time spent running varied among both HR and C lines, and showed a sex-by-linetype interaction, with the HR/C ratio being 1.52 for males but only 1.17 for females. Thus, females (speed) and males (speed, but also time) evolved differently, as did the replicate selected lines. Speed and time showed a trade-off among HR but not among C lines. These results demonstrate that uniform selection on a complex trait can cause consistent responses in the trait under direct selection while promoting divergence in the lower-level components of that trait.

Differential response to a selective cannabinoid receptor antagonist (SR141716: rimonabant) in female mice from lines selectively bred for high voluntary wheel-running behaviour.

Exercise is a naturally rewarding behaviour in human beings and can be associated with feelings of euphoria and analgesia. The endocannabinoid system may play a role in the perception of neurobiological rewards during and after prolonged exercise. Mice from lines that have been selectively bred for high voluntary wheel running (high runner or HR lines) may have evolved neurobiological mechanisms that increase the incentive salience of endurance-type exercise. Here, we test the hypothesis that endocannabinoid signalling has been altered in the four replicate HR lines as compared with four nonselected control lines. After 18 days of acclimation to cages with attached wheels, we injected mice with rimonabant (SR141716), a selective cannabinoid CB1 receptor antagonist. During the time of normal peak running, each mouse received, in a randomized order, intraperitoneal injection of rimonabant (0.1 or 3.0 mg/kg) or vehicle, over 9 days. Drug response was quantified as wheel revolutions, time and speed 10-70 min postinjection. Rimonabant decreased running in all mice; however, female HR mice differentially decreased running speed and distance (but not time) as compared with control females. We conclude that altered endocannabinoid signalling plays a role in the high wheel running of female HR mice.