Retinal optic flow during natural locomotion.

We examine the structure of the visual motion projected on the retina during natural locomotion in real world environments. Bipedal gait generates a complex, rhythmic pattern of head translation and rotation in space, so without gaze stabilization mechanisms such as the vestibular-ocular-reflex (VOR) a walker's visually specified heading would vary dramatically throughout the gait cycle. The act of fixation on stable points in the environment nulls image motion at the fovea, resulting in stable patterns of outflow on the retinae centered on the point of fixation. These outflowing patterns retain a higher order structure that is informative about the stabilized trajectory of the eye through space. We measure this structure by applying the curl and divergence operations on the retinal flow velocity vector fields and found features that may be valuable for the control of locomotion. In particular, the sign and magnitude of foveal curl in retinal flow specifies the body's trajectory relative to the gaze point, while the point of maximum divergence in the retinal flow field specifies the walker's instantaneous overground velocity/momentum vector in retinotopic coordinates. Assuming that walkers can determine the body position relative to gaze direction, these time-varying retinotopic cues for the body's momentum could provide a visual control signal for locomotion over complex terrain. In contrast, the temporal variation of the eye-movement-free, head-centered flow fields is large enough to be problematic for use in steering towards a goal. Consideration of optic flow in the context of real-world locomotion therefore suggests a re-evaluation of the role of optic flow in the control of action during natural behavior.

Control of gaze in natural environments: effects of rewards and costs, uncertainty and memory in target selection.

The development of better eye and body tracking systems, and more flexible virtual environments have allowed more systematic exploration of natural vision and contributed a number of insights. In natural visually guided behaviour, humans make continuous sequences of sensory-motor decisions to satisfy current goals, and the role of vision is to provide the relevant information in order to achieve those goals. This paper reviews the factors that control gaze in natural visually guided actions such as locomotion, including the rewards and costs associated with the immediate behavioural goals, uncertainty about the state of the world and prior knowledge of the environment. These general features of human gaze control may inform the development of artificial systems.

Gaze and the Control of Foot Placement When Walking in Natural Terrain.

Human locomotion through natural environments requires precise coordination between the biomechanics of the bipedal gait cycle and the eye movements that gather the information needed to guide foot placement. However, little is known about how the visual and locomotor systems work together to support movement through the world. We developed a system to simultaneously record gaze and full-body kinematics during locomotion over different outdoor terrains. We found that not only do walkers tune their gaze behavior to the specific information needed to traverse paths of varying complexity but that they do so while maintaining a constant temporal look-ahead window across all terrains. This strategy allows walkers to use gaze to tailor their energetically optimal preferred gait cycle to the upcoming path in order to balance between the drive to move efficiently and the need to place the feet in stable locations. Eye movements and locomotion are intimately linked in a way that reflects the integration of energetic costs, environmental uncertainty, and momentary informational demands of the locomotor task. Thus, the relationship between gaze and gait reveals the structure of the sensorimotor decisions that support successful performance in the face of the varying demands of the natural world. VIDEO ABSTRACT.

The critical phase for visual control of human walking over complex terrain.

To walk efficiently over complex terrain, humans must use vision to tailor their gait to the upcoming ground surface without interfering with the exploitation of passive mechanical forces. We propose that walkers use visual information to initialize the mechanical state of the body before the beginning of each step so the resulting ballistic trajectory of the walker's center-of-mass will facilitate stepping on target footholds. Using a precision stepping task and synchronizing target visibility to the gait cycle, we empirically validated two predictions derived from this strategy: (1) Walkers must have information about upcoming footholds during the second half of the preceding step, and (2) foot placement is guided by information about the position of the target foothold relative to the preceding base of support. We conclude that active and passive modes of control work synergistically to allow walkers to negotiate complex terrain with efficiency, stability, and precision.

The biomechanics of walking shape the use of visual information during locomotion over complex terrain.

The aim of this study was to examine how visual information is used to control stepping during locomotion over terrain that demands precision in the placement of the feet. More specifically, we sought to determine the point in the gait cycle at which visual information about a target is no longer needed to guide accurate foot placement. Subjects walked along a path while stepping as accurately as possible on a series of small, irregularly spaced target footholds. In various conditions, each of the targets became invisible either during the step to the target or during the step to the previous target. We found that making targets invisible after toe off of the step to the target had little to no effect on stepping accuracy. However, when targets disappeared during the step to the previous target, foot placement became less accurate and more variable. The findings suggest that visual information about a target is used prior to initiation of the step to that target but is not needed to continuously guide the foot throughout the swing phase. We propose that this style of control is rooted in the biomechanics of walking, which facilitates an energetically efficient strategy in which visual information is primarily used to initialize the mechanical state of the body leading into a ballistic movement toward the target foothold. Taken together with previous studies, the findings suggest the availability of visual information about the terrain near a particular step is most essential during the latter half of the preceding step, which constitutes a critical control phase in the bipedal gait cycle.

Humans exploit the biomechanics of bipedal gait during visually guided walking over complex terrain.

How do humans achieve such remarkable energetic efficiency when walking over complex terrain such as a rocky trail? Recent research in biomechanics suggests that the efficiency of human walking over flat, obstacle-free terrain derives from the ability to exploit the physical dynamics of our bodies. In this study, we investigated whether this principle also applies to visually guided walking over complex terrain. We found that when humans can see the immediate foreground as little as two step lengths ahead, they are able to choose footholds that allow them to exploit their biomechanical structure as efficiently as they can with unlimited visual information. We conclude that when humans walk over complex terrain, they use visual information from two step lengths ahead to choose footholds that allow them to approximate the energetic efficiency of walking in flat, obstacle-free environments.