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W4387043978 abstract "Full text Figures and data Side by side Abstract Editor's evaluation Introduction Results Discussion Methods Data availability References Decision letter Author response Article and author information Abstract Locomotor movements cause visual images to be displaced across the eye, a retinal slip that is counteracted by stabilizing reflexes in many animals. In insects, optomotor turning causes the animal to turn in the direction of rotating visual stimuli, thereby reducing retinal slip and stabilizing trajectories through the world. This behavior has formed the basis for extensive dissections of motion vision. Here, we report that under certain stimulus conditions, two Drosophila species, including the widely studied Drosophila melanogaster, can suppress and even reverse the optomotor turning response over several seconds. Such ‘anti-directional turning’ is most strongly evoked by long-lasting, high-contrast, slow-moving visual stimuli that are distinct from those that promote syn-directional optomotor turning. Anti-directional turning, like the syn-directional optomotor response, requires the local motion detecting neurons T4 and T5. A subset of lobula plate tangential cells, CH cells, show involvement in these responses. Imaging from a variety of direction-selective cells in the lobula plate shows no evidence of dynamics that match the behavior, suggesting that the observed inversion in turning direction emerges downstream of the lobula plate. Further, anti-directional turning declines with age and exposure to light. These results show that Drosophila optomotor turning behaviors contain rich, stimulus-dependent dynamics that are inconsistent with simple reflexive stabilization responses. Editor's evaluation The present study provides a valuable new perspective on the optomotor response based on an inversion of the behavior under specific (non-natural) conditions that may help elucidate the principles of this specific behavior. The evidence provided is convincing. https://doi.org/10.7554/eLife.86076.sa0 Decision letter Reviews on Sciety eLife's review process Introduction Visual navigation requires active mechanisms to stabilize locomotor trajectories through the world. Insects exhibit an optomotor turning response, a behavior in which they rotate their bodies in the direction of visual patterns that rotate about them (Hassenstein and Reichardt, 1956; Götz and Wenking, 1973; Buchner, 1976). This behavior is analogous to optomotor turning responses in fish (Clark, 1981) and the optokinetic response in mammals (Koerner and Schiller, 1972). In insects, this response is thought to be a course stabilizing mechanism that minimizes retinal slip, allowing animals to maintain their trajectory in the face of external or unexpected rotational forces (Götz and Wenking, 1973; Götz, 1975). For instance, if an insect attempts to walk in a straight line, it may slip and turn to the right. From the point of view of the insect, this turn is observed as optic flow rotating to the left. By responding to this leftward optic flow with a leftward turn, the insect can recover its original trajectory. In fruit flies, the optomotor response relies on well-characterized circuitry (Yang and Clandinin, 2018). Photoreceptor signals are split into parallel ON and OFF pathways in the lamina and medulla (Joesch et al., 2010; Clark et al., 2011; Behnia et al., 2014; Strother et al., 2014), which are not direction-selective. These signals provide input to T4 and T5 cells, which compute direction-selective responses along four directions at every point in the fly visual field (Bausenwein et al., 1992; Maisak et al., 2013; Takemura et al., 2013; Shinomiya et al., 2019; Henning et al., 2022). The outputs of T4 and T5 cells are then summed across visual space by lobula plate tangential cells (LPTCs) (Joesch et al., 2008; Schnell et al., 2012; Maisak et al., 2013; Mauss et al., 2015; Barnhart et al., 2018). Different LPTCs provide distinct signals about the overall pattern of motion surrounding the fly, and have been linked to head and body movements (Krapp and Hengstenberg, 1996; Haikala et al., 2013; Kim et al., 2017). There have been several reports of flies turning in the direction opposite to what is predicted by the optomotor turning response. In some cases, these counter-intuitive behaviors were observed using periodic stimuli with spatial wavelengths smaller than the receptive field of individual ommatidia, and thus can be accounted for by aliasing (Götz, 1964; Götz, 1970; Buchner, 1976). Work in a tethered flight simulator showed that when a moving pattern is presented in front of the fly, the animal turned in the direction of the stimulus motion (Tammero et al., 2004), as expected (Götz, 1968). However, if the moving pattern was presented behind the fly, it attempted to turn in the direction opposite to stimulus motion (Tammero et al., 2004). In a different experimental preparation, rotational patterns were