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• W2071663937 abstract "•Zebrafish hunting responses are triggered by conjunctions of visual features•Tectal neurons show non-linear mixed selectivity for prey-like visual stimuli•Tectal assemblies show premotor activity specifically preceding hunting responses Visuomotor circuits filter visual information and determine whether or not to engage downstream motor modules to produce behavioral outputs. However, the circuit mechanisms that mediate and link perception of salient stimuli to execution of an adaptive response are poorly understood. We combined a virtual hunting assay for tethered larval zebrafish with two-photon functional calcium imaging to simultaneously monitor neuronal activity in the optic tectum during naturalistic behavior. Hunting responses showed mixed selectivity for combinations of visual features, specifically stimulus size, speed, and contrast polarity. We identified a subset of tectal neurons with similar highly selective tuning, which show non-linear mixed selectivity for visual features and are likely to mediate the perceptual recognition of prey. By comparing neural dynamics in the optic tectum during response versus non-response trials, we discovered premotor population activity that specifically preceded initiation of hunting behavior and exhibited anatomical localization that correlated with motor variables. In summary, the optic tectum contains non-linear mixed selectivity neurons that are likely to mediate reliable detection of ethologically relevant sensory stimuli. Recruitment of small tectal assemblies appears to link perception to action by providing the premotor commands that release hunting responses. These findings allow us to propose a model circuit for the visuomotor transformations underlying a natural behavior. Visuomotor circuits filter visual information and determine whether or not to engage downstream motor modules to produce behavioral outputs. However, the circuit mechanisms that mediate and link perception of salient stimuli to execution of an adaptive response are poorly understood. We combined a virtual hunting assay for tethered larval zebrafish with two-photon functional calcium imaging to simultaneously monitor neuronal activity in the optic tectum during naturalistic behavior. Hunting responses showed mixed selectivity for combinations of visual features, specifically stimulus size, speed, and contrast polarity. We identified a subset of tectal neurons with similar highly selective tuning, which show non-linear mixed selectivity for visual features and are likely to mediate the perceptual recognition of prey. By comparing neural dynamics in the optic tectum during response versus non-response trials, we discovered premotor population activity that specifically preceded initiation of hunting behavior and exhibited anatomical localization that correlated with motor variables. In summary, the optic tectum contains non-linear mixed selectivity neurons that are likely to mediate reliable detection of ethologically relevant sensory stimuli. Recruitment of small tectal assemblies appears to link perception to action by providing the premotor commands that release hunting responses. These findings allow us to propose a model circuit for the visuomotor transformations underlying a natural behavior. To generate visually guided behavior, the nervous system extracts task-relevant information from the retinal image to select and control an appropriate response. Over 50 years ago, neuroethologists introduced the idea that specific behaviors can be triggered by “key stimuli,” delivered under appropriate conditions [1Tinbergen N. The Herring Gull’s World. Collins, London1953Google Scholar, 2Ingle D. Crews D. Vertebrate neuroethology: definitions and paradigms.Annu. Rev. Neurosci. 1985; 8: 457-494Crossref PubMed Scopus (20) Google Scholar]. In this context, individual neurons have been discovered in visual pathways that are proposed to function as “feature detectors.” Such neurons are selective for specific spatiotemporal patterns within the visual scene and include neurons tuned to visual features that define key stimuli. Notably, stimulus-response pathways are subject to various modulating influences, and consequently “key stimuli” do not always trigger a response. Motivational state, arousal, attention, recent experience, and long-term memory can influence response probability, stimulus preference, and the choice of motor outputs (e.g., [3Laming P.R. Cairns C. Effects of food, glucose, and water ingestion on feeding activity in the toad (Bufo bufo).Behav. Neurosci. 