Development and organization of the retinal orientation selectivity map.

Orientation or axial selectivity, the property of neurons in the visual system to respond preferentially to certain angles of visual stimuli, plays a pivotal role in our understanding of visual perception and information processing. This computation is performed as early as the retina, and although much work has established the cellular mechanisms of retinal orientation selectivity, how this computation is organized across the retina is unknown. Using a large dataset collected across the mouse retina, we demonstrate functional organization rules of retinal orientation selectivity. First, we identify three major functional classes of retinal cells that are orientation selective and match previous descriptions. Second, we show that one orientation is predominantly represented in the retina and that this predominant orientation changes as a function of retinal location. Third, we demonstrate that neural activity plays little role on the organization of retinal orientation selectivity. Lastly, we use in silico modeling followed by validation experiments to demonstrate that the overrepresented orientation aligns along concentric axes. These results demonstrate that, similar to direction selectivity, orientation selectivity is organized in a functional map as early as the retina.

Magnetic orientation in juvenile Atlantic herring (Clupea harengus) could involve cryptochrome 4 as a potential magnetoreceptor.

The Earth's magnetic field can provide reliable directional information, allowing migrating animals to orient themselves using a magnetic compass or estimate their position relative to a target using map-based orientation. Here we show for the first time that young, inexperienced herring (Clupea harengus, Ch) have a magnetic compass when they migrate hundreds of kilometres to their feeding grounds. In birds, such as the European robin (Erithacus rubecula), radical pair-based magnetoreception involving cryptochrome 4 (ErCRY4) was demonstrated; the molecular basis of magnetoreception in fish is still elusive. We show that cry4 expression in the eye of herring is upregulated during the migratory season, but not before, indicating a possible use for migration. The amino acid structure of herring ChCRY4 shows four tryptophans and a flavin adenine dinucleotide-binding site, a prerequisite for a magnetic receptor. Using homology modelling, we successfully reconstructed ChCRY4 of herring, DrCRY4 of zebrafish (Danio rerio) and StCRY4 of brown trout (Salmo trutta) and showed that ChCRY4, DrCRY4 and ErCRY4a, but not StCRY4, exhibit very comparable dynamic behaviour. The electron transfer could take place in ChCRY4 in a similar way to ErCRY4a. The combined behavioural, transcriptomic and simulation experiments provide evidence that CRY4 could act as a magnetoreceptor in Atlantic herring.

Tactile adaptation to orientation produces a robust tilt aftereffect and exhibits crossmodal transfer when tested in vision.

Orientation processing is one of the most fundamental functions in both visual and somatosensory perception. Converging findings suggest that orientation processing in both modalities is closely linked: somatosensory neurons share a similar orientation organisation as visual neurons, and the visual cortex has been found to be heavily involved in tactile orientation perception. Hence, we hypothesized that somatosensation would exhibit a similar orientation adaptation effect, and this adaptation effect would be transferable between the two modalities, considering the above-mentioned connection. The tilt aftereffect (TAE) is a demonstration of orientation adaptation and is used widely in behavioural experiments to investigate orientation mechanisms in vision. By testing the classic TAE paradigm in both tactile and crossmodal orientation tasks between vision and touch, we were able to show that tactile perception of orientation shows a very robust TAE, similar to its visual counterpart. We further show that orientation adaptation in touch transfers to produce a TAE when tested in vision, but not vice versa. Additionally, when examining the test sequence following adaptation for serial effects, we observed another asymmetry between the two conditions where the visual test sequence displayed a repulsive intramodal serial dependence effect while the tactile test sequence exhibited an attractive serial dependence. These findings provide concrete evidence that vision and touch engage a similar orientation processing mechanism. However, the asymmetry in the crossmodal transfer of TAE and serial dependence points to a non-reciprocal connection between the two modalities, providing further insights into the underlying processing mechanism.

Albatrosses employ orientation and routing strategies similar to yacht racers.