presented on a dome around freely walking flies (Williamson et al., 2018). Under these conditions, flies generally turned in the direction of motion of the stimulus, but these rotations were often punctuated by brief, large-magnitude saccades in the opposite direction. Similarly, experiments using flight simulators have reported spikes in the torque in the direction opposite the stimulus rotation (Wolf and Heisenberg, 1990). Interestingly, zebrafish have also been observed to turn in the opposite direction of optic flow under certain conditions (Bak-Coleman et al., 2015). Here, we show that rotational stimuli can elicit strong, consistent anti-directional turning behavior in two drosophilid species, Drosophila melanogaster and Drosophila yakuba. We report that flies respond to high-contrast, high-luminance rotational motion stimuli by first turning in the direction of stimulus motion, and then reversing their trajectory after approximately 1 s, depending on the species. In D. melanogaster, we characterize the dynamics of this behavior and the stimuli that drive it, showing that it is distinct from prior observations of anti-directional turning. The behavior depends critically on adaptation to back-to-front motion. We use the genetic tools available in D. melanogaster to show that this behavior relies on the motion detecting neurons T4 and T5. Silencing horizontal system (HS) neurons and CH neurons, two wide-field neurons downstream of T4 and T5, resulted in small changes in this complex turning behavior. However, the visually evoked responses of these direction-selective neurons could not account for the anti-directional behavior. Thus, the observed reversal must be mediated by downstream circuitry. Overall, these results show that circuits in the fly generate behaviors that oppose the direction of wide-field visual motion, showing that Drosophila turning responses are more complex than a simple stabilizing reflex. Results Anti-directional turning responses to high-contrast stimuli Optomotor turning responses are central to gaze stabilization, so we sought to examine this response across different conditions. Many studies have investigated this behavior using stimuli with low contrast, low light intensity, or both (Götz and Wenking, 1973; Buchner, 1976; Rister et al., 2007; Seelig et al., 2010; Bahl et al., 2013; Bosch et al., 2015), at a variety of different speeds. However, natural scenes can have relatively high contrast and luminance, and such conditions have been poorly explored in the laboratory. In this experiment, we presented flies with rotational stimuli using high contrast and relatively high luminance. We tethered individual female D. melanogaster above a freely rotating ball to characterize the optomotor response (Buchner, 1976; Creamer et al., 2019; Figure 1a). As expected, low-contrast, slow-moving sinusoidal gratings caused flies to turn in the same direction as the moving gratings via the classical optomotor turning response (Figure 1b; Hassenstein and Reichardt, 1956; Götz, 1964; Buchner, 1976; Tammero et al., 2004; Seelig et al., 2010; Clark et al., 2011; Bahl et al., 2013; Silies et al., 2013; Clark et al., 2014; Bahl et al., 2015; Leonhardt et al., 2016; Salazar-Gatzimas et al., 2016; Strother et al., 2017; Creamer et al., 2018; Strother et al., 2018). However, when we changed the stimulus to high-contrast sinusoidal gratings (nominal 100% Weber contrast), flies turned in the stimulus direction for approximately 1 s, but then reversed course, and turned in the direction opposite to the stimulus motion for the duration of the stimulus presentation. Because this turning response is in the opposite direction of stimulus and the syn-directional optomotor turning response, we refer to it as anti-directional turning. Figure 1 with 2 supplements see all Download asset Open asset Flies turn opposite to the stimulus direction in high-contrast conditions. (a) We measured fly turning behavior as they walked on an air-suspended ball. Stimuli were presented over 270° around the fly. (b) We presented drifting sinusoidal gratings for 5 s (shaded region) with either high contrast (c=1.0) or low contrast (c=0.25). When high-contrast sinusoidal gratings were presented, flies initially turned in the same direction as the stimulus, then started turning in the opposite direction after ~1 s of stimulation. Under low-contrast conditions, flies turned continuously in the same direction as the stimulus. In these experiments, the sine waves had a wavelength of 60° and a temporal frequency of 1 Hz. Shaded patches represent ±1 SEM. N=10 flies. (c) We swept contrast between 0 and 1 and measured the mean turning response during the first 0.5 s (purple, purple bar in b) and during the last 4 s of the stimulus (brown, brown line in b). The response in the first 0.5 s increased with increasing contrast, while the response in the last 4 s increased from c=0 to c=0.25, and then decreased with increasing contrast, until flies turned in the direction opposite the