Abstract
Typically, locomotion has been studied by restricting the animal’s path and/or speed, focusing on stride and step kinematics. Here we incorporate measurements of the legs and trunk in the support and swing phases, during trotting with various speeds and curvatures. This paradigm releases the animal from the confines of the treadmill and runway into the open space. The diagonal step, a new unit of locomotion, is defined by regarding the line between the two supporting diagonal legs as a frame of reference for the description of the dynamics of the virtual line between the two swinging diagonal legs. This analysis reveals that during free trotting the mouse uses three types of steps: fixating, opening, and closing steps. During progression along a straight path, the mouse uses fixating steps, in which the swinging diagonal maintains a fixed direction, landing on the supporting foreleg; during progression along a curved path the mouse uses opening and closing steps alternately. If two steps of the same type are performed sequentially, they engender an abrupt change of direction. Our results reveal how steering with the swinging diagonal, while using a virtually bipedal gait, engenders the whole repertoire of free-trotting behavior.
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Acknowledgments
We thank Prof. Mark Shik for his encouragement and critical comments. All animals were maintained in facilities fully accredited by NIH Animal Welfare Assurance Number A5010-01 (TAU). The studies were conducted in accordance with the Guide for Care and Use of Laboratory Animals provided by the NIH ‘Principles of Laboratory Animal Care’ (NIH publication # 86-23, 1996).
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Trot recognition algorithm
The first stage of the algorithm was to identify the support phase of diagonal steps. This was done by calculating the speed of the mid-point of each diagonal in each frame and by recognizing, within each diagonal, segments of complete arrest. In the second stage, the separate sequences of support-swing of the two diagonals were interwoven into a single sequence marking the supporting diagonal for each segment. To sift out frames in which all 4 legs were in contact with the ground, segments in which both diagonals were at arrest were marked as irrelevant; to treat instances in which a leg belonging to one diagonal started to step before the other diagonal stopped, segments in which both diagonals moved were added to the preceding support segment. The support-swing segments obtained at this stage became the candidates for genuine diagonal steps.
The third stage was one of inspection and correction. Step candidates in which 3 or more feet were in contact with the ground for more than 3 frames were marked as irrelevant and discarded. While the legs in the trot gait are expected to release and establish contact 'at the same instant', the 'same instant' is a matter of measurement precision. In light of the level of precision we had, we decided to allow a tolerance of one frame at the start of the step, one frame at the end, and an additional frame for the switch between the heavy diagonals. In addition, step candidates in which the mouse progressed in a non-diagonal manner (such as partial strides of bound or pace) were excluded from the general sequence.
The final stage was the sequence-recognition and edge-correction stage, in which steps identified at the third stage were concatenated into locomotion bouts. In these bouts the mouse trotted continuously. Steps located at the beginning or end of a bout were checked to verify that they were not part of a longer, irregular movement, as when the mouse raises a foreleg, pauses, and only then proceeds with a diagonal step. As in the third stage, we allowed for one frame of tolerance.
For each step, we calculate the difference in the timing of release of contact of the 2 legs that were heavy and become light, and the difference in the timing of establishment of contact of the 2 legs that were light and become heavy. The variance is the squared sum of these differences.
Fig9
Figure S1: Foot-fall Pattern juxtaposed with amount of Shift of Front and trotting variability. This plot depicts the entire sequence of a specific mouse. Black rectangles represent frames in which the corresponding foot was in contact with the ground. The color gradient represents the magnitude of per-step shifts of front, with yellow representing smaller than median (3.5°) values, orange-values between median and upper quartile (8°), red-values between upper quartile and quantile 0.9 (14.2°), dark red- between quantile 0.9 and maximal (38°) values, and white representing frames which were not recognized as trotting steps. The blue line represents the trot variability (see ‘Calculation of trot variability’). The predominance of the checkerboard pattern implies the use of the trot gait throughout different speeds (reflected by the length of black rectangles, i.e., length of support phase), and different curvatures (yellow and red bars). Note that there is no direct correlation between curvature, speed, and trot variability. (JPEG Image 236 kb)
Fig10
Figure S2: The Supporting Diagonal as a system of reference and its graphical representation. (a) A single diagonal step progressing from left to right. The green segment on the supporting diagonal represents the Normalized Intersection Point, with the supporting fore leg defined as zero and the supporting hind leg as one. The green sector between the diagonals represents the Direction of the swinging diagonal. (b) A curve representing the step in A. Each point corresponds to a single frame. Note that the points progress from right to left (towards the fore leg). (JPEG Image 104 kb)
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Gruntman, E., Benjamini, Y. & Golani, I. Coordination of steering in a free-trotting quadruped. J Comp Physiol A 193, 331–345 (2007). https://doi.org/10.1007/s00359-006-0187-5
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DOI: https://doi.org/10.1007/s00359-006-0187-5