The way goal-oriented birds adjust their travel direction and route in response to wind significantly affects their travel costs. This is expected to be particularly pronounced in pelagic seabirds, which utilize a wind-dependent flight style called dynamic soaring. Dynamic soaring seabirds in situations without a definite goal, e.g. searching for prey, are known to preferentially fly with crosswinds or quartering-tailwinds to increase the speed and search area, and reduce travel costs. However, little is known about their reaction to wind when heading to a definite goal, such as homing. Homing tracks of wandering albatrosses (Diomedea exulans) vary from beelines to zigzags, which are similar to those of sailboats. Here, given that both albatrosses and sailboats travel slower in headwinds and tailwinds, we tested whether the time-minimizing strategies used by yacht racers can be compared to the locomotion patterns of wandering albatrosses. We predicted that when the goal is located upwind or downwind, albatrosses should deviate their travel directions from the goal on the mesoscale and increase the number of turns on the macroscale. Both hypotheses were supported by track data from albatrosses and racing yachts in the Southern Ocean confirming that albatrosses qualitatively employ the same strategy as yacht racers. Nevertheless, albatrosses did not strictly minimize their travel time, likely making their flight robust against wind fluctuations to reduce flight costs. Our study provides empirical evidence of tacking in albatrosses and demonstrates that man-made movement strategies provide a new perspective on the laws underlying wildlife movement.

Visual boundary cues suffice to anchor place and grid cells in virtual reality.

The hippocampal formation contains neurons responsive to an animal's current location and orientation, which together provide the organism with a neural map of space.1,2,3 Spatially tuned neurons rely on external landmark cues and internally generated movement information to estimate position.4,5 An important class of landmark cue are the boundaries delimiting an environment, which can define place cell field position6,7 and stabilize grid cell firing.8 However, the precise nature of the sensory information used to detect boundaries remains unknown. We used 2-dimensional virtual reality (VR)9 to show that visual cues from elevated walls surrounding the environment are both sufficient and necessary to stabilize place and grid cell responses in VR, when only visual and self-motion cues are available. By contrast, flat boundaries formed by the edges of a textured floor did not stabilize place and grid cells, indicating only specific forms of visual boundary stabilize hippocampal spatial firing. Unstable grid cells retain internally coherent, hexagonally arranged firing fields, but these fields "drift" with respect to the virtual environment over periods >5 s. Optic flow from a virtual floor does not slow drift dynamics, emphasizing the importance of boundary-related visual information. Surprisingly, place fields are more stable close to boundaries even with floor and wall cues removed, suggesting invisible boundaries are inferred using the motion of a discrete, separate cue (a beacon signaling reward location). Subsets of place cells show allocentric directional tuning toward the beacon, with strength of tuning correlating with place field stability when boundaries are removed.

Serial dependence requires visual awareness: Evidence from continuous flash suppression.

The visual system often undergoes a relatively stable perception even in a noisy visual environment. This crucial function was reflected in a visual perception phenomenon-serial dependence, in which recent stimulus history systematically biases current visual decisions. Although serial dependence effects have been revealed in numerous studies, few studies examined whether serial dependence would require visual awareness. By using the continuous flash suppression (CFS) technique to render grating stimuli invisible, we investigated whether serial dependence effects could emerge at the unconscious levels. In an orientation adjustment task, subjects viewed a randomly oriented grating and reported their orientation perception via an adjustment response. Subjects performed a series of three type trial pairs. The first two trial pairs, in which subjects were instructed to make a response or no response toward the first trial of the pairs, respectively, were used to measure serial dependence at the conscious levels; the third trial pair, in which the grating stimulus in the first trial of the pair was masked by a CFS stimulus, was used to measure the serial dependence at the unconscious levels. One-back serial dependence effects for the second trial of the pairs were evaluated. We found significant serial dependence effects at the conscious levels, whether absence (Experiment 1) or presence (Experiment 2) of CFS stimuli, but failed to find the effects at the unconscious levels, corroborating the view that serial dependence requires visual awareness.

Discriminating orientation information with phase consistency in alpha and low-gamma frequency bands: an EEG study.