stimulus direction at the highest contrasts. N=20 flies. (d) We repeated the presentation of drifting sinusoidal gratings, this time in the lab of author TRC, using a similar behavioral apparatus. Stimulus parameters were as described in (b). In these experiments, the population average shows that flies proceeded to zero net turning at high contrasts, but some individual flies exhibited anti-directional turning responses. N=20 flies. (e) We repeated the experiments with D. yakuba, also in the lab of TRC, and observed that this species exhibited a robust anti-directional turning response to high-contrast gratings and a classical syn-directional turning response to low-contrast gratings. N=11 flies. We swept a range of contrasts and compared the fly turning in the first 500 ms to the turning after 1 s (Figure 1c). As contrast increased, the flies turned faster during the first half second of stimulus presentation, reaching a plateau at around 0.5 contrast, consistent with previous results (McCann and MacGinitie, 1965; Buchner, 1976; Heisenberg and Buchner, 1977; Duistermars et al., 2007; Bahl et al., 2015; Strother et al., 2017). Fly behavior after the first second of stimulation was more complex. At contrasts between 0 and 0.25, flies turned in the same direction as the stimulus, with faster turning as the contrast increased. When the contrast was greater than 0.25, turning decreased, lowering to no net sustained turning at around 0.8 contrast. Above a contrast of 0.8, flies began to turn in the direction opposite the stimulus. These initial experiments took place in the lab of author DAC. To confirm that these unexpected responses did not reflect some idiosyncrasy of one specific behavioral apparatus or environment, we repeated these experiments in a second lab, that of author TRC. Under similar conditions, using the same strain of D. melanogaster, we reproduced the rapid deceleration after an initial, transient syn-directional response (Figure 1d), with some individual flies exhibiting significant anti-directional turning (Figure 1—figure supplement 1). This demonstrates that the key features of this behavioral response are stable across experimental systems and laboratories, though the magnitude of anti-directional turning behavior in D. melanogaster is sensitive to some unknown experimental parameter differences between the laboratories. Individual strains of D. melanogaster, and other drosophilid species, display significant variation in their locomotor patterns during walking (York et al., 2022). Indeed, when we tested a Canton-S D. melanogaster strain, we observed milder but significant anti-directional turning at long timescales (Figure 1—figure supplement 2b). We reasoned that a strong test of the generality of anti-directional turning would be to examine turning behavior in another species, and selected D. yakuba. Strikingly, D. yakuba also displayed anti-directional turning behavior under similar conditions (Figure 1e). Thus, this behavior is not an idiosyncratic feature of a single laboratory strain. Conditions for anti-directional turning behaviors While anti-directional turning behaviors have been reported before, other groups have presented similar stimuli without observing anti-directional behavior (Götz and Wenking, 1973; Buchner, 1976; Rister et al., 2007; Seelig et al., 2010; Bahl et al., 2013; Bosch et al., 2015). We wondered what aspects of our experimental setup could lead to these behavioral differences. In our experiments, anti-directional turning was strongly linked to display brightness (Figure 1—figure supplement 2a). When the mean brightness of the screens was reduced from 100 cd/m2 to 1 cd/m2, we saw no anti-directional turning in 5 s trials (though average optomotor behavior did decrease over the course of the stimulus presentation). When we further reduced the mean brightness to 0.1 cd/m2, flies persisted in their optomotor behavior throughout the stimulus presentation. We note that in these low-luminance experiments, low levels of ambient light in the nominally dark experimental rig could also reduce the effective contrast of the stimulus. We tested a variety of other factors that might affect anti-directional turning. Anti-directional turning occurred when experiments were run both at hot temperatures and at room temperature (Figure 1—figure supplement 2b). We also observed anti-directional behavior when flies were reared in the dark and on different media. We also tested several other experiment conditions (Figure 1—figure supplement 2c). Flies responded with anti-directional turning to high-contrast stimuli presented at both blue and green wavelengths. We glued fly heads to their thorax to ensure stimuli could not be affected by head movements (Haikala et al., 2013; Kim et al., 2017), but found no difference between head-fixed and head-free flies. We did find a few factors that modulated anti-directional turning behavior. In particular, rearing D. melanogaster at 25°C instead of 20°C or testing flies that were 2 weeks