Biology subjects

Knoblauch, K.

Publications and source records attributed to Knoblauch, K..

16 recordsLinked to original sources

bioRxiv2025

Effects of color-enhancing filters on color salience in normal trichromats

Notch filters can alter color contrasts by selectively filtering different spectral bands of the stimulus and have been developed to enhance reddish-greenish contrasts for color-deficient observers with anomalous trichromacy. We examined the effects of such filters on color salience for normal trichromats, using a visual search paradigm where the task was to locate a color target superimposed on a variegated chromatic background, similar to foraging for fruits among foliage. Background colors varied along a bluish-yellowish or purpliish to yellow-green (short-wave cone isolating) axis, roughly spanning the range of dominant color variations in arid or lush environments. Target colors sampled a wide range of hue angles and contrasts. Testing was conducted on a computer monitor, with the filter effects simulated by calculating corresponding chromaticities with or without the filter for naturalistic (Munsell) reflectances. The filter evaluated (Enchroma SuperX(R) glasses) was designed to increase color contrast along a magenta-green axis. Consistent with this, search times for targets on the blue-yellow background were significantly faster for the filter condition, because the filter increased the target-background color difference. Alternatively, overall differences in search times were not observed for the S-cone background. The differences on the two backgrounds could be qualitatively accounted for by the relative salience of the stimuli predicted by a perceptual color space (CIELAB). Our results demonstrate the efficacy of the filters for enhancing visual performance for normal trichromats and naturalistic tasks, and illustrate how these effects depend on the potential color characteristics of the environment.

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bioRxiv2024

A principled approach to community detection in interareal cortical networks

Interareal structural connectivity explored by tract-tracing reveals super-dense, weighted, directed, and spatially embedded complex networks. These properties make the extraction and alignment of the community structure of these networks with brain function challenging. Here, we overcome these difficulties using a principled, information theoretic approach that distinguishes connectivity profiles using the Hellinger distance. Applying it to retrograde tract-tracing data in the macaque, we show that the cortex at the interareal level is organized into a nested hierarchy of link-communities alongside a node-community hierarchy. We find that the [1/2]-Renyi divergence of connection profiles, a non-linear transform of the Hellinger metric, follows a Weibull-like distribution and scales linearly with interareal distances, establishing a quantitative expression between functional organization and cortical geometry. Finally, we show that the loop entropy along the hierarchy is maximized at a community description level that is neither excessively detailed, nor non-specific, defining a "Goldilocks" level that we hypothesize, is optimal for system-wide efficient information processing. Author SummaryUnderstanding how functional organization emerges from connectivity alone is a central challenge in neuroscience. The cortex forms a dense network, making it difficult to identify meaningful groups of areas using standard methods. Here, we introduce a principled information-theoretical approach that compares the connectivity patterns of brain areas to uncover their community structure and organization. By first detecting the link communities and then deriving the grouping of areas from it, our method reveals a hierarchical structure based on patterns of reciprocal communication. Applied to the retrograde tract-tracing database of the macaque cortical network, this approach identifies communities that share similar interaction patterns, offering new insights into large-scale brain organization. More broadly, it provides a general framework for analyzing complex networks with dense, directed, and weighted connections.

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bioRxiv2023

Modulation of motion signals across the visual cortical hierarchy during bistable perception

Top-down influences play a central role in perception. In multi-stable sensory phenomena subjective interpretation fluctuates spontaneously over time and the perceptual switches reflect internally driven representations engaging perceptual decision-making. Using functional magnetic resonance imaging (fMRI), we investigated how perceptual states are represented across early visual areas during perception of a bistable moving plaid. While subjects maintained central fixation, perception of the stimulus--two superimposed, obliquely oriented moving gratings--spontaneously alternated between component motion (2 gratings sliding obliquely over each other) and pattern motion (a single coherent pattern moving laterally). Direction-selective clusters were reliably modulated by perceptual state across both early (V1) and higher-order (V2 and the human medial temporal complex (hMT+)) visual regions. Pattern modulation was significantly stronger in primary visual cortex (V1) than in higher-order areas, suggesting that coherent percept formation engages mechanisms beyond simple bottom-up feature integration. Enhanced V1 modulation points to a prominent role for top-down influences from higher-order regions, consistent with predictive coding frameworks in which higher-level areas convey predictions to lower levels to disambiguate sensory input and drive perceptual inference under ambiguous conditions. Eye tracking confirmed that reflexive eye movement direction did not predict perceptual state, excluding an oculomotor confound. Interpretation of the results in light of current knowledge of the physiology of motion integration in early visual areas suggests circuits at which feedback would be predicted to act.