Recent studies suggest that noninvasive imaging methods (EEG, MEG) in the human brain scalp can decode the content of visual features information (orientation, color, motion, etc.) in Visual-Working Memory (VWM). Previous work demonstrated that with the sustained low-frequency Event-Related Potential (ERP under 6 Hz) of scalp EEG distributions, it is possible to accurately decode the content of orientation information in VWM during the delay interval. In addition, previous studies showed that the raw data captured by a combination of the occi-parietal electrodes could be used to decode the orientation. However, it is unclear whether the orientation information is available in other frequency bands (higher than 6 Hz) or whether this information is feasible with fewer electrodes. Furthermore, the exploration of orientation information in the phase values of the signal has not been well-addressed. In this study, we propose that orientation information is also accessible through the phase consistency of the occipital region in the alpha band frequency. Our results reveal a significant difference between orientations within 200 ms after stimulus offset in early visual sensory processing, with no apparent effect in power and Event-Related Oscillation (ERO) during this period. Additionally, in later periods (420-500 ms after stimulus offset), a noticeable difference is observed in the phase consistency of low gamma-band activity in the occipital area. Importantly, our findings suggest that phase consistency between trials of the orientation feature in the occipital alpha and low gamma-band can serve as a measure to obtain orientation information in VWM. Furthermore, the study demonstrates that phase consistency in the alpha and low gamma band can reflect the distribution of orientation-selective neuron numbers in the four main orientations in the occipital area.

When experience with scenes foils attentional orienting: ERP evidence against flexible target-context mapping in visual search.

Visual search is speeded when a target is repeatedly presented in an invariant scene context of nontargets (contextual cueing), demonstrating observers' capability for using statistical long-term memory (LTM) to make predictions about upcoming sensory events, thus improving attentional orienting. In the current study, we investigated whether expectations arising from individual, learned environmental structures can encompass multiple target locations. We recorded event-related potentials (ERPs) while participants performed a contextual cueing search task with repeated and non-repeated spatial item configurations. Notably, a given search display could be associated with either a single target location (standard contextual cueing) or two possible target locations. Our result showed that LTM-guided attention was always limited to only one target position in single- but also in the dual-target displays, as evidenced by expedited reaction times (RTs) and enhanced N1pc and N2pc deflections contralateral to one ("dominant") target of up to two repeating target locations. This contrasts with the processing of non-learned ("minor") target positions (in dual-target displays), which revealed slowed RTs alongside an initial N1pc "misguidance" signal that then vanished in the subsequent N2pc. This RT slowing was accompanied by enhanced N200 and N400 waveforms over fronto-central electrodes, suggesting that control mechanisms regulate the competition between dominant and minor targets. Our study thus reveals a dissociation in processing dominant versus minor targets: While LTM templates guide attention to dominant targets, minor targets necessitate control processes to overcome the automatic bias towards previously learned, dominant target locations.

Orienting role of the putative human posterior infero-temporal area in visual attention.

The dorsal attention network (DAN) is a network of brain regions essential for attentional orienting, which includes the lateral intraparietal area (LIP) and frontal eye field (FEF). Recently, the putative human dorsal posterior infero-temporal area (phPITd) has been identified as a new node of the DAN. However, its functional relationship with other areas of the DAN and its specific role in visual attention remained unclear. In this study, we analyzed a large publicly available neuroimaging dataset to investigate the intrinsic functional connectivities (FCs) of the phPITd with other brain areas. The results showed that the intrinsic FCs of the phPITd with the areas of the visual network and the DAN were significantly stronger than those with the ventral attention network (VAN) areas and areas of other networks. We further conducted individual difference analyses with a sample size of 295 participants and a series of attentional tasks to investigate which attentional components each phPITd-based DAN edge predicts. Our findings revealed that the intrinsic FC of the left phPITd with the LIPv could predict individual ability in attentional orienting, but not in alerting, executive control, and distractor suppression. Our results not only provide direct evidence of the phPITd's functional relationship with the LIPv, but also offer a comprehensive understanding of its specific role in visual attention.

Feature variability determines specificity and transfer in multiorientation feature detection learning.

Historically, in many perceptual learning experiments, only a single stimulus is practiced, and learning is often specific to the trained feature. Our prior work has demonstrated that multi-stimulus learning (e.g., training-plus-exposure procedure) has the potential to achieve generalization. Here, we investigated two important characteristics of multi-stimulus learning, namely, roving and feature variability, and their impacts on multi-stimulus learning and generalization. We adopted a feature detection task in which an oddly oriented target bar differed by 16° from the background bars. The stimulus onset asynchrony threshold between the target and the mask was measured with a staircase procedure. Observers were trained with four target orientation search stimuli, either with a 5° deviation (30°-35°-40°-45°) or with a 45° deviation (30°-75°-120°-165°), and the four reference stimuli were presented in a roving manner. The transfer of learning to the swapped target-background orientations was evaluated after training. We found that multi-stimulus training with a 5° deviation resulted in significant learning improvement, but learning failed to transfer to the swapped target-background orientations. In contrast, training with a 45° deviation slowed learning but produced a significant generalization to swapped orientations. Furthermore, a modified training-plus-exposure procedure, in which observers were trained with four orientation search stimuli with a 5° deviation and simultaneously passively exposed to orientations with high feature variability (45° deviation), led to significant orientation learning generalization. Learning transfer also occurred when the four orientation search stimuli with a 5° deviation were presented in separate blocks. These results help us to specify the condition under which multistimuli learning produces generalization, which holds potential for real-world applications of perceptual learning, such as vision rehabilitation and expert training.

