A Fine-Scale Functional Logic to Convergence from Retina to Thalamus.

Numerous well-defined classes of retinal ganglion cells innervate the thalamus to guide image-forming vision, yet the rules governing their convergence and divergence remain unknown. Using two-photon calcium imaging in awake mouse thalamus, we observed a functional arrangement of retinal ganglion cell axonal boutons in which coarse-scale retinotopic ordering gives way to fine-scale organization based on shared preferences for other visual features. Specifically, at the ∼6 µm scale, clusters of boutons from different axons often showed similar preferences for either one or multiple features, including axis and direction of motion, spatial frequency, and changes in luminance. Conversely, individual axons could "de-multiplex" information channels by participating in multiple, functionally distinct bouton clusters. Finally, ultrastructural analyses demonstrated that retinal axonal boutons in a local cluster often target the same dendritic domain. These data suggest that functionally specific convergence and divergence of retinal axons may impart diverse, robust, and often novel feature selectivity to visual thalamus.

Dysfunction of the magnocellular subdivision of the visual thalamus in developmental dyslexia.

Developmental dyslexia (DD) is one of the most common learning disorders, affecting millions of children and adults worldwide. To date, scientific research has attempted to explain DD primarily based on pathophysiological alterations in the cerebral cortex. In contrast, several decades ago, pioneering research on five post-mortem human brains suggested that a core characteristic of DD might be morphological alterations in a specific subdivision of the visual thalamus - the magnocellular LGN (M-LGN). However, due to considerable technical challenges in investigating LGN subdivisions non-invasively in humans, this finding was never confirmed in-vivo, and its relevance for DD pathology remained highly controversial. Here, we leveraged recent advances in high-resolution magnetic resonance imaging (MRI) at high field strength (7 Tesla) to investigate the M-LGN in DD in-vivo. Using a case-control design, we acquired data from a large sample of young adults with DD (n = 26; age 28 ± 7 years; 13 females) and matched control participants (n = 28; age 27 ± 6 years; 15 females). Each participant completed a comprehensive diagnostic behavioral test battery and participated in two MRI sessions, including three functional MRI experiments and one structural MRI acquisition. We measured blood-oxygen-level-dependent responses and longitudinal relaxation rates to compare both groups on LGN subdivision function and myelination. Based on previous research, we hypothesized that the M-LGN is altered in DD and that these alterations are associated with a key DD diagnostic score, i.e., rapid letter and number naming (RANln). The results showed aberrant responses of the M-LGN in DD compared to controls, which was reflected in a different functional lateralization of this subdivision between groups. These alterations were associated with RANln performance, specifically in male DD. We also found lateralization differences in the longitudinal relaxation rates of the M-LGN in DD relative to controls. Conversely, the other main subdivision of the LGN, the parvocellular LGN (P-LGN), showed comparable blood-oxygen-level-dependent responses and longitudinal relaxation rates between groups. The present study is the first to unequivocally show that M-LGN alterations are a hallmark of DD, affecting both the function and microstructure of this subdivision. It further provides a first functional interpretation of M-LGN alterations and a basis for a better understanding of sex-specific differences in DD with implications for prospective diagnostic and treatment strategies.

The emergence of the EEG dominant rhythm across the first year of life.

The spectral composition of EEG provides important information on the function of the developing brain. For example, the frequency of the dominant rhythm, a salient features of EEG data, increases from infancy to adulthood. Changes of the dominant rhythm during infancy are yet to be fully characterized, in terms of their developmental trajectory and spectral characteristics. In this study, the development of dominant rhythm frequency was examined during a novel sustained attention task across 6-month-old (n = 39), 9-month-old (n = 30), and 12-month-old (n = 28) infants. During this task, computer-generated objects and faces floated down a computer screen for 10 s after a 5-second fixation cross. The peak frequency in the range between 5 and 9 Hz was calculated using center of gravity (CoG) and examined in response to faces and objects. Results indicated that peak frequency increased from 6 to 9 to 12 months of age in face and object conditions. We replicated the same result for the baseline. There was high reliability between the CoGs in the face, object, and baseline conditions across all channels. The developmental increase in CoG was more reliable than measures of mode frequency across different conditions. These findings suggest that CoG is a robust index of brain development across infancy.

Sound Improves Neuronal Encoding of Visual Stimuli in Mouse Primary Visual Cortex.

