Because incoming light, in both the left and right eyes, contracts their rhabdomeres in the back-to-front direction, photoreceptors can transmit information about the field rotation or the fly’s orientation changes to the brain. Specifically, the output from the eye towards which the fly turns phase leads, and thus neurally signals rotation direction. Here we illustrate the underlying mechanism by using a brief 205 o/s left saccade (in a world of 6.8o black-and-white gratings), as an example. (A) During the left saccade, the flow field facing the left eye moves back-to-front (red arrow) across its photoreceptors’ receptive fields (i). The image of the moving flow field is inverted by the ommatidium lens, and so moves front-to-back (blue arrow) while being sampled by the rhabdomere (ii). Light contracts the rhabdomeres in back-to-front direction (red arrow). With the projected image and the photon-sampler (rhabdomere) moving against each other, the light input from two bright bars (iii) becomes narrowed and accelerated. Accordingly, photoreceptor output shows two prominent peaks, in which distance to each other is compacted in time (iv). (B) During the left saccade, the flow field facing the right eye moves front-to-back (blue arrow) across its photoreceptors’ receptive fields (i). Again, the ommatidium lens inverts the image, which now moves back-to-front (red arrow) over the rhabdomere (ii), which contracts in the same direction (red arrow). With the projected image and the rhabdomere moving together, the light input from two bright bars (iii) excites the photoreceptor longer, in which output shows two prominent peaks elongated in time (iv). (C) The light input to the left eye photoreceptors (red trace) rise and decay faster than the corresponding light input (blue trace) to the right eye photoreceptors. (D) R1-R6 output in the left eye (red trace) is faster and briefer than the photoreceptor output (blue trace) in the right eye. (E) Consequently, by correlating the outputs from the left and right eye photoreceptors in time, the fly brain can obtain information about the directional changes in the visual space, in respect to its head orientation.