Electrodermal and central measures of the tonic orienting reflex (OR).

Sokolov described both phasic and tonic aspects of the Orienting Reflex (OR), but subsequent research and theory development has focussed primarily on the phasic OR at the expense of the tonic OR. The present study used prestimulus skin conductance level (SCL) during a dishabituation paradigm to model the tonic OR, examining its amplitude patterning over repeated standard stimulus presentations and a change stimulus. We expected sensitisation (increased amplitude) following the initial and change trials, and habituation (decrement) over the intervening trials. Prestimulus EEG alpha level was explored as a potential central measure of the tonic OR (as an inverse correlate), examining its pattern over stimulus repetition and change in relation to the SCL model. We presented a habituation series of innocuous auditory stimuli to two groups (each N = 20) at different ISIs (Long 13-15 s and Short 5-7 s) and recorded electrodermal and EEG data during two counterbalanced conditions; Indifferent: no task requirements; Significant: silent counting. Across groups and conditions, prestimulus SCLs and alpha amplitudes generally showed the expected trials patterns, confirming our main hypotheses. Findings have important implications for including the assessment of Sokolov's tonic OR in modelling central and autonomic nervous system interactions of fundamental attention and learning processes.

Neurotoxic effects of home radon exposure on oscillatory dynamics serving attentional orienting in children and adolescents.

Radon is a naturally occurring gas that contributes significantly to radiation in the environment and is the second leading cause of lung cancer globally. Previous studies have shown that other environmental toxins have deleterious effects on brain development, though radon has not been studied as thoroughly in this context. This study examined the impact of home radon exposure on the neural oscillatory activity serving attention reorientation in youths. Fifty-six participants (ages 6-14 years) completed a classic Posner cuing task during magnetoencephalography (MEG), and home radon levels were measured for each participant. Time-frequency spectrograms indicated stronger theta (3-7 Hz, 300-800 ms), alpha (9-13 Hz, 400-900 ms), and beta responses (14-24 Hz, 400-900 ms) during the task relative to baseline. Source reconstruction of each significant oscillatory response was performed, and validity maps were computed by subtracting the task conditions (invalidly cued - validly cued). These validity maps were examined for associations with radon exposure, age, and their interaction in a linear regression design. Children with greater radon exposure showed aberrant oscillatory activity across distributed regions critical for attentional processing and attention reorientation (e.g., dorsolateral prefrontal cortex, and anterior cingulate cortex). Generally, youths with greater radon exposure exhibited a reverse neural validity effect in almost all regions and showed greater overall power relative to peers with lesser radon exposure. We also detected an interactive effect between radon exposure and age where youths with greater radon exposure exhibited divergent developmental trajectories in neural substrates implicated in attentional processing (e.g., bilateral prefrontal cortices, superior temporal gyri, and inferior parietal lobules). These data suggest aberrant, but potentially compensatory neural processing as a function of increasing home radon exposure in areas critical for attention and higher order cognition.

Brain parasites and misorientation of migratory birds.

Haemoproteus blood parasites of birds are thought to be relatively benign. Recent findings show that infections may develop in the brain of birds, possibly distorting their orientation sense. Misdirected migration may lead migrants outside their range where they are recognized as vagrants and can transmit parasites to novel hosts.

Suppression on the basis of template for rejection is reactive: Evidence from human electrophysiology.

