10The bias for a particular direction of rotation of the flagellar motor is a sensitive readout of chemotaxis signaling, which mediates bacterial migration towards favorable chemical environments. The rotational bias has not been characterized in Helicobacter pylori, which limits our understanding of the signaling dynamics. Here, we determined that H. pylori swim faster (slower) whenever their flagella rotate counterclockwise (clockwise) by analyzing their 15 hydrodynamic interactions with bounding surfaces. The anisotropy in swimming speeds helped quantify the fraction of the time that the cells swam slower to report the first measurements of the bias. A stochastic model of run-reversals indicated that the anisotropy promotes faster spread compared to isotropic swimmers. The approach further revealed that the diffuse spread of H. pylori is likely limited at the physiological temperature due to increased reversal frequencies. 20 Thus, anisotropic run-reversals make it feasible to study signal-response relations in the chemotaxis network in non-model bacterial species. Page 2 of 22 Impact Statement Anisotropy in run and reversal swimming speeds promotes the spread of H. pylori and reveals temperature-dependent behavior of the flagellar switch. cell was observed, B C , to yield: FG HC*3,C = J K LM J KThe error associated with the calculation of FG HC*3,C values decreases with increasing B C . But, 15 different cells were observed for different durations; hence the FG HC*3,C values were allocated weights that corresponded to their respective durations:
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