What Functions Do Reflexes Serve During Human Locomotion?

Studies on the reflex modulation of vertebrate locomotion have been conducted in many different laboratories and with many different preparations: for example, lamprey swimming, bird flight, quadrupedal walking in cats and bipedal walking in humans. Emerging concepts are that reflexes are task-, phase- and context-dependent. To function usefully in a behaviour such as locomotion wherein initial conditions change from step to step, reflexes would have to show modulation. Papers are reviewed in which the study of different reflexes have been conducted during different behaviours, with an emphasis on experiments in humans. A framework is developed in which the modulation and flexibility of reflexes are demonstrated. Alterations in cutaneous, and muscle (stretch and load receptor) reflexes between sitting, standing and walking are discussed. Studies in which both electrical, mechanical and 'natural' receptor activation have been conducted during walking are reviewed. Reflexes are shown to have important regulatory functions during human locomotion. A framework for discussion of reflex function throughout the step cycle is developed. The function of a given reflex pathway changes dynamically throughout the locomotor cycle. While all reflexes act in concert to a certain extent, generally cutaneous reflexes act to alter swing limb trajectory to avoid stumbling and falling. Stretch reflexes act to stabilize limb trajectory and assist force production during stance. Load receptor reflexes are shown to have an effect on both stance phase body weight support and step cycle timing. After neurotrauma or in disease, reflexes no longer function as during normal locomotion, but still have the potential to be clinically exploited in gait modification regimens.