1998; 112: 1266-1272Crossref PubMed Scopus (9) Google Scholar, 4Brzoska J. Schneider H. Modification of prey-catching behavior by learning in the common toad (Bufo b. bufo [L], Anura, Amphibia): Changes in responses to visual objects and effects of auditory stimuli.Behav. Processes. 1978; 3: 125-136Crossref PubMed Scopus (22) Google Scholar]). Therefore, to understand how sensorimotor circuits link perception to action, it is necessary to monitor neural activity and behavior simultaneously. In larval zebrafish, the small size and optical transparency of the nervous system allows functional imaging of neural activity at cellular resolution and throughout the brain, during behavior [5O’Malley D.M. Kao Y.H. Fetcho J.R. Imaging the functional organization of zebrafish hindbrain segments during escape behaviors.Neuron. 1996; 17: 1145-1155Abstract Full Text Full Text PDF PubMed Scopus (246) Google Scholar, 6Ahrens M.B. Li J.M. Orger M.B. Robson D.N. Schier A.F. Engert F. Portugues R. Brain-wide neuronal dynamics during motor adaptation in zebrafish.Nature. 2012; 485: 471-477PubMed Google Scholar, 7Portugues R. Feierstein C.E. Engert F. Orger M.B. Whole-brain activity maps reveal stereotyped, distributed networks for visuomotor behavior.Neuron. 2014; 81: 1328-1343Abstract Full Text Full Text PDF PubMed Scopus (171) Google Scholar]. In this study, we used two-photon (2P) calcium imaging to examine how perception of prey-like visual cues leads to initiation of hunting. In larval zebrafish, prey catching is a visually guided behavior [8Borla M.A. Palecek B. Budick S. O’Malley D.M. Prey capture by larval zebrafish: evidence for fine axial motor control.Brain Behav. Evol. 2002; 60: 207-229Crossref PubMed Scopus (106) Google Scholar, 9McElligott M.B. O’malley D.M. Prey tracking by larval zebrafish: axial kinematics and visual control.Brain Behav. Evol. 2005; 66: 177-196Crossref PubMed Scopus (102) Google Scholar, 10Gahtan E. Tanger P. Baier H. Visual prey capture in larval zebrafish is controlled by identified reticulospinal neurons downstream of the tectum.J. Neurosci. 2005; 25: 9294-9303Crossref PubMed Scopus (227) Google Scholar]. Several studies have examined the locomotor and oculomotor components of hunting routines including the kinematic features of orienting turns (described as J-turns in [9McElligott M.B. O’malley D.M. Prey tracking by larval zebrafish: axial kinematics and visual control.Brain Behav. Evol. 2005; 66: 177-196Crossref PubMed Scopus (102) Google Scholar]), capture swims [8Borla M.A. Palecek B. Budick S. O’Malley D.M. Prey capture by larval zebrafish: evidence for fine axial motor control.Brain Behav. Evol. 2002; 60: 207-229Crossref PubMed Scopus (106) Google Scholar, 11Patterson B.W. Abraham A.O. MacIver M.A. McLean D.L. Visually guided gradation of prey capture movements in larval zebrafish.J. Exp. Biol. 2013; 216: 3071-3083Crossref PubMed Scopus (59) Google Scholar], and coordinated pectoral fin movements [12McClenahan P. Troup M. Scott E.K. Fin-tail coordination during escape and predatory behavior in larval zebrafish.PLoS ONE. 2012; 7: e32295Crossref PubMed Scopus (37) Google Scholar]. Of particular relevance to this study, zebrafish larvae perform a specialized oculomotor behavior, eye convergence, specifically during hunting. A convergent saccade defines the onset of all hunting routines, and the eyes maintain a high vergence angle until after the strike at prey [13Bianco I.H. Kampff A.R. Engert F. Prey capture behavior evoked by simple visual stimuli in larval zebrafish.Front. Syst. Neurosci. 2011; 5: 101Crossref PubMed Scopus (161) Google Scholar]. After the initial convergent saccade, vergence angle further increases during prey tracking, in relation to target proximity [11Patterson B.W. Abraham A.O. MacIver M.A. McLean D.L. Visually guided gradation of prey capture movements in larval zebrafish.J. Exp. Biol. 2013; 216: 3071-3083Crossref PubMed Scopus (59) Google Scholar]. By increasing the extent of the binocular visual field and advancing it close to the nose of the animal, eye convergence might enable a stereopsis mechanism for judging target distance and triggering the final capture event [13Bianco I.H. Kampff A.R. Engert F. Prey capture behavior evoked by simple visual stimuli in larval zebrafish.Front. Syst. Neurosci. 