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bioRxiv2022

Retinotopic organization of feedback projections in primate early visual cortex: implications for active vision

Feedback connections play a major role in many theories of brain function. Previous studies of feedback connections to early visual areas have mainly concentrated on the representation of central visual fields. Here, injections of tracers at different eccentricities in areas V1 and V2 revealed retinotopically organized feedback. Peripheral injections revealed projections from 15 areas that are not labeled by central injections. Connection strengths for the majority of projections vary significantly with eccentricity in a systematic fashion with respect to distance and origin; whereas projections to central and upper visual fields are significantly stronger from ventral stream areas, peripheral and lower field projections are stronger from the dorsal stream. Non-invasive functional connectivity suggests a similar anatomical organization in humans. These features are discussed with respect to the cognitive and perceptual roles of these feedback pathways.

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bioRxiv2020

Predictability of cortical-cortical connections in the mammalian brain

Despite a five-order magnitude range in size, the mammalian brain exhibits many shared anatomical and functional characteristics that should translate into cortical network commonalities. Here we develop a framework employing machine learning to quantify the degree of predictability of the weighted interareal cortical matrix. Data were obtained with retrograde tract-tracing experiments supplemented by projection length measurements. Using this framework with consistent and edge-complete empirical datasets in the macaque and mouse cortex, we show that there is significant amount of predictability embedded in the interareal cortical networks of both species. At the binary level, links are predictable with an Area Under the ROC curve of at least 0.8 for the macaque. At the weighted level, strengths of the medium and strong links are predictable with at least 85-90% accuracy in mouse and 70-80% in macaque, whereas weak links are not predictable in either species. These observations suggest that the formation and evolution of the cortical network at the mesoscale is to a large extent, rule-based, motivating further research on the architectural invariants of the cortical connectome.

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bioRxiv2020

A dopamine gradient controls access to distributed working memory in monkey cortex

Dopamine is critical for working memory. However, its effects throughout the large-scale primate cortex are poorly understood. Here we report that dopamine receptor density per neuron, measured by receptor autoradiography in the macaque monkey cortex, displays a macroscopic gradient along the cortical hierarchy. We developed a connectome- and biophysically-based model for distributed working memory that incorporates multiple neuron types and a dopamine gradient. The model captures an inverted U-shaped dependence of working memory on dopamine. The spatial distribution of mnemonic persistent activity matches that observed in over 90 experimental studies. We show that dopamine filters out irrelevant stimuli by enhancing inhibition of pyramidal cell dendrites. The level of cortical dopamine can also determine whether memory encoding is through persistent activity or an internal synaptic state. Taken together, our work represents a cross-level understanding that links molecules, cell types, recurrent circuit dynamics and a core cognitive function distributed across the cortex.

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bioRxiv2020

Cortical Hierarchy and The Dual Counterstream Architecture.