In everyday life, we integrate visual and auditory information in routine tasks such as navigation and communication. While concurrent sound can improve visual perception, the neuronal correlates of audiovisual integration are not fully understood. Specifically, it remains unclear whether neuronal firing patters in the primary visual cortex (V1) of awake animals demonstrate similar sound-induced improvement in visual discriminability. Furthermore, presentation of sound is associated with movement in the subjects, but little is understood about whether and how sound-associated movement affects audiovisual integration in V1. Here, we investigated how sound and movement interact to modulate V1 visual responses in awake, head-fixed mice and whether this interaction improves neuronal encoding of the visual stimulus. We presented visual drifting gratings with and without simultaneous auditory white noise to awake mice while recording mouse movement and V1 neuronal activity. Sound modulated activity of 80% of light-responsive neurons, with 95% of neurons increasing activity when the auditory stimulus was present. A generalized linear model (GLM) revealed that sound and movement had distinct and complementary effects of the neuronal visual responses. Furthermore, decoding of the visual stimulus from the neuronal activity was improved with sound, an effect that persisted even when controlling for movement. These results demonstrate that sound and movement modulate visual responses in complementary ways, improving neuronal representation of the visual stimulus. This study clarifies the role of movement as a potential confound in neuronal audiovisual responses and expands our knowledge of how multimodal processing is mediated at a neuronal level in the awake brain.SIGNIFICANCE STATEMENT Sound and movement are both known to modulate visual responses in the primary visual cortex; however, sound-induced movement has largely remained unaccounted for as a potential confound in audiovisual studies in awake animals. Here, authors found that sound and movement both modulate visual responses in an important visual brain area, the primary visual cortex, in distinct, yet complementary ways. Furthermore, sound improved encoding of the visual stimulus even when accounting for movement. This study reconciles contrasting theories on the mechanism underlying audiovisual integration and asserts the primary visual cortex as a key brain region participating in tripartite sensory interactions.

Brain State-Dependent Modulation of Thalamic Visual Processing by Cortico-Thalamic Feedback.

The behavioral state of a mammal impacts how the brain responds to visual stimuli as early as in the dorsolateral geniculate nucleus of the thalamus (dLGN), the primary relay of visual information to the cortex. A clear example of this is the markedly stronger response of dLGN neurons to higher temporal frequencies of the visual stimulus in alert as compared with quiescent animals. The dLGN receives strong feedback from the visual cortex, yet whether this feedback contributes to these state-dependent responses to visual stimuli is poorly understood. Here, we show that in male and female mice, silencing cortico-thalamic feedback profoundly reduces state-dependent differences in the response of dLGN neurons to visual stimuli. This holds true for dLGN responses to both temporal and spatial features of the visual stimulus. These results reveal that the state-dependent shift of the response to visual stimuli in an early stage of visual processing depends on cortico-thalamic feedback.SIGNIFICANCE STATEMENT Brain state affects even the earliest stages of sensory processing. A clear example of this phenomenon is the change in thalamic responses to visual stimuli depending on whether the animal's brain is in an alert or quiescent state. Despite the radical impact that brain state has on sensory processing, the underlying circuits are still poorly understood. Here, we show that both the temporal and spatial response properties of thalamic neurons to visual stimuli depend on the state of the animal and, crucially, that this state-dependent shift relies on the feedback projection from visual cortex to thalamus.

Viewpoint independent face recognition via extrapolation in paper wasps.

Visual recognition of three-dimensional signals, like faces, is challenging because the signals appear different from different viewpoints. A flexible, but cognitively challenging solution is viewpoint independent recognition, where receivers identify signals from novel viewing angles. Here, we use same/different concept learning to test viewpoint independent face recognition in Polistes fuscatus, a wasp that uses facial patterns to individually identify conspecifics. We find that wasps use extrapolation to identify novel views of conspecific faces. For example, wasps identify a pair of pictures of the same wasp as the 'same', even if the pictures are taken from different views (e.g. one face 0° rotation, one face 60° rotation). This result is notable because it provides the first evidence of view invariant recognition via extrapolation in an invertebrate. The results suggest that viewpoint independent recognition via extrapolation may be a widespread strategy to facilitate individual face recognition.

Individual prediction tendencies do not generalize across modalities.