2011; 5: 101Crossref PubMed Scopus (161) Google Scholar]. The optic tectum (OTc) is the largest retinorecipient structure in the brain of teleost fish and is likely to be of central importance for hunting behavior. Visual space is retinotopically mapped across the OTc in register with the tectal motor map and as such the OTc is well suited to control goal-directed behaviors toward specific points in space [14Stuermer C.A. Retinotopic organization of the developing retinotectal projection in the zebrafish embryo.J. Neurosci. 1988; 8: 4513-4530PubMed Google Scholar]. These include orienting and avoidance behaviors [15Torres B. Luque M.A. Pérez-Pérez M.P. Herrero L. Visual orienting response in goldfish: a multidisciplinary study.Brain Res. Bull. 2005; 66: 376-380Crossref PubMed Scopus (14) Google Scholar], saccadic eye movements [16Klier E.M. Wang H. Crawford J.D. The superior colliculus encodes gaze commands in retinal coordinates.Nat. Neurosci. 2001; 4: 627-632Crossref PubMed Scopus (137) Google Scholar], and prey-catching behaviors including striking at prey [17Chen Q. Deng H. Brauth S.E. Ding L. Tang Y. Reduced performance of prey targeting in pit vipers with contralaterally occluded infrared and visual senses.PLoS ONE. 2012; 7: e34989Crossref PubMed Scopus (20) Google Scholar]. Indeed, neural activity in the OTc of larval zebrafish was recently observed in response to live prey [18Muto A. Ohkura M. Abe G. Nakai J. Kawakami K. Real-time visualization of neuronal activity during perception.Curr. Biol. 2013; 23: 307-311Abstract Full Text Full Text PDF PubMed Scopus (156) Google Scholar]. Zebrafish hunting is greatly reduced by ablating the retinal input to the tectum [10Gahtan E. Tanger P. Baier H. Visual prey capture in larval zebrafish is controlled by identified reticulospinal neurons downstream of the tectum.J. Neurosci. 2005; 25: 9294-9303Crossref PubMed Scopus (227) Google Scholar], silencing a specific population of tectal interneurons [19Del Bene F. Wyart C. Robles E. Tran A. Looger L. Scott E.K. Isacoff E.Y. Baier H. Filtering of visual information in the tectum by an identified neural circuit.Science. 2010; 330: 669-673Crossref PubMed Scopus (181) Google Scholar], or a genetic mutation that disrupts the spatial and temporal fidelity of retinotectal transmission [20Smear M.C. Tao H.W. Staub W. Orger M.B. Gosse N.J. Liu Y. Takahashi K. Poo M.M. Baier H. Vesicular glutamate transport at a central synapse limits the acuity of visual perception in zebrafish.Neuron. 2007; 53: 65-77Abstract Full Text Full Text PDF PubMed Scopus (74) Google Scholar]. Larvae respond to prey located within the frontal region of visual space (the “reactive perceptive field” [13Bianco I.H. Kampff A.R. Engert F. Prey capture behavior evoked by simple visual stimuli in larval zebrafish.Front. Syst. Neurosci. 2011; 5: 101Crossref PubMed Scopus (161) Google Scholar]), which is represented in the anterior portion of the visuotopic tectal space map [14Stuermer C.A. Retinotopic organization of the developing retinotectal projection in the zebrafish embryo.J. Neurosci. 1988; 8: 4513-4530PubMed Google Scholar, 21Niell C.M. Smith S.J. Functional imaging reveals rapid development of visual response properties in the zebrafish tectum.Neuron. 2005; 45: 941-951Abstract Full Text Full Text PDF PubMed Scopus (168) Google Scholar]. Notably, optogenetic stimulation of the anterior-ventral OTc is sufficient to evoke convergent saccades and J-turns [22Fajardo O. Zhu P. Friedrich R.W. Control of a specific motor program by a small brain area in zebrafish.Front Neural Circuits. 2013; 7: 67Crossref PubMed Scopus (32) Google Scholar]. By contrast, projection neurons in the posterior tectum have been reported to be dispensable for prey catching [19Del Bene F. Wyart C. Robles E. Tran A. Looger L. Scott E.K. Isacoff E.Y. Baier H. Filtering of visual information in the tectum by an identified neural circuit.Science. 