old instead of 12–60 hr old both reduced overall turning behavior and eliminated anti-directional turning. In these cases, optomotor turning still decreased over the course of the 5 s, high-contrast trials, but did not reverse. As details of rearing temperature and the age at which behavior tests are run often vary across labs, it is possible that these factors, as well as stimulus brightness, account for the differences between our observations and the previous literature. Distinct spatiotemporal tuning of the anti-directional behavioral response To further characterize the anti-directional response, we swept the spatial and temporal frequency of the sinusoidal grating stimulus. Using only Weber contrasts of 1, we compared the early response (first quarter second, Figure 2a) to the late response (after 1 s, Figure 2b). D. melanogaster always turned in the optomotor direction during the early stimulus response. In this early response, flies turned most vigorously to stimuli with short spatial frequencies (~20° wavelength) and fast temporal frequencies (~8 Hz), in agreement with earlier studies (Tammero et al., 2004; Creamer et al., 2018; Strother et al., 2018). However, during the longer-timescale response to high-contrast stimuli, flies only turned in the optomotor direction at very high temporal frequencies (>~16 Hz) and at very low temporal frequencies (<0.5 Hz). At intermediate temporal frequencies, flies showed a sustained anti-directional response. The maximal anti-directional response was achieved at 1 Hz and 45° wavelength, distinct from the conditions for peak classical turning responses. Interestingly, the stimuli that elicit the strongest anti-directional response appear similar to those that maximally activate T4 and T5 neurons when those neurons are measured in head-fixed flies (Maisak et al., 2013; Leong et al., 2016; Arenz et al., 2017; Creamer et al., 2018; Strother et al., 2018; Wienecke et al., 2018). Figure 2 with 1 supplement see all Download asset Open asset Anti-directional turning behavior has distinct tuning and is driven by adaptation. (a) Heatmap of fly turning velocity during the first 0.5 s of sinusoidal grating stimulation under high-contrast conditions and variable temporal and spatial frequencies. The flies turned in the direction of the stimulus across all conditions and responded most to 8 Hz, 22° stimuli. N=16, 21, 17, 21, 7, and 22 flies for spatial frequencies 1/120°, 1/90°, 1/60°, 1/45°, 1/30°, and 1/22° respectively. (b) Heatmap as in (a), measured during the last 4 s of stimulation. Flies turned in the same direction as the stimulus at high and low temporal frequencies, but in the opposite direction of the stimulus at intermediate temporal frequencies, with a maximal anti-directional response at wavelengths between 30° and 60°. (c) Switching stimulus contrast from high to low after 5 s caused flies to revert to syn-directional behavior after the anti-directional response. N=7 flies. (d) Presenting rotating random binary patterns (5° vertical strips rotating at 150 °/s) induced anti-directional turning similar to that elicited by rotating sine wave gratings. N=7 flies. (e) We presented flies with 5 s of ‘translational’ stimuli (dark shaded region), with high-contrast sinusoidal gratings moving either front-to-back or back-to-front, bilaterally, for 5 s. After that, we presented high-contrast rotational sinusoidal grating stimuli (60° wavelength, 1 Hz). Front-to-back stimulation did not affect the subsequent response to rotational stimuli, but back-to-front stimuli caused flies to turn immediately in the opposite direction of the stimulus. N=18 flies. Anti-directional turning results from adaptation effects We were intrigued by the switch from syn-directional to anti-directional turning behavior. To investigate the dynamics of these changes, we presented a rotating sinusoidal stimulus at contrast 1 for 5 s, and then changed the contrast to 0.25 (Figure 2c). After the switch to low contrast, the flies quickly reverted classical, syn-directional optomotor behavior, demonstrating that no long-term switch in directional turning occurs during high-contrast stimulus presentation. This effect did not depend on the periodic nature of these stimuli: a rotating stimulus consisting of 5°-wide vertical bars with randomly chosen, binary contrasts (Clark et al., 2014) yielded similar behavioral responses (Figure 2d). To further isolate the causes of this switch in behavior, we developed a protocol to adapt the fly to different stimuli before presenting the high-contrast rotational sinusoidal gratings to elicit the anti-directional turning response. This adapting protocol consisted of 5 s presentations of an adaptor stimulus, followed by a high-contrast rotational stimulus (Figure 2e, Figure 2—figure supplement 1). The adaptor stimuli we tested were a uniform gray screen, a stationary high-contrast sinusoid, a closed-loop high-contrast sinusoid, or a high-contrast ‘translational’ sinusoidal stimulus. During the closed-loop