Predictive processing theories, which model the brain as a "prediction machine", explain a wide range of cognitive functions, including learning, perception and action. Furthermore, it is increasingly accepted that aberrant prediction tendencies play a crucial role in psychiatric disorders. Given this explanatory value for clinical psychiatry, prediction tendencies are often implicitly conceptualized as individual traits or as tendencies that generalize across situations. As this has not yet explicitly been shown, in the current study, we quantify to what extent the individual tendency to anticipate sensory features of high probability generalizes across modalities. Using magnetoencephalography (MEG), we recorded brain activity while participants were presented with a sequence of four different (either visual or auditory) stimuli, which changed according to predefined transitional probabilities of two entropy levels: ordered vs. random. Our results show that, on a group-level, under conditions of low entropy, stimulus features of high probability are preactivated in the auditory but not in the visual modality. Crucially, the magnitude of the individual tendency to predict sensory events seems not to correlate between the two modalities. Furthermore, reliability statistics indicate poor internal consistency, suggesting that the measures from the different modalities are unlikely to reflect a single, common cognitive process. In sum, our findings suggest that quantification and interpretation of individual prediction tendencies cannot be generalized across modalities.

Haptic and visuo-haptic impairments for object recognition in children with autism spectrum disorder: focus on the sensory and multisensory processing dysfunctions.

Dysfunctions in sensory processing are widely described in individuals with autism spectrum disorder (ASD), although little is known about the developmental course and the impact of these difficulties on the learning processes during the preschool and school ages of ASD children. Specifically, as regards the interplay between visual and haptic information in ASD during developmental age, knowledge is very scarce and controversial. In this study, we investigated unimodal (visual and haptic) and cross-modal (visuo-haptic) processing skills aimed at object recognition through a behavioural paradigm already used in children with typical development (TD), with cerebral palsy and with peripheral visual impairments. Thirty-five children with ASD (age range: 5-11 years) and thirty-five age-matched and gender-matched typically developing peers were recruited. The procedure required participants to perform an object-recognition task relying on only the visual modality (black-and-white photographs), only the haptic modality (manipulation of real objects) and visuo-haptic transfer of these two types of information. Results are consistent with the idea that visuo-haptic transfer may be significantly worse in ASD children than in TD peers, leading to significant impairment in multisensory interactions for object recognition facilitation. Furthermore, ASD children tended to show a specific deficit in haptic information processing, while a similar trend of maturation of visual modality between the two groups is reported. This study adds to the current literature by suggesting that ASD differences in multisensory processes also regard visuo-haptic abilities necessary to identify and recognise objects of daily life.

Effects of general anaesthesia in early childhood on local and global visual processing: a post hoc analysis of the APEX cohort study.

BACKGROUND: Preclinical studies suggest that early exposure to anaesthesia alters the visual system in mice and non-human primates. We investigated whether exposure to general anaesthesia leads to visual attention processing changes in children, which could potentially impact essential life skills, including learning. METHODS: This was a post hoc analysis of data from the APprentissages EXécutifs et cerveau chez les enfants d'âge scolaire (APEX) cohort study. A total of 24 healthy 9-10-yr-old children who were or were not exposed to general anaesthesia (for surgery) by a mean age of 3.8 (2.6) yr performed a visual attention task to evaluate ability to process either local details or general global visual information. Whether children were distracted by visual interference during global and local information processing was also assessed. RESULTS: Participants included in the analyses (n=12 participants exposed to general anaesthesia and n=12 controls) successfully completed (>90% of correct answers) the trial tasks. Children from both groups were equally distracted by visual interference. However, children who had been exposed to general anaesthesia were more attracted to global visual information than were control children (P=0.03). CONCLUSIONS: These findings suggest lasting effects of early-life exposure to general anaesthesia on visuospatial abilities. Further investigations of the mechanisms by which general anaesthesia could have delayed effects on how children perceive their visual environment are needed.

New methods needed to investigate the potential adverse effects of anaesthesia on neurological development in childhood.

The issue of potentially harmful effects of neurotoxicity or anaesthesia management on children undergoing general anaesthesia is still not resolved. Studies have so far been limited by methodological problems. In a retrospective cohort study, a new noninvasive method was used to demonstrate visual processing changes in children with a single previous exposure to anaesthesia. We need new noninvasive methods that can be used before and after exposure to anaesthesia and surgery to detemine possible effects on long-term neurodevelopment.

Visual attention and processing function in relation to executive functioning in very preterm-born children aged 3 years: a prospective cohort study.

