Shift In Tuning Research Articles

Surround suppression and temporal processing of visual signals.

Extraclassical surround suppression strongly modulates responses of neurons in the retina, lateral geniculate nucleus (LGN), and primary visual cortex. Although a great deal is known about the spatial properties of extraclassical suppression and the role it serves in stimulus size tuning, relatively little is known about how extraclassical suppression shapes visual processing in the temporal domain. We recorded the spiking activity of retinal ganglion cells and LGN neurons in the cat to test the hypothesis that extraclassical suppression influences temporal features of visual responses in the early visual system. Our results demonstrate that extraclassical suppression not only shifts the distribution of interspike intervals in a manner that decreases the efficacy of neuronal communication, it also decreases the reliability of neuronal responses to visual stimuli and it decreases the duration of visual responses, an effect that underlies a rightward shift in the temporal frequency tuning of LGN neurons. Taken together, these results reveal a dynamic relationship between extraclassical suppression and the temporal features of neuronal responses.

Plasticity during motherhood: changes in excitatory and inhibitory layer 2/3 neurons in auditory cortex.

Maternal behavior can be triggered by auditory and olfactory cues originating from the newborn. Here we report how the transition to motherhood affects excitatory and inhibitory neurons in layer 2/3 (L2/3) of the mouse primary auditory cortex. We used in vivo two-photon targeted cell-attached recording to compare the response properties of parvalbumin-expressing neurons (PVNs) and pyramidal glutamatergic neurons (PyrNs). The transition to motherhood shifts the average best frequency of PVNs to higher frequency by a full octave, with no significant effect on average best frequency of PyrNs. The presence of pup odors significantly reduced the spontaneous and evoked activity of PVN. This reduction of feedforward inhibition coincides with a complimentary increase in spontaneous and evoked activity of PyrNs. The selective shift of PVN frequency tuning should render pup odor-induced disinhibition more effective for high-frequency stimuli, such as ultrasonic vocalizations. Indeed, pup odors increased neuronal responses of PyrNs to pup ultrasonic vocalizations. We conclude that plasticity in the mothers is mediated, at least in part, via modulation of the feedforward inhibition circuitry in the auditory cortex.

Critical band masking reveals the effects of optical distortions on the channel mediating letter identification.

There is evidence that letter identification is mediated by only a narrow band of spatial frequencies and that the center frequency of the neural channel thought to underlie this selectivity is related to the size of the letters. When letters are spatially filtered (at a fixed size) the channel tuning characteristics change according to the properties of the spatial filter (Majaj et al., 2002). Optical aberrations in the eye act to spatially filter the image formed on the retina—their effect is generally to attenuate high frequencies more than low frequencies but often in a non-monotonic way. We might expect the change in the spatial frequency spectrum caused by the aberration to predict the shift in channel tuning observed for aberrated letters. We show that this is not the case. We used critical-band masking to estimate channel-tuning in the presence of three types of aberration—defocus, coma and secondary astigmatism. We found that the maximum masking was shifted to lower frequencies in the presence of an aberration and that this result was not simply predicted by the spatial-frequency-dependent degradation in image quality, assessed via metrics that have previously been shown to correlate well with performance loss in the presence of an aberration. We show that if image quality effects are taken into account (using visual Strehl metrics), the neural channel required to model the data is shifted to lower frequencies compared to the control (no-aberration) condition. Additionally, we show that when spurious resolution (caused by π phase shifts in the optical transfer function) in the image is masked, the channel tuning properties for aberrated letters are affected, suggesting that there may be interference between visual channels. Even in the presence of simulated aberrations, whose properties change from trial-to-trial, observers exhibit flexibility in selecting the spatial frequencies that support letter identification.

