Dynamics of chemo-receptor activity with time-periodic attractant field

When exposed to a time-periodic chemical signal, an \textit{E.~coli} cell responds by modulating its receptor activity in a similar time-periodic manner. However, there exists a phase lag between the applied signal and the activity response. We study the variation of the activity amplitude and phase lag as a function of the applied frequency~$\omega$, using numerical simulations. The amplitude increases with~$\omega$, reaches a plateau, and then decreases again for large~$\omega$. The phase lag increases monotonically with~$\omega$ and finally saturates to $3\pi/2$ when~$\omega$ is large. The activity is no longer a single-valued function of the attractant signal, and plotting activity versus attractant concentration over one complete time period generates a loop. We monitor the loop area as a function of~$\omega$ and find two peaks for small and large~$\omega$, and a sharp minimum at intermediate~$\omega$ values. We explain these results as an interplay between the time scales associated with adaptation, activity switching, and applied signal variation. In particular, for very large~$\omega$, the quasi-equilibrium approximation for activity dynamics breaks down, a regime that has not been explored in earlier studies. We perform analytical calculations in this limit and find good agreement with our simulation results.