2010; 330: 669-673Crossref PubMed Scopus (181) Google Scholar]. In this study, we performed functional imaging in the anterior tectum of tethered larval zebrafish, while the animal engaged in virtual hunting behavior that was evoked by presentation of artificial visual cues [13Bianco I.H. Kampff A.R. Engert F. Prey capture behavior evoked by simple visual stimuli in larval zebrafish.Front. Syst. Neurosci. 2011; 5: 101Crossref PubMed Scopus (161) Google Scholar]. By systematically varying four features of the visual stimuli, we found that prey-catching behavior was selectively evoked by specific conjunctions of visual features. Unbiased clustering of visual response vectors revealed that tectal neurons show mixed selectivity for multiple stimulus features. Furthermore, we could identify cells that showed non-linear mixed feature selectivity that closely matched the stimulus tuning of hunting responses. To investigate how activation of these feature-analyzing neurons might be linked to initiation of prey-catching behavior, we compared neural activity in response trials versus non-response trials. This enabled us to uncover tectal population activity that was specifically associated with hunting responses. Assemblies of tectal neurons produced bursts of activity in advance of, or concurrent with, the initiation of behavior, were confined to the left or right tectal hemisphere and their laterality correlated with asymmetries in the oculomotor parameters of convergent saccades. Consequently, these population dynamics likely represent premotor activity controlling the release of hunting responses. In summary, by imaging neural activity at cellular resolution during naturalistic behavior, we have functionally identified circuit components that are likely to mediate the perceptual recognition of ethologically relevant stimuli and the release of an adaptive behavioral response. To monitor neural activity during the recognition of prey-like visual cues and the initiation of hunting routines, we combined a virtual hunting assay for tethered larval zebrafish [13Bianco I.H. Kampff A.R. Engert F. Prey capture behavior evoked by simple visual stimuli in larval zebrafish.Front. Syst. Neurosci. 2011; 5: 101Crossref PubMed Scopus (161) Google Scholar] with in vivo 2P functional imaging in transgenic larvae expressing a genetically encoded fluorescent calcium indicator under the control of a pan-neuronal promoter, Tg(elavl3:GCaMP5G)a4598 [23Ahrens M.B. Orger M.B. Robson D.N. Li J.M. Keller P.J. Whole-brain functional imaging at cellular resolution using light-sheet microscopy.Nat. Methods. 2013; 10: 413-420Crossref PubMed Scopus (834) Google Scholar] (Figure 1). In our assay, larval zebrafish were tethered in agarose gel but able to freely move their eyes and tail, and visual cues were projected onto a diffusive screen in front of the animal (Figures 1A and 1D). We previously showed that larvae respond to simple moving spots with hunting-associated oculomotor and locomotor behaviors [13Bianco I.H. Kampff A.R. Engert F. Prey capture behavior evoked by simple visual stimuli in larval zebrafish.Front. Syst. Neurosci. 2011; 5: 101Crossref PubMed Scopus (161) Google Scholar]. Specifically, larvae perform a convergent saccade and an orienting turn, comprising multiple unilateral tail bends directed toward the visual target. The kinematics of these behaviors closely match those observed for freely swimming fish hunting live prey. Because every hunting routine (for both free swimming and tethered larvae) commences with eye convergence, and the spontaneous rate of convergent saccades is very low, we could use eye tracking alone to determine that the animal has initiated hunting behavior in response to a visual target (Figures 1B and 1E). Furthermore, the relatively high failure rate of stimulus-evoked hunting responses (5%–10% response rate for the best stimuli) allowed us to disambiguate visually evoked activity from neural activity related to the release of a behavioral response. During the virtual hunting assay, we performed 2P calcium imaging to monitor neural activity in the optic tecta (Figure 1C). Hunting responses generated only small motion artifacts that could be corrected during post-processing (see the Experimental Procedures), and 2P imaging had no apparent detrimental impact on larval health or behavior. This approach