Hierarchy is a major organizational principle of the cortex and underscores modern computational theories of cortical function. The local microcircuit amplifies long-distance inter-areal input, which show distance-dependent changes in their laminar profiles. Statistical modeling of these changes in laminar profiles demonstrates that inputs from multiple hierarchical levels to their target areas show remarkable consistency, allowing the construction of a cortical hierarchy based on a principle of hierarchical distance. The statistical modeling that is applied to structure can also be applied to laminar differences in the oscillatory coherence between areas thereby determining a functional hierarchy of the cortex. Close examination of the anatomy of inter-areal connectivity reveals a dual counterstream architecture with well-defined distance-dependent feedback and feedforward pathways in both the supra- and infragranular layers, suggesting a multiplicity of feedback pathways with well-defined functional properties. These findings are consistent with feedback connections providing a generative network involved in a wide range of cognitive functions. A dynamical model constrained by connectivity data shed insights into the experimentally observed signatures of frequency-dependent Granger causality for feedforward versus feedback signaling. Concerted experiments capitalizing on recent technical advances and combining tract-tracing, high-resolution fMRI, optogenetics and mathematical modeling hold the promise of a much improved understanding of lamina-constrained mechanisms of neural computation and cognition. However, because inter-areal interactions involve cortical layers that have been the target of important evolutionary changes in the primate lineage, these investigations will need to include human and non-human primates comparisons. PlanO_LIIntroduction C_LIO_LIHierarchy - signatures of inputs to the local circuits. C_LIO_LIModels of hierarchy C_LIO_LIHierarchy - input consistency C_LIO_LIDual stream architecture. C_LIO_LIFunctional characteristics of FF and FB pathways. C_LIO_LIThe predictive brain and the importance of top-down generative networks. C_LIO_LIConclusion. C_LI

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bioRxiv2020

Canonical and noncanonical features of the mouse visual cortical hierarchy

Neocortical circuit computations underlying active vision are performed by a distributed network of reciprocally connected, functionally specialized areas. Mouse visual cortex is a dense, hierarchically organized network, comprising subnetworks that form preferentially interconnected processing streams. To determine the detailed layout of the mouse visual hierarchy, laminar patterns formed by interareal axonal projections, originating in each of ten visual areas were analyzed. Reciprocally connected pairs of areas, and shared targets of pairs of source areas, exhibited structural features consistent with a hierarchical organization. Beta regression analyses, which estimated a continuous measure of hierarchical distance, indicated that the network comprises multiple hierarchies embedded within overlapping processing levels. Single unit recordings showed that within each processing stream, receptive field sizes typically increased with increasing hierarchical level; however, ventral stream areas showed overall larger receptive field diameters. Together, the results reveal canonical and noncanonical hierarchical network motifs in mouse visual cortex.

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bioRxiv2020

Voice and Face Gender Perception engages multimodal integration via multiple feedback pathways.

Multimodal integration provides an ideal framework for investigating top-down influences in perceptual integration. Here, we investigate mechanisms and functional networks participating in face-voice multimodal integration during gender perception by using complementary behavioral (Maximum Likelihood Conjoint Measurement) and brain imaging (Dynamic Causal Modeling of fMRI data) techniques. Thirty-six subjects were instructed to judge pairs of face-voice stimuli either according to the gender of the face (face task), the voice (voice task) or the stimulus (stimulus task; no specific modality instruction given). Face and voice contributions to the tasks were not independent, as both modalities significantly contributed to all tasks. The top-down influences in each task could be modeled as a differential weighting of the contributions of each modality with an asymmetry in favor of the auditory modality in terms of magnitude of the effect. Additionally, we observed two independent interaction effects in the decision process that reflect both the coherence of the gender information across modalities and the magnitude of the gender difference from neutral. In a second experiment we investigated with functional MRI the modulation of effective connectivity between the Fusiform Face Area (FFA) and the Temporal Voice Area (TVA), two cortical areas implicated in face and voice processing. Twelve participants were presented with multimodal face-voice stimuli and instructed to attend either to face, voice or any gender information. We found specific changes in effective connectivity between these areas in the same conditions that generated behavioral interactions. Taken together, we interpret these results as converging evidence supporting the existence of multiple parallel hierarchical systems in multi-modal integration.