According to most theories of attention, the selection of task-relevant visual information can be enhanced by holding them in visual working memory (VWM). However, there has been a long-standing debate concerning whether similar optimization can also be achieved for task-irrelevant information, known as a "template for rejection". The present study aimed to explore this issue by examining the consequence of cue distractors before visual search tasks. For this endeavor, we manipulated the display heterogeneity by using two distractor conditions, salient and non-salient, to explore the extent to which holding the distractor color in VWM might affect attentional selection. We measured the reaction times of participants while their EEG activity was recorded. The results showed that WM-matched distractors did not improve reaction times but rather slowed them down in both tasks. Event-related potential (ERP) results showed that the display heterogeneity had no modulatory effect on the degree of distractor suppression. Even in the salient distractor condition, the WM-matched distractor received no greater suppression. Furthermore, the WM-matched distractor but not the neutral distractor elicited an N2pc before the PD in salient distractor conditions. This suggests that the template for rejection operates reactively since suppression occurs after extra attentional processes to the distractor. Moreover, the presence of WM-matched distractors led to a reduction of P3b, indicating a competition between target processing and WM-matched distractor rejection. Our findings provide insights into the mechanisms underlying the optimization of attentional selection, and have implications for future studies aimed at understanding the role of VWM in cognition.

Sense of direction in vestibular disorders.

BACKGROUND: Our sense of direction (SOD) ability relies on the sensory integration of both visual information and self-motion cues from the proprioceptive and vestibular systems. Here, we assess how dysfunction of the vestibular system impacts perceived SOD in varying vestibular disorders, and secondly, we explore the effects of dizziness, migraine and psychological symptoms on SOD ability in patient and control groups. METHODS: 87 patients with vestibular disorder and 69 control subjects were assessed with validated symptom and SOD questionnaires (Santa Barbara Sense of Direction scale and the Object Perspective test). RESULTS: While patients with vestibular disorders performed significantly worse than controls at the group level, only central and functional disorders (vestibular migraine and persistent postural perceptual dizziness), not peripheral disorders (benign-paroxysmal positional vertigo, bilateral vestibular failure and Meniere's disease) showed significant differences compared to controls on the level of individual vestibular groups. Additionally, orientational abilities associated strongly with spatial anxiety and showed clear separation from general dizziness and psychological factors in both patient and control groups. CONCLUSIONS: SOD appears to be less affected by peripheral vestibular dysfunction than by functional and/or central diagnoses, indicating that higher level disruptions to central vestibular processing networks may impact SOD more than reductions in sensory peripheral inputs. Additionally, spatial anxiety is highly associated with orientational abilities in both patients and control subjects.

Adaptation-induced sharpening of orientation tuning curves in the mouse visual cortex.

OBJECTIVE: Orientation selectivity is an emergent property of visual neurons across species with columnar and noncolumnar organization of the visual cortex. The emergence of orientation selectivity is more established in columnar cortical areas than in noncolumnar ones. Thus, how does orientation selectivity emerge in noncolumnar cortical areas after an adaptation protocol? Adaptation refers to the constant presentation of a nonoptimal stimulus (adapter) to a neuron under observation for a specific time. Previously, it had been shown that adaptation has varying effects on the tuning properties of neurons, such as orientation, spatial frequency, motion and so on. BASIC METHODS: We recorded the mouse primary visual neurons (V1) at different orientations in the control (preadaptation) condition. This was followed by adapting neurons uninterruptedly for 12 min and then recording the same neurons postadaptation. An orientation selectivity index (OSI) for neurons was computed to compare them pre- and post-adaptation. MAIN RESULTS: We show that 12-min adaptation increases the OSI of visual neurons ( n  = 113), that is, sharpens their tuning. Moreover, the OSI postadaptation increases linearly as a function of the OSI preadaptation. CONCLUSION: The increased OSI postadaptation may result from a specific dendritic neural mechanism, potentially facilitating the rapid learning of novel features.

Involvement of the superior colliculi in crossmodal correspondences.

There is an increasing body of evidence suggesting that there are low-level perceptual processes involved in crossmodal correspondences. In this study, we investigate the involvement of the superior colliculi in three basic crossmodal correspondences: elevation/pitch, lightness/pitch, and size/pitch. Using a psychophysical design, we modulate visual input to the superior colliculus to test whether the superior colliculus is required for behavioural crossmodal congruency effects to manifest in an unspeeded multisensory discrimination task. In the elevation/pitch task, superior colliculus involvement is required for a behavioural elevation/pitch congruency effect to manifest in the task. In the lightness/pitch and size/pitch task, we observed a behavioural elevation/pitch congruency effect regardless of superior colliculus involvement. These results suggest that the elevation/pitch correspondence may be processed differently to other low-level crossmodal correspondences. The implications of a distributed model of crossmodal correspondence processing in the brain are discussed.