therefore enabled us to monitor neural activity during the sensorimotor transformations linking the recognition of prey-like visual objects to the initiation of a hunting response. To examine the stimulus tuning of hunting responses, we presented a panel of moving spot stimuli that differed in terms of four stimulus features: direction, size, speed, and contrast polarity. For each feature, we tested two values, for a total of 16 unique stimuli. Specifically, moving spots could differ in direction (left-right or right-left motion), speed (fast 30°/s or slow 15°/s), size (small 3.5°, large 13.2°), or contrast polarity (a bright spot on a dark background, or a dark spot on a bright background) (see the Experimental Procedures). Stimuli were presented in the frontal portion of visual space, spanning the region where freely swimming larvae attend to live prey (approximately −60° (left) to +60° (right) [13Bianco I.H. Kampff A.R. Engert F. Prey capture behavior evoked by simple visual stimuli in larval zebrafish.Front. Syst. Neurosci. 2011; 5: 101Crossref PubMed Scopus (161) Google Scholar]). Thus, moving spots appeared at 100° to the left or right of the animal’s extended midsagittal plane and then swept 200° to the right, or left, respectively. Behavioral responses indicating the initiation of hunting routines were defined as convergent saccades in which both eyes rotated nasally (Experimental Procedures) (n = 361 events in 48 fish). Larvae responded to stimuli most frequently when they were almost directly ahead (median azimuth, −5.56° [left], Wilcoxon signed rank test versus median of 0°, p = 0.19; Figure 2A). There was no significant difference in the spatial location of targets at the time of convergent saccades for left- versus rightward-moving spots (p = 0.59, Kolmogorov-Smirnov test; Figure 2B), nor for slow- versus fast-moving spots (p = 0.06). Convergent saccades increased ocular vergence angle by 19.03° ± 0.49° (mean ± SEM), with mean vergence angle after saccade of 44.4° ± 0.43°, similar to our previous study [13Bianco I.H. Kampff A.R. Engert F. Prey capture behavior evoked by simple visual stimuli in larval zebrafish.Front. Syst. Neurosci. 2011; 5: 101Crossref PubMed Scopus (161) Google Scholar] (Figure 2C). The eye contralateral to the stimulus at the time of the saccade tended to show a larger nasal rotation and adopted a more nasal post-saccadic position (in agreement with [24Trivedi C.A. Bollmann J.H. Visually driven chaining of elementary swim patterns into a goal-directed motor sequence: a virtual reality study of zebrafish prey capture.Front Neural Circuits. 2013; 7: 86Crossref PubMed Scopus (55) Google Scholar]). For example, convergent saccades triggered by visual cues located on the right, usually involved greater rotation of the left eye (p = 9.13 × 10−9, contralateral versus ipsilateral eye position after the saccade; p = 6.88 × 10−5, contralateral versus ipsilateral change in eye position, paired t tests; Figure 2D). In summary, our behavioral assay allowed us to present visual cues to tethered larval zebrafish to evoke oculomotor responses associated with the initiation of natural hunting routines, during 2P functional imaging. The probability of evoking hunting responses varied substantially across our panel of visual stimuli. We quantified response rate (R) as the proportion of stimulus presentations that evoked a convergent saccade (Figure 2E). The most effective stimuli were spots for which polarity was inverse (dark spots) and size was large. Fast-moving large, dark spots were also more effective than otherwise identical slow-moving stimuli. These results suggest that hunting responses are sensitive to multiple stimulus features. We used logistic regression to model the relationship between response rate and the four visual features. For each type of feature, we used a binary coding scheme to represent the two feature levels (e.g., fast [1], slow [0]) such that each stimulus was described by a vector of four binary values (Figure 2E, bottom table). Using stepwise regression, we identified the model described in Figure 2F as producing the most accurate description of the data. To compare alternative models, we used a cross-validation approach in which we fit model coefficients on half the data set (training set) and