Interestingly, the steepness of the temperature gradient can induce different behavioral outputs, indicating that C. elegans can recognize temperature changes in time and space

To apply temperature stimuli to animals, recent studies have used agar pads to mount animals and then modulate the surrounding environmental temperature with the aid of a thermocouple (
8
,
9
,
10
). However, these methods can only deliver the stimuli with temporal resolution on the order of 10 s, if not longer. Alternative approaches typically use either a sinusoidal or nonchanging temperature stimulus (
8
,
9
). These methods also require manually gluing worms onto the pad (
10
), which is time consuming and can affect the animals' responses. Laser stimulation has also been used, but only for higher temperatures and typically with the purpose of inducing a pain-like response, and the exact temperature is unknown (
11
). Therefore, there is a need for a novel platform that can deliver temperature stimuli (both hot and cold), without appreciable spatial gradients, and, importantly, with a sub-second temporal resolution, while allowing the simultaneous high-resolution monitoring of neuronal responses.