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bioRxiv2019

Suprathreshold contrast response in normal and anomalous trichromats

Maximum Likelihood Difference Scaling was used to measure suprathreshold contrast response difference scales for low-frequency Gabor patterns modulated along luminance and L-M color directions in normal, protanomalous, and deuteranomalous observers. Based on a signal-detection model, perceptual scale values, parameterized as d', were estimated by maximum likelihood. The difference scales were well fit by a Michaelis-Menten model, permitting estimates of response and contrast gain parameters for each subject. Anomalous observers showed no significant differences in response or contrast gain from normal observers for luminance contrast. For chromatic modulations, however, anomalous observers displayed higher contrast and lower response gain compared to normal observers. These effects cannot be explained by simple pigment shift models and support a compensation mechanism to optimize the mapping of the input contrast range to the neural response range. A linear relation between response and contrast gain suggests a neural trade-off between them.

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bioRxiv2019

Unique Features of Sub-Cortical Circuits in A Macaque Model of Congenital Blindness

There is extensive modification of the functional organization of the brain in the congenital blind human, although there is little understanding of the structural underpinnings of these changes. The visual system of macaque has been extensively characterized both anatomically and functionally. We have taken advantage of this to examine the influence of the congenital blindness in macaque resulting from the removal of the retina during in utero development. Developmental anophthalmia in macaque effectively removes the normal influence of the thalamus on cortical development leading to an induced hybrid cortex (HC) combining features of primary visual and extrastriate cortex. Here we show that retrograde tracers injected in early visual areas including hybrid cortex reveals a drastic reduction of cortical projections of the reduced lateral geniculate nucleus. In addition, there is an important expansion of projections from the pulvinar complex to the hybrid cortex, compared to the controls. These findings show that the functional consequences of congenital blindness need to be considered in terms of both modifications of the inter-areal cortical network and the ascending visual pathways.

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bioRxiv2019

Refinement of the primate corticospinal pathway during prenatal development

Perturbation of the developmental refinement of the corticospinal pathway leads to motor disorders. In non-primates developmental refinement is well documented, however in primates invasive investigations of the developing corticospinal pathway have been confined to neonatal and postnatal stages when refinement is relatively modest.\n\nHere, we investigated the developmental changes in the distribution of corticospinal projection neurons in cynomolgus monkey. Injections of retrograde tracer at the cervical levels of the spinal cord at embryonic day (E) 95 and E105 show that (i) areal distribution of back-labeled neurons is more extensive than in the neonate and dense labeling is found in prefrontal, limbic, temporal and occipital cortex; (ii) distributions of contra- and ipsilateral projecting corticospinal neurons are comparable in terms of location and numbers of labeled neurons, in contrast to the adult where the contralateral projection is an order of magnitude higher than the ipsilateral projection. Findings from one largely restricted injection suggest a hitherto unsuspected early innervation of the gray matter.\n\nIn the fetus there was in addition dense labeling in the central nucleus of the amygdala, the hypothalamus, the subthalamic nucleus and the adjacent region of the zona incerta, subcortical structures with only minor projections in the adult control.

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bioRxiv2017

Effects of background and contour luminance on the hue and brightness of the Watercolor effect

Conjoint measurement was used to investigate the joint influences of the luminance of the background and the inner contour on hue- and brightness filling-in for a stimulus configuration generating a water-color effect (WCE), i.e., a wiggly bi-chromatic contour enclosing a region with the lower luminance component on the exterior. Two stimuli with the background and inner contour luminances covarying independently were successively presented, and in separate experiments, the observer judged which member of the pairs interior regions contained a stronger hue or was brighter. Braided-contour control stimuli that generated little or no perceptual filling-in were also used to assess whether observers were judging the interior regions and not the contours themselves. Three nested models of the contributions of the background and inner contour to the judgments were fit to the data by maximum likelihood and evaluated by likelihood ratio tests. Both stimulus components contributed to both the hue and brightness of the interior region with increasing luminance of the inner contour generating an assimilative filling-in for the hue judgments but a contrast effect for the brightness judgments. Control analyses showed negligible effects for the order of the luminance of the background or inner contour on the judgments. An additive contribution of both components was rejected in favor of a saturated model in which the responses depended on the levels of both stimulus components. For the hue judgments, increased background luminance led to greater hue filling-in at higher luminances of the interior contour. For the brightness judgments, the higher background luminance generated less brightness filling-in at higher luminances of the interior contour. The results indicate different effects of the inner contour and background on the induction of the brightness and coloration percepts of the WCE, suggesting that they are mediated by different mechanisms.