assessed model predictions against the other unseen half (test data set) to estimate a cross-validated R2 (Experimental Procedures). The model in Figure 2F had a cross-validated R2 of 0.82 and indicates that hunting responses are strongly modulated by size and contrast polarity. Large stimuli increase the odds of response by 3.8-fold (given by eβ1) and dark stimuli by 6.5-fold (eβ2). In addition, the interaction term in the model indicates that, when the stimulus is both dark and large and fast, the odds of a response are increased by 2.5-fold (eβ3). We conclude that larval zebrafish respond differentially to moving visual cues as a function of multiple stimulus features and are sensitive to the coincidence of particular feature values (feature compounds). Specifically, size, contrast polarity and speed of motion interact, such that stimuli that are large, dark, and fast are most effective in triggering hunting responses. To investigate how different stimuli—and individual stimulus features—are encoded by neural activity, we performed 2P calcium imaging in the rostral portion of the optic tecta (and adjacent regions) (Figure 3A). In addition to the 16 moving spot stimuli, we included two control stimuli, which were 3 s “whole-field” light flashes at two different intensities. We imaged activity at ten to 15 dorsoventral levels and at each focal plane presented five to eight repetitions of each of the 18 stimuli, in a pseudo-random sequence, while simultaneously monitoring behavior. To characterize the visual response profiles of individual neurons, we computed a visual response vector for each cell as follows. First, imaging planes were automatically segmented to define regions-of-interest that corresponded well to single somata (Figure S1). Regions of interest (ROIs) localized to the synaptic neuropil layers of the OTc were excluded. Next, we computed the mean fluorescent calcium signal (ΔF/F) across the repeated presentations of each visual stimulus and finally concatenated these average responses to produce a visual response vector (VRV). The VRV therefore summarizes the visual responses of each neuron in the form of the full response time course to the 18 visual stimuli (684 time points per cell). To examine the diversity of visual response profiles, we used an unbiased clustering method to group visually responsive cells from 14 fish based on the similarity of their VRVs, as measured by correlation (Experimental Procedures). Our method produced 20 clusters, each of which contained cells from a minimum of six fish (Figure 3B; Figure S2; Table S1). These clusters contained neurons with more coherent visual tuning properties than we could obtain using k-means clustering. From a total of 169,371 ROIs (14 fish), our method clustered only 5,092 visually responsive cells (∼3%). This relatively small sample set is most likely not exhaustive but allowed us to identify groups of neurons with feature selective visual tuning that were found consistently across multiple fish. Notably, an alternative clustering approach based on Gaussian mixture modeling identified very similar clusters but also isolated only a relatively small number of cells (1,035 cells from 101,656 in 10 fish, ∼1%; Figure S3; Experimental Procedures). Figure 3B shows the 20 clusters identified using our correlation-based clustering approach to measure the similarity of VRVs, at a minimum correlation coefficient threshold of 0.75 (see Figure S2 and Table S1 for additional cluster details). Clusters could be broadly divided into those modulated by changes in background luminance and clusters selective for moving spots. A step increase in background luminance occurs during presentation of negative polarity (dark) moving spots (starting 2 s before spot appearance and ending 2 s after spot disappearance; Experimental Procedures), as well as during the control light-flash stimuli. The six clusters responsive to changes in background luminance (c15–20) showed a diversity of response properties and temporal dynamics. These include negative modulation (a decrease in fluorescence signal, which we presume corresponds to a decrease in tonic firing rate) in response to an increase in luminance (cluster 20): constituent cells were found in the habenulae and torus longitudinalis (TL) as well as the