Auditory Processing of Intonational Rises and Falls in German: Rises Are Special in Attention Orienting.

This article investigates the processing of intonational rises and falls when presented unexpectedly in a stream of repetitive auditory stimuli. It examines the neurophysiological correlates (ERPs) of attention to these unexpected stimuli through the use of an oddball paradigm where sequences of repetitive stimuli are occasionally interspersed with a deviant stimulus, allowing for elicitation of an MMN. Whereas previous oddball studies on attention toward unexpected sounds involving pitch rises were conducted on nonlinguistic stimuli, the present study uses as stimuli lexical items in German with naturalistic intonation contours. Results indicate that rising intonation plays a special role in attention orienting at a pre-attentive processing stage, whereas contextual meaning (here a list of items) is essential for activating attentional resources at a conscious processing stage. This is reflected in the activation of distinct brain responses: Rising intonation evokes the largest MMN, whereas falling intonation elicits a less pronounced MMN followed by a P3 (reflecting a conscious processing stage). Subsequently, we also find a complex interplay between the phonological status (i.e., accent/head marking vs. boundary/edge marking) and the direction of pitch change in their contribution to attention orienting: Attention is not oriented necessarily toward a specific position in prosodic structure (head or edge). Rather, we find that the intonation contour itself and the appropriateness of the contour in the linguistic context are the primary cues to two core mechanisms of attention orienting, pre-attentive and conscious orientation respectively, whereas the phonological status of the pitch event plays only a supplementary role.

Molecular signatures of attention networks.

Attention network theory proposes three distinct types of attention-alerting, orienting, and control-that are supported by separate brain networks and modulated by different neurotransmitters, that is, norepinephrine, acetylcholine, and dopamine. Here, we explore the extent of cortical, genetic, and molecular dissociation of these three attention systems using multimodal neuroimaging. We evaluated the spatial overlap between fMRI activation maps from the attention network test (ANT) and cortex-wide gene expression data from the Allen Human Brain Atlas. The goal was to identify genes associated with each of the attention networks in order to determine whether specific groups of genes were co-expressed with the corresponding attention networks. Furthermore, we analyzed publicly available PET-maps of neurotransmitter receptors and transporters to investigate their spatial overlap with the attention networks. Our analyses revealed a substantial number of genes (3871 for alerting, 6905 for orienting, 2556 for control) whose cortex-wide expression co-varied with the activation maps, prioritizing several molecular functions such as the regulation of protein biosynthesis, phosphorylation, and receptor binding. Contrary to the hypothesized associations, the ANT activation maps neither aligned with the distribution of norepinephrine, acetylcholine, and dopamine receptor and transporter molecules, nor with transcriptomic profiles that would suggest clearly separable networks. Independence of the attention networks appeared additionally constrained by a high level of spatial dependency between the network maps. Future work may need to reconceptualize the attention networks in terms of their segregation and reevaluate the presumed independence at the neural and neurochemical level.

The Anterior Thalamus Preferentially Drives Allocentric But Not Egocentric Orientation Tuning in Postrhinal Cortex.

Navigating a complex world requires integration of multiple spatial reference frames, including information about one's orientation in both allocentric and egocentric coordinates. Combining these two information sources can provide additional information about one's spatial location. Previous studies have demonstrated that both egocentric and allocentric spatial signals are reflected by the firing of neurons in the rat postrhinal cortex (POR), an area that may serve as a hub for integrating allocentric head direction (HD) cell information with egocentric information from center-bearing and center-distance cells. However, we have also demonstrated that POR HD cells are uniquely influenced by the visual properties and locations of visual landmarks, bringing into question whether the POR HD signal is truly allocentric as opposed to simply being a response to visual stimuli. To investigate this issue, we recorded HD cells from the POR of female rats while bilaterally inactivating the anterior thalamus (ATN), a region critical for expression of the "classic" HD signal in cortical areas. We found that ATN inactivation led to a significant decrease in both firing rate and tuning strength for POR HD cells, as well as a disruption in the encoding of allocentric location by conjunctive HD/egocentric cells. In contrast, POR egocentric cells without HD tuning were largely unaffected in a consistent manner by ATN inactivation. These results indicate that the POR HD signal originates at least partially from projections from the ATN and supports the view that the POR acts as a hub for the integration of egocentric and allocentric spatial representations.