Muscle fiber architecture, i.e., the physical arrangement of fibers within a muscle, is an important determinant of a muscle's mechanical function. In pennate muscles, fibers are oriented at an angle to the muscle's line of action and rotate as they shorten, becoming more oblique such that the fraction of force directed along the muscle's line of action decreases throughout a contraction. Fiber rotation decreases a muscle's output force but increases output velocity by allowing the muscle to function at a higher gear ratio (muscle velocity/fiber velocity). The magnitude of fiber rotation, and therefore gear ratio, depends on how the muscle changes shape in the dimensions orthogonal to the muscle's line of action. Here, we show that gear ratio is not fixed for a given muscle but decreases significantly with the force of contraction (P < 0.0001). We find that dynamic muscle-shape changes promote fiber rotation at low forces and resist fiber rotation at high forces. As a result, gearing varies automatically with the load, to favor velocity output during low-load contractions and force output for contractions against high loads. Therefore, muscle-shape changes act as an automatic transmission system allowing a pennate muscle to shift from a high gear during rapid contractions to low gear during forceful contractions. These results suggest that variable gearing in pennate muscles provides a mechanism to modulate muscle performance during mechanically diverse functions.biomechanics ͉ force-velocity tradeoff ͉ gear ratio ͉ muscle architecture T he force, speed, and power that can be harnessed from skeletal muscles to power movement are ultimately limited by the mechanical behavior of myofibers, the muscles' contractile cells. Two features of the contractile behavior of myofibers that likely constrain locomotor performance are the well defined limits to contraction velocity and contractile force. The maximum shortening velocity of a myofiber can be characterized by allowing the muscle to contract against near-zero loads, and a maximum isometric force can be defined when a muscle contracts at a fixed length. To understand how these limits to speed and force at the level of skeletal muscle cells translate to limits to speed or force of movement requires a consideration of musculoskeletal components that modulate force and velocity ''downstream'' of the force-producing cells. The most familiar of these are the skeletal lever systems, which define the mechanical advantage (gearing) through which muscle force is transmitted (1, 2). Like any lever or gear, skeletal gearing influences the ratio of muscle force or velocity to output force or velocity.A less obvious potential gearing mechanism resides within the architecture of the muscles themselves (3). It is generally recognized that skeletal muscle architecture influences the force and velocity of a muscle, but in most cases, the analysis of the effects of muscle architecture on force or velocity output has been limited to predictions based on static anatomy. Dynamic changes...