Intention estimation in brain–machine interfaces

Objective. The objective of this work was to quantitatively investigate the mechanisms underlying the performance gains of the recently reported ‘recalibrated feedback intention-trained Kalman Filter’ (ReFIT-KF). Approach. This was accomplished by designing variants of the ReFIT-KF algorithm and evaluating training and online data to understand the neural basis of this improvement. We focused on assessing the contribution of two training set innovations of the ReFIT-KF algorithm: intention estimation and the two-stage training paradigm. Main results. Within the two-stage training paradigm, we found that intention estimation independently increased target acquisition rates by 37% and 59%, respectively, across two monkeys implanted with multiunit intracortical arrays. Intention estimation improved performance by enhancing the tuning properties and the mutual information between the kinematic and neural training data. Furthermore, intention estimation led to fewer shifts in channel tuning between the training set and online control, suggesting that less adaptation was required during online control. Retraining the decoder with online BMI training data also reduced shifts in tuning, suggesting a benefit of training a decoder in the same behavioral context; however, retraining also led to slower online decode velocities. Finally, we demonstrated that one- and two-stage training paradigms performed comparably when intention estimation is applied. Significance. These findings highlight the utility of intention estimation in reducing the need for adaptive strategies and improving the online performance of BMIs, helping to guide future BMI design decisions.

Attention during natural vision warps semantic representation across the human brain

Little is known about how attention changes the cortical representation of sensory information in humans. Based on neurophysiological evidence, we hypothesized that attention causes tuning changes to expand the representation of attended stimuli at the cost of unattended stimuli. To investigate this issue we used functional MRI (fMRI) to measure how semantic representation changes when searching for different object categories in natural movies. We find that many voxels across occipito-temporal and fronto-parietal cortex shift their tuning toward the attended category. These tuning shifts expand the representation of the attended category and of semantically-related but unattended categories, and compress the representation of categories semantically-dissimilar to the target. Attentional warping of semantic representation occurs even when the attended category is not present in the movie, thus the effect is not a target-detection artifact. These results suggest that attention dynamically alters visual representation to optimize processing of behaviorally relevant objects during natural vision.

Adaptation to Speed in Macaque Middle Temporal and Medial Superior Temporal Areas

The response of a sensory neuron to an unchanging stimulus typically adapts, showing decreases in response gain that are accompanied by changes in the shape of tuning curves. It remains unclear whether these changes arise purely due to spike rate adaptation within single neurons or whether they are dependent on network interactions between neurons. Further, it is unclear how the timescales of neural and perceptual adaptation are related. To examine this issue, we compared speed tuning of middle temporal (MT) and medial superior temporal neurons in macaque visual cortex after adaptation to two different reference speeds. For 75% of speed-tuned units, adaptation caused significant changes in tuning that could be explained equally well as lateral shifts, vertical gain changes, or both. These tuning changes occurred rapidly, as both neuronal firing rate and Fano factor showed no evidence of changing beyond the first 500 ms after motion onset, and the magnitude of tuning curve changes showed no difference between trials with adaptation durations shorter or longer than 1 s. Importantly, the magnitude of tuning shifts was correlated with the transient-sustained index, which measures a well characterized form of rapid response adaptation in MT, and is likely associated with changes at the level of neuronal networks. Tuning curves changed in a manner that increased neuronal sensitivity around the adapting speed, consistent with improvements in human and macaque psychophysical performance that we observed over the first several hundred ms of adaptation.

Octopaminergic modulation of neuronal responses to flight-induced naturalistic image sequences

