(A) Key partners of the sensorimotor pathway bridging the Or42a-expressing olfactory sensory neuron (OSN, yellow) to the descending neuron PDM-DN (black), down to the A27h premotor circuit (green) in the ventral nerve cord. (B) Putative sensorimotor transformation of positive changes in odor concentration (C(t)) during up-gradient runs. Positive gradients detected by the Or42a OSN promote the OSN activity, which is expected to strongly active its cognate uniglomerular projection neuron (uPN, orange). Due to the cholinergic nature of olfactory projection neurons, excitation of the Or42a uPN promotes the activity of the LH-LN1 neuron (pink) located in the lateral horn region. In turn, the activity of LH-LN1 controls both the LH-LN2 and PDM-DN neuron. We speculate that the 3-element feedforward motif formed by LH-LN1, LH-LN2 and PDM-DN converts the activity of Or42a uPN into an inhibition of PDM-DN. As PDM-DN is cholinergic, the lack of activity of PDM-DN is expected to leave its downstream GABAergic partner SEZ-DN1 (blue) inactive. As a result, the sensorimotor pathway mediated by PDM-DN has no inhibitory effect on the A27h premotor circuit (green), which promotes forward peristalsis (runs). As indicated in the legend, neurons with a contour in black are thought to be excited. Neurons with a contour in red are thought to be inhibited or inactive. (C) Same as B upon detection of negative odor changes during down-gradient runs. Negative odor gradients inhibit the activity of Or42a OSN, which is thought to leave the Or42a uPN inactive. As a result, we speculate that the 3-element feedforward motif formed by LH-LN1, LH-LN2 and PDM-DN converts the inactivity of Or42a uPN into an excitation of PDM-DN. In turn, the firing activity of the cholinergic PDM-DN is thought to excite SEZ-DN1. Due to the GABAergic nature of SEZ-DN1, excitation of PDM-DN and SEZ-DN1 represses the activity of the A27h premotor circuit (green) in the posterior segments of the ventral nerve cord (VNC), thereby triggering an interruption in forward peristalsis (stop). Videos.