(
A) Schematic diagram of the
C. elegans nervous system. (
B) Multi-neuron calcium imaging in freely behaving animals (P
nmr-1/P
lim-4/P
lgc-55/P
npr-9::NLS::wCherry::SL2::GCaMP6). Interneurons in the backward and forward modules were simultaneously imaged using a confocal microscopy (see Materials and methods for details). Data are aligned to the onset of a reversal (up, vertical dashed lines,
t = 0) or the end of a turn (bottom, vertical dashed lines,
t = 0). Heat map across trials and Δ
R(
t)/
R0 (Mean ± SEM) across trials are shown. (
C) Ramping rate of AIB calcium activity during thermally-induced escape responses. Ramping rate is significantly higher in type-II transition (nine animals, each point has more than three trials). *p<0.05, two-way ANOVA. (
D–E) Characterization of the probability of type-II transition during optogenetic stimulation of two different interneurons AIB (left) and RIM (right) for varying durations. Longer activation of AIB consistently led to a higher probability of type-II transition (left). The control groups are animals performing spontaneous motor state transitions without optogenetic stimulation. A blue laser (473 nm, 14.71 mW/mm
2) was used to stimulate ChR2 expressed neurons. (
F–G) Hyperpolarization of RIM (P
tdc-1::Arch) or AIB (P
npr-9::Arch) both affected reversal duration (bar graph), but only inhibiting AIB affected the probability of type-II transition (pie chart). In
F), simultaneous optogenetic activation of AIB (or RIM) (635 nm, 4.46 mW/mm
2 for AIB or 473 nm, 14.71 mW/mm
2 for RIM) and inhibition (561 nm, 6.80 mW/mm
2) of RIM (or AIB) were carried out for 7 (or 5) seconds. In
G), RIM were activated for varying durations, while AIB were inactivated through an exogenous expression of the potassium channel TWK-18. (
H–I) Reversal length distribution and transition rates after optogenetic ablation of AIB. ALM/AVM were optogenetically activated for 1.5 s to trigger avoidance behavior. AIB were ablated by exogenous expression of a membrane targeted PH-miniSOG. Data from
Figure 1C) are used as control. For reversal duration, error bars indicate SEMs. Mann–Whitney U test. For probability of turn, error bars indicate 95% binomial proportion confidence interval. χ
2 test. *p<0.05, **p<0.01, ***p<0.001, ****p<0.0001. All multiple comparisons were adjusted using Bonferroni correction.