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We present a high-throughput optogenetic illumination system capable ofsimultaneous closed-loop light delivery to specified targets in populations ofmoving Caenorhabditis elegans. The instrument addresses three technicalchallenges: it delivers targeted illumination to specified regions of theanimal's body such as its head or tail; it automatically delivers stimulitriggered upon the animal's behavior; and it achieves high throughput bytargeting many animals simultaneously. The instrument was used tooptogenetically probe the animal's behavioral response to competingmechanosensory stimuli in the the anterior and posterior soft touch receptorneurons. Responses to more than $10^4$ stimulus events from a range ofanterior-posterior intensity combinations were measured. The animal'sprobability of sprinting forward in response to a mechanosensory stimulusdepended on both the anterior and posterior stimulation intensity, while theprobability of reversing depended primarily on the posterior stimulationintensity. We also probed the animal's response to mechanosensory stimulationduring the onset of turning, a relatively rare behavioral event, by deliveringstimuli automatically when the animal began to turn. Using this closed-loopapproach, over $10^3$ stimulus events were delivered during turning onset at arate of 9.2 events per worm-hour, a greater than 25-fold increase in throughputcompared to previous investigations. These measurements validate with greaterstatistical power previous findings that turning acts to gate mechanosensoryevoked reversals. Compared to previous approaches, the current system offerstargeted optogenetic stimulation to specific body regions or behaviors withmany-fold increases in throughput to better constrain quantitative models ofsensorimotor processing.

Closed-loop targeted optogenetic stimulation of C. elegans populations