adapter, the stimulus position was yoked to the fly’s own turning to better simulate real rotation, a situation we hypothesized might yield less dependence on stimulus contrast (following Leonhardt et al., 2016). The translational stimulus had both left and right hemifields moving either front-to-back or back-to-front across the fly’s two eyes (Creamer et al., 2018). These translational stimuli resulted in no net turning (Silies et al., 2013; Creamer et al., 2018), but specifically adapt motion detectors in both eyes selective for front-to-back or back-to-front motion. We found that adapting with a stationary or closed-loop high-contrast sinusoid had little effect on the behavior (Figure 2—figure supplement 1). Similarly, adapting the fly with front-to-back stimuli did not have a strong effect on the subsequent response to rotational stimuli. However, adapting with back-to-front stimuli generated responses that no longer showed an initial syn-directional turning response, but instead exhibited anti-directional turning immediately after the rotational stimulus began. This result indicates that the anti-directional turning results from slow-timescale changes that depend on strong back-to-front motion. Anti-directional turning is elicited when stimuli are presented in front of the fly A previous report of anti-directional turning behavior in flying tethered flies showed that flies turn in the opposite direction to stimuli that are presented behind their midline (Tammero et al., 2004). To test whether our results were caused by this effect, we split our stimulus into three regions: 90° in front of the fly, 45° in front of the midline on either side of the fly, and 45° behind the midline on either side of the fly (Figure 3a). We found that flies displayed anti-directional turning when presented with stimuli only in the front region or only just in front of the midline (Figure 3b, c). They did not display anti-directional turning when moving stimuli were presented behind the midline (Figure 3b, c). This suggests a different mechanism from the behaviors that depend on posterior spatial location to elicit reverse turning (Tammero et al., 2004). Figure 3 with 1 supplement see all Download asset Open asset Anti-directional turning is driven by stimuli in the forward-facing visual field and is not driven by saccades. (a) We divided our panoramic display into three sections: the front 90°, the 45° behind the fly on either side, and a middle 45°. (b) High-contrast sinusoidal gratings were presented on each of these three display sections, with the remaining sections blank. Flies turned syn-directionally when stimuli were presented behind the fly, and turned anti-directionally when stimuli were presented in front of the fly. Shaded patches represent ±1 SEM. N=55 flies. (c) Average turning in the last 4 s of the stimulus (black bar in b), in low-contrast and high-contrast conditions. Shaded patches in the time trace plots represent ±1 SEM. N=55 flies. (d) A single fly responds to many trials of sinusoidal grating stimuli at high contrast (blue bar) and low contrast (orange bar). We show a heatmap of the fly’s responses over time (horizontal axis) and across trials (vertical axis). (e) We can ignore the magnitude of the turning and instead only quantify whether the fly was turning in the same direction as the stimulus (white area) or in the opposite direction (dark gray area). This shows sustained anti-directional turning, not brief saccades. (f) Averaging the direction (but not magnitude) of turning across trials and across flies yields a turning index for each point in time. Shaded patches in the time trace plots represent ±1 SEM. N=7 flies. Anti-directional responses do not depend on saccades Anti-directional saccades have been reported in walking and flying flies (Wolf and Heisenberg, 1990; Williamson et al., 2018). In walking flies (Williamson et al., 2018), flies largely turned in syn-directionally, but these turns were sometimes interrupted by brief, high-amplitude saccades in the opposite direction, against the stimulus direction. If such saccades were frequent or high amplitude, the net effect could shift the average turning we measured, creating apparent anti-directional turning. To investigate this possibility, we plotted the turning response on a per-trial basis (Figure 3d). We then discarded information about the magnitude of the turns and considered only the direction of the turning at each point in time (Figure 3e). Strikingly, in many trials, flies continued to turn opposite to the stimulus for several seconds, a behavior unlike brief saccades. This prolonged turning against the stimulus was also observable in the distribution of turning over time in all trials over all flies (Figure 3—figure supplement 1). Next, we calculated a turning index for each response timepoint (sampled at 60 Hz). This turning index represents the fraction of trials in which the fly turned in the direction of the stimulus at each timepoint minus the fraction of trials in which