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bioRxiv2017

Neural circuits for long-range color filling-in

Surface color appearance depends on both local surface chromaticity and global context. How are these inter-dependencies supported by cortical networks? Combining functional imaging and psychophysics, we examined if color from long-range filling-in engages distinct pathways from responses caused by a field of uniform chromaticity. We find that color from filling-in is best classified and best correlated with appearance by two dorsal areas, V3A and V3B/KO. In contrast, a field of uniform chromaticity is best classified by ventral areas hV4 and LO. Dynamic causal modeling revealed feedback modulation from area V3A to areas V1 and LO for filling-in, contrasting with feedback from LO modulating areas V1 and V3A for a matched uniform chromaticity. These results indicate a dorsal stream role in color filling-in via feedback modulation of area V1 coupled with a cross-stream modulation of ventral areas suggesting that local and contextual influences on color appearance engage distinct neural networks.

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bioRxiv2017

Cortical Connectivity In A Macaque Model Of Congenital Blindness

AbstractBrain-mapping of the congenitally blind human reveals extensive plasticity(1). The visual cortex of the blind has been observed to support higher cognitive functions including language and numerical processing(2, 3). This functional shift is hypothesized to reflect a metamodal cortical function, where computations are defined by the local network. In the case of developmental deafferentation, local circuits are considered to implement higher cognitive functions by accommodating diverse long-distance inputs(4-7). However, the extent to which visual deprivation triggers a reorganization of the large-scale network in the cortex is still controversial(8). Here we show that early prenatal ablation of the retina, an experimental model of anophthalmia in macaque, leads to a major reduction of area V1 and the creation of a default extrastriate cortex (DEC)(9, 10). Anophthalmic and normal macaques received retrograde tracer injections in DEC, as well as areas V2 and V4 post-natally. This revealed a six-fold expansion of the spatial extent of local connectivity in the DEC and a surprisingly high location of the DEC derived from a computational model of the cortical hierarchy(11). In the anophthalmic the set of areas projecting to the DEC, area V2 and V4 does not differ from that of normal adult controls, but there is a highly significant increase in the relative cumulative weight of the ventral stream areas input to the early visual areas. These findings show that although occupying the territory that would have become primary visual cortex the DEC exhibits features of a higher order area, thus reflecting a combination of intrinsic and extrinsic factors on cortical specification. Understanding the interaction of these contributing factors will shed light on cortical plasticity during primate development and the neurobiology of blindness.

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bioRxiv2017

The Mouse Cortical Interareal Network Reveals Well Defined Connectivity Profiles and an Ultra Dense Cortical Graph

The inter-areal wiring pattern of mouse cerebral cortex was analyzed in relation to an accurate parcellation of cortical areas. Twenty-seven retrograde tracer injections were made in 19 areas of a 41 area (plus 7 sub-area) parcellation of the mouse neo-, parahippocampal and perirhinal cortex. Flat mounts of the cortex and multiple histological markers enabled detailed counts of labeled neurons in individual areas. A weight index was determined for each area-to-area pathway based on the Fraction of Extrinsically Labeled Neurons (FLNe). Data analysis allowed cross species comparison with the macaque. Estimation of FLNe statistical variability based on repeat injections revealed high consistency across individuals and justifies using a single injection per area to characterize connectivity. The observed lognormal distribution of connections to each cortical area spanned 5 orders of magnitude and revealed a distinct connectivity profile for each area, analogous to that observed in macaque. The resulting graph has a density of 97% (i.e. 97% of connections that can exist do exist), considerably higher than the 66% density reported for the macaque. Our results provide more sharply defined connectivity profiles and a markedly higher graph density than shown in a recent probabilistic mouse connectome.

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