optic tecta (Figure 3C). Cluster 19 showed positive modulation in response to decreases in luminance (“dimming detectors”), and cluster 16, which contained the largest number of neurons of any cluster, displayed positive modulation in response to increasing whole-field luminance (“ON” response). This was evident in the response to changes in background light level during control stimuli and dark spot presentations and in response to large, bright, moving spots. A large proportion of these neurons (41%) were located in the TL, specifically at its rostral pole (Figure 4A). We identified 14 clusters that were responsive to moving spots and showed minimal modulation to changes in background luminance. Inspection of cluster centroids (the average VRV of cells within the cluster) revealed that clusters respond differentially across the panel of 16 moving spot stimuli and show direction, size, and polarity selectivity (Figure 3B; Table S1). We quantified feature tuning by computing, for each cell, four selectivity indices (for direction, speed, size, and polarity) based on the maximal mean calcium signal across the panel of 16 stimuli (Experimental Procedures). Mirror-symmetric clusters could be identified in the left and right tectal hemispheres, with similar feature tuning. For example, clusters 9–12 show size, polarity, and direction selectivity, with a net preference for large, bright spots moving either leftward or rightward (Figures 4B and 4C). Clusters 9 and 10 prefer right-left-moving spots. Despite otherwise similar tuning, clusters 9 and 10 were segregated because they respond at different times to spots sweeping across the visual field, from +100° (right) to −100° (left). The retinotectal projection is entirely crossed in larval zebrafish such that the left OTc is innervated by retinal ganglion cells deriving from the right eye and the right OTc receives input from the left eye. Accordingly, cluster 9 is exclusively located in the left OTc and responds earlier during presentation of right-left visual cues (moving tail-nose), whereas cluster 10 is confined to the right OTc and responds later, after the cue has crossed to the left visual hemifield (nose-tail motion; Figure 4B). Clusters 11 and 12 show the opposite direction selectivity (preferring left-right motion) and are similarly located in the left and right OTc, respectively (Figure 4C). Consequently, clusters 9 and 12 form a mirror-symmetric pair tuned to tail-nose motion, located on the left and right, respectively. Clusters 10 and 11 form a second pair tuned to nose-tail-moving spots. Hunting responses were evoked most frequently by large, dark, fast-moving spots. Our unbiased clustering procedure identified tectal neurons tuned to large, dark spots, which additionally showed direction selectivity. Clusters 1 and 2 show a preference for leftward-moving large, dark spots and are located in the left and right OTc, respectively (Figure 4D). Clusters 3–6 show the opposite direction selectivity, preferring rightward motion. These four clusters have similar tuning and were divided not only by tectal laterality, but also by rostrocaudal tectal location, based on the differential timing of their calcium responses (Figure 4E). In accordance with the retinotopic mapping of visual space, clusters at more caudal positions (clusters 3 and 6) responded when visual cues were at more peripheral locations. Our expectation is that other clusters (e.g., clusters 1 and 2) could be similarly subdivided if temporal resolution was higher or the correlation threshold of our clustering procedure was increased. In conclusion, we find that tectal neurons show mixed selectivity and are sensitive to combinations of visual features (feature compounds). These include direction-selective cells with a preference for large, dark, moving spots that we found to be among the most effective stimuli in evoking hunting responses. Hunting responses displayed mixed feature selectivity and were most effectively triggered by large, dark, fast-moving spots. Based on these behavioral observations, we developed an ap" @default.
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• W2071663937 title "Visuomotor Transformations Underlying Hunting Behavior in Zebrafish" @default.
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