The ability to orient attention to one's environment is a prerequisite for developing executive functions (EF) from preschool age. Very preterm children are vulnerable for delays in visual orienting function (VOF) and EF deficits. This study aimed to investigate associations between objective VOF and subjective parent-reported EF in very preterm-born children at 3 years corrected age (CA). In a prospective cohort study (BOND) involving 90 children born < 30 weeks, VOF and EF were assessed using an eye tracking-based method and BRIEF-P questionnaire. Associations between abnormal VOF (viewing reaction times) and EF scores (BRIEF-P scores) were studied using multivariable regression. Using a modified Delphi method, a subset of eight items related to VOF and EF was explored. Abnormal VOF was observed in 31% of the children and abnormal global EF composite scores in 41%. Abnormal VOF was not associated with global or domain-level EF scores. However, children with abnormal VOF more frequently had EF problems related to attentional behavior: "easily sidetracked" (OR 4.18 (CI: 1.21-14.41), p = 0.02) and "short attention span" (OR 4.52 (CI: 1.34-15.22), p = 0.02). Conclusion: Although abnormal VOF was not associated with global, parent-reported, EF at 3 years CA, secondary analyses did show a relation to specific attention and concentration span items. Further research is needed to study the role of VOF in objectifying preschool EF assessments in very preterm born children. What is known: • Very preterm children are vulnerable for executive function (EF) deficits and for delays in visual orienting function, as compared to term-born children • The ability to orient attention to one's visual environment is an important prerequisite for EF early in life and is easy to measure objectively with visual orienting function (VOF) assessment What is new: • VOF measurement at 3 years of age relates to parent-reported attention and concentration span items, important aspects of executive functions • Future work should explore the clinical additional value of early VOF measurement in children at risk for EF deficits.

Subjective visual vertical/horizontal and video head impulse test in dyslexic children.

Different studies have tried to establish a relationship between dyslexia and the vestibular system function. Subjective Visual Vertical/Horizontal (SVV and SVH) and Video Head Impulse Test (VHIT) are useful for studying the vestibular system and can be easily performed in children. Our aim was to evaluate the vestibular function in dyslexic children by SVV/SVH and VHIT. We enrolled 18 dyslexic children (10M/8F; mean age 10.7 ± 2.3 years; range 7-14 years) and 18 age-matched children with typical development of learning abilities. All children performed VHIT, SVV and SVH. We found normal gain and symmetry of vestibulo-ocular-reflex both in dyslexic and typically developing children. Fifteen out of 18 dyslexic children (83.3%) showed a difference of at least one amongst SVV or SVH. The mean value of SVV was 2.3° and the mean value of SVH was 2.6°. Statistical analysis showed a significant difference between typically developing and dyslexic children for both SVV and SVH. We confirm a relationship between dyslexia and the alteration of SVV and SVH. Our results could be related to the pathogenetic hypothesis of a visual processing impairment related to a dysfunction of the magnocellular pathway or to a general deficit related to a multimodal cortical network.

A hierarchy of visual processing deficits in body dysmorphic disorder: a conceptual review and empirical investigation.

OBJECTIVE: Abnormal visual processing has been proposed as a mechanism underlying excessive focus on minor appearance flaws in body dysmorphic disorder (BDD). Existing BDD research has not differentiated the various stages of face processing (featural, first-order configural, holistic and second-order configural) that are required for higher-order processes such as emotion recognition. This study investigated a hierarchical visual processing model to examine the nature of abnormalities in face processing in BDD. METHOD: Thirty BDD participants and 27 healthy controls completed the Navon task, a featural and configural face processing task and a facial emotion labelling task. RESULTS: BDD participants performed similarly to controls when processing global and local non-face stimuli on the Navon task, when detecting subtle changes in the features and spacing of a target face, and when labelling emotional faces. However, BDD participants displayed poorer performance when viewing inverted faces, indicating difficulties in configural processing. CONCLUSIONS: The findings only partially support prior work. However, synthesis of results with previous findings indicates that heterogenous task methodologies may contribute to inconsistent findings. Recommendations are provided regarding the task parameters that appear most sensitive to abnormalities in BDD.

Visual-Motor Processing Speed and Reaction Time Differences between Medically-At-Risk Drivers and Healthy Controls.