the fly turned in the opposite direction (Figure 3f). Since this turning index does not include the magnitude of turning, it is strongly affected by sustained low-amplitude turns and discounts any brief high-amplitude saccades. When presented with high-contrast stimuli, flies maintained a negative turning index, indicating that sustained turns, and not high-velocity saccades, underlie this anti-directional turning behavior. As such, it appears distinct from the reports of anti-directional saccades. Anti-directional turning requires elementary motion detectors What neurons are involved in this anti-directional turning behavior? Previous work demonstrated that T4 and T5 are required for directional neural responses (Schnell et al., 2012), as well as for optomotor turning (Maisak et al., 2013; Salazar-Gatzimas et al., 2016; Salazar-Gatzimas et al., 2018), for walking speed regulation (Creamer et al., 2018), and for responses to visual looming stimuli (Schilling and Borst, 2015). We silenced the neurons T4 and T5 using shibirets (Kitamoto, 2001) and measured responses to sinusoidal stimuli that switched from high to low contrast (Figure 4a). Flies in which T4 and T5 had been silenced displayed only minimal responses to motion stimuli, with anti-directional turning suppressed along with classical syn-directional turning. Thus, we conclude that, like optomotor turning behaviors, this anti-directional behavior depends critically on signals from T4 and T5. Figure 4 Download asset Open asset Syn-directional and anti-directional turning share common circuitry. (a) We silenced T4 and T5 neurons by expressing shibirets selectively in those neurons. We measured turning behavior during a contrast-switching stimulus (as in Figure 2c). Results from flies with T4 and T5 silenced shown in dark red, while controls are in light red and gray. Average fly behavior during the last 4 s of the first contrast (black bar on left) shown as bars on the right, with individual fly behavior shown as dots. Note that the data labeled ‘low contrast’ are from experiments in which the low-contrast stimulus was shown before the high-contrast stimulus. Shaded patches in the time trace plots represent ±1 SEM, as do vertical lines on bar plots. *** indicates experimental results are significantly different from results, p<0.001 via a two-sample Student’s t-test. * indicates p<0.05. N=17, 24, 19 flies with genotypes T4T5/Shibirets, T4T5/+, +/Shibirets. (b) Results from horizontal system (HS) neuron silencing as in (a). Silencing HS neurons reduced syn-directional turning behavior (p<0.001) but did not have a strong effect on anti-directional turning. N=34, 21, 19 flies with genotypes HS/Shibirets, HS/+, +/Shibirets. (c) Results from CH neuron silencing as in (a). CH neuron silencing reduced the degree of anti-directional turning (p<0.001). N=63, 57, 70 flies with genotypes CH/Shibirets, CH/+, +/Shibirets. Anti-directional turning requires the CH LPTC Since the switch from optomotor to anti-directional behavior seems to be dependent on the direction of motion adaptation (Figure 2e), we reasoned that neurons involved in this behavior were likely to be downstream from T4 and T5. Relatively little is known about circuitry that connects the neurons T4 and T5 to optomotor turning behavior. However, horizontal system (HS) cells are well-studied postsynaptic partners of T4 and T5 (Joesch et al., 2008; Joesch et al., 2010). These LPTCs integrate information from front-to-back and back-to-front selective T4 and T5 cells across the fly’s visual field (Mauss et al., 2015). HS cells have been implicated in visually evoked head turns (Kim et al., 2017) and body rotations in flight (Haikala et al., 2013) and in maintenance of direction during walking (Fujiwara et al., 2022). When we silenced HS neurons, we found small deficits in syn-directional turning behavior, consistent with prior results, but no deficits in anti-directional turning (Figure 4b), indicating that HS cells synaptic output is not required specifically for anti-directional turning behavior. Next, we turned to the CH LPTCs. These cells are GABAergic and are both pre-synaptic and post-synaptic in the lobula plate (Wei et al., 2020). In blowflies, these neurons play an inhibitory role in an interconnected LPTC circuit that shapes behavior (Borst and Weber, 2011). When we silenced CH neurons, we found a small increase in syn-directional turning and a decrease in anti-directional turning (Figure 4c). Overall, silencing this neuron type caused the flies to turn more in the direction of motion. This result suggests that CH activity contributes to the ant" @default.
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W4387043978 date "2023-06-08" @default.
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W4387043978 title "Decision letter: Long-timescale anti-directional rotation in Drosophila optomotor behavior" @default.
W4387043978 doi "https://doi.org/10.7554/elife.86076.sa1" @default.
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