This cross-sectional study compared visual-motor processing speed and reaction times between medically-at-risk drivers and normal controls to determine if the time in seconds distinguished between drivers who pass, fail, or need restrictions based on a road test. The medically-at-risk drivers' data (N = 35, 28-89 years) were collected as part of a comprehensive driving evaluation and coded by diagnosis (e.g., cognitive, neurological, medical) and driving outcome. The healthy control (N = 121, 21-79 years) data were collected in previous studies. The Vision Coach™ Full Field 60 task was used to collect reaction times in seconds between the two groups. Independent t-tests showed a significant difference (p < .001) in trial times between healthy controls and medically-at-risk adults. No significant difference (p = .141) was found between the three diagnoses groups. The resulting scores from the Vision Coach™ demonstrated a significant different (p < .001) between those who were determined fit to drive without restrictions and those who were determined not fit to drive after a comprehensive driving evaluation, showing the potential to be used as a screening tool for determining driving risk.

Brief Stimuli Cast a Persistent Long-Term Trace in Visual Cortex.

Visual processing is strongly influenced by recent stimulus history, a phenomenon termed adaptation. Prominent theories cast adaptation as a consequence of optimized encoding of visual information by exploiting the temporal statistics of the world. However, this would require the visual system to track the history of individual briefly experienced events, within a stream of visual input, to build up statistical representations over longer timescales. Here, using an openly available dataset from the Allen Brain Observatory, we show that neurons in the early visual cortex of the mouse indeed maintain long-term traces of individual past stimuli that persist despite the presentation of several intervening stimuli, leading to long-term and stimulus-specific adaptation over dozens of seconds. Long-term adaptation was selectively expressed in cortical, but not in thalamic, neurons, which only showed short-term adaptation. Early visual cortex thus maintains concurrent stimulus-specific memory traces of past input, enabling the visual system to build up a statistical representation of the world to optimize the encoding of new information in a changing environment.SIGNIFICANCE STATEMENT In the natural world, previous sensory input is predictive of current input over multisecond timescales. The visual system could exploit these predictabilities by adapting current visual processing to the long-term history of visual input. However, it is unclear whether the visual system can track the history of individual briefly experienced images, within a stream of input, to build up statistical representations over such long timescales. Here, we show that neurons in early visual cortex of the mouse brain exhibit remarkably long-term adaptation to brief stimuli, persisting over dozens of seconds, and despite the presentation of several intervening stimuli. The visual cortex thus maintains long-term traces of individual briefly experienced past images, enabling the formation of statistical representations over extended timescales.

Optimal parameters for rapid (invisible) frequency tagging using MEG.

Frequency tagging has been demonstrated to be a useful tool for identifying representational-specific neuronal activity in the auditory and visual domains. However, the slow flicker (<30 Hz) applied in conventional frequency tagging studies is highly visible and might entrain endogenous neuronal oscillations. Hence, stimulation at faster frequencies that is much less visible and does not interfere with endogenous brain oscillatory activity is a promising new tool. In this study, we set out to examine the optimal stimulation parameters of rapid frequency tagging (RFT/RIFT) with magnetoencephalography (MEG) by quantifying the effects of stimulation frequency, size and position of the flickering patch. Rapid frequency tagging using flickers above 50 Hz results in almost invisible stimulation which does not interfere with slower endogenous oscillations; however, the signal is weaker as compared to tagging at slower frequencies so certainty over the optimal parameters of stimulation delivery are crucial. The here presented results examining the frequency range between 60 Hz and 96 Hz suggest that RFT induces brain responses with decreasing strength up to about 84 Hz. In addition, even at the smallest flicker patch (2°) focally presented RFT induces a significant and measurable oscillatory brain signal (steady state visual evoked potential/field, SSVEP/F) at the stimulation frequency (66 Hz); however, the elicited response increases with patch size. While focal RFT presentation elicits the strongest response, off-centre presentations do generally mainly elicit a measureable response if presented below the horizontal midline. Importantly, the results also revealed considerable individual differences in the neuronal responses to RFT stimulation. Finally, we discuss the comparison of oscillatory measures (coherence and power) and sensor types (planar gradiometers and magnetometers) in order to achieve optimal outcomes. Based on our extensive findings we set forward concrete recommendations for using rapid frequency tagging in human cognitive neuroscience investigations.

Visual processing and collective motion-related decision-making in desert locusts.

Collectively moving groups of animals rely on the decision-making of locally interacting individuals in order to maintain swarm cohesion. However, the complex and noisy visual environment poses a major challenge to the extraction and processing of relevant information. We addressed this challenge by studying swarming-related decision-making in desert locust last-instar nymphs. Controlled visual stimuli, in the form of random dot kinematograms, were presented to tethered locust nymphs in a trackball set-up, while monitoring movement trajectory and walking parameters. In a complementary set of experiments, the neurophysiological basis of the observed behavioural responses was explored. Our results suggest that locusts use filtering and discrimination upon encountering multiple stimuli simultaneously. Specifically, we show that locusts are sensitive to differences in speed at the individual conspecific level, and to movement coherence at the group level, and may use these to filter out non-relevant stimuli. The locusts also discriminate and assign different weights to different stimuli, with an observed interactive effect of stimulus size, relative abundance and motion direction. Our findings provide insights into the cognitive abilities of locusts in the domain of decision-making and visual-based collective motion, and support locusts as a model for investigating sensory-motor integration and motion-related decision-making in the intricate swarm environment.

Evolution of the visual system in ray-finned fishes.

The vertebrate eye allows to capture an enormous amount of detail about the surrounding world which can only be exploited with sophisticated central information processing. Furthermore, vision is an active process due to head and eye movements that enables the animal to change the gaze and actively select objects to investigate in detail. The entire system requires a coordinated coevolution of its parts to work properly. Ray-finned fishes offer a unique opportunity to study the evolution of the visual system due to the high diversity in all of its parts. Here, we are bringing together information on retinal specializations (fovea), central visual centers (brain morphology studies), and eye movements in a large number of ray-finned fishes in a cladistic framework. The nucleus glomerulosus-inferior lobe system is well developed only in Acanthopterygii. A fovea, independent eye movements, and an enlargement of the nucleus glomerulosus-inferior lobe system coevolved at least five times independently within Acanthopterygii. This suggests that the nucleus glomerulosus-inferior lobe system is involved in advanced object recognition which is especially well developed in association with a fovea and independent eye movements. None of the non-Acanthopterygii have a fovea (except for some deep sea fish) or independent eye movements and they also lack important parts of the glomerulosus-inferior lobe system. This suggests that structures for advanced visual object recognition evolved within ray-finned fishes independent of the ones in tetrapods and non-ray-finned fishes as a result of a coevolution of retinal, central, and oculomotor structures.

Preliminary evidence of an increased susceptibility to face pareidolia in postpartum women.

As primates, we are hypersensitive to faces and face-like patterns in the visual environment, hence we often perceive illusory faces in otherwise inanimate objects, such as burnt pieces of toast and the surface of the moon. Although this phenomenon, known as face pareidolia, is a common experience, it is unknown whether our susceptibility to face pareidolia is static across our lifespan or what factors would cause it to change. Given the evidence that behaviour towards face stimuli is modulated by the neuropeptide oxytocin (OT), we reasoned that participants in stages of life associated with high levels of endogenous OT might be more susceptible to face pareidolia than participants in other stages of life. We tested this hypothesis by assessing pareidolia susceptibility in two groups of women; pregnant women (low endogenous OT) and postpartum women (high endogenous OT). We found evidence that postpartum women report seeing face pareidolia more easily than women who are currently pregnant. These data, collected online, suggest that our sensitivity to face-like patterns is not fixed and may change throughout adulthood, providing a crucial proof of concept that requires further research.

Higher order visual areas enhance stimulus responsiveness in mouse primary visual cortex.

Over the past few years, the various areas that surround the primary visual cortex (V1) in the mouse have been associated with many functions, ranging from higher order visual processing to decision-making. Recently, some studies have shown that higher order visual areas influence the activity of the primary visual cortex, refining its processing capabilities. Here, we studied how in vivo optogenetic inactivation of two higher order visual areas with different functional properties affects responses evoked by moving bars in the primary visual cortex. In contrast with the prevailing view, our results demonstrate that distinct higher order visual areas similarly modulate early visual processing. In particular, these areas enhance stimulus responsiveness in the primary visual cortex, by more strongly amplifying weaker compared with stronger sensory-evoked responses (for instance specifically amplifying responses to stimuli not moving along the direction preferred by individual neurons) and by facilitating responses to stimuli entering the receptive field of single neurons. Such enhancement, however, comes at the expense of orientation and direction selectivity, which increased when the selected higher order visual areas were inactivated. Thus, feedback from higher order visual areas selectively amplifies weak sensory-evoked V1 responses, which may enable more robust processing of visual stimuli.