Event Abstract Back to Event Octopaminergic modulation of neuronal responses to flight-induced naturalistic image sequences Diana Rien1, Roland Kern1 and Rafael Kurtz1* 1 Bielefeld University, Neurobiology, Germany Sensory input experienced by animals during behavioral action may differ fundamentally during different states of locomotor activity. Given the limited working range of neurons, adjustments of their sensitivity and stimulus tuning to the current locomotor state might be advantageous. Such state-dependent modulation, which has recently received much attention in various animal models (1), was demonstrated in visual motion-sensitive neurons of different fly species (2-4). The modulation of these neurons is mediated by octopamine, as was recently shown directly by manipulating the activity of a set of octopaminergic neurons (5). Flight or walking activity, as well as application of an octopamine agonist caused enhanced resting activity, shorter latencies in the response to abrupt motion onsets, and larger stimulus-induced responses (2,3,5-10). The strongest response boosts were observed with sustained presentation of high temporal frequencies, thus leading in some studies to a shift in the temporal frequency tuning toward higher values (2,7). However, the functional relevance of these state-dependent changes is unclear, because experimenter-defined stimuli have been used instead of natural optic flow. We assessed the functional significance of octopaminergic neuromodulation for the processing of optic flow as experienced during flight. Naturalistic image sequences were reconstructed based on the head position and gaze orientation during flight (11) and replayed with a panoramic stimulus device (12). We studied how the responses of the V1 neuron in Calliphora vicina to these stimuli are modified by octopaminergic modulation. Spontaneous as well as stimulus- induced spike rates were increased by the octopamine agonist CDM and decreased by an antagonist, epinastine. Moreover, small but consistent shifts in the peak positions of cross- correlograms between the different drug-induced states were present, consistent with an octopamine-induced acceleration of responses to optic flow cues. However, cross-correlations as well as coherence values between the responses recorded in different states of octopaminergic modulation were almost as high as those between traces obtained within one state. Thus, our results suggest that naturalistic optic-flow processing is slightly modified, but not fundamentally altered, by state-dependent neuromodulation. Figure 1

Activation of the basolateral amygdala induces long-term enhancement of specific memory representations in the cerebral cortex

The basolateral amygdala (BLA) modulates memory, particularly for arousing or emotional events, during post-training periods of consolidation. It strengthens memories whose substrates in part or whole are stored remotely, in structures such as the hippocampus, striatum and cerebral cortex. However, the mechanisms by which the BLA influences distant memory traces are unknown, largely because of the need for identifiable target mnemonic representations. Associative tuning plasticity in the primary auditory cortex (A1) constitutes a well-characterized candidate specific memory substrate that is ubiquitous across species, tasks and motivational states. When tone predicts reinforcement, the tuning of cells in A1 shifts toward or to the signal frequency within its tonotopic map, producing an over-representation of behaviorally important sounds. Tuning shifts have the cardinal attributes of forms of memory, including associativity, specificity, rapid induction, consolidation and long-term retention and are therefore likely memory representations. We hypothesized that the BLA strengthens memories by increasing their cortical representations. We recorded multiple unit activity from A1 of rats that received a single discrimination training session in which two tones (2.0s) separated by 1.25 octaves were either paired with brief electrical stimulation (400ms) of the BLA (CS+) or not (CS−). Frequency response areas generated by presenting a matrix of test tones (0.5–53.82kHz, 0–70dB) were obtained before training and daily for 3weeks post-training. Tuning both at threshold and above threshold shifted predominantly toward the CS+ beginning on Day 1. Tuning shifts were maintained for the entire 3weeks. Absolute threshold and bandwidth decreased, producing less enduring increases in sensitivity and selectivity. BLA-induced tuning shifts were associative, highly specific and long-lasting. We propose that the BLA strengthens memory for important experiences by increasing the number of neurons that come to best represent that event. Traumatic, intrusive memories might reflect abnormally extensive representational networks due to hyper-activity of the BLA consequent to the release of excessive amounts of stress hormones.

The Electric Field Tuning Characteristics in the Strain-Mediated Ferrite-Piezoelectric Laminated Magnetoelectric Microwave Devices

This research focused on the numerical simulation of electric field tuning characteristics of the magnetoelectric microwave devices with the core of the laminated ferrite-piezoelectric magnetoelectric materials. Firstly, we proposed an expression for the shift of ferromagnetic resonance (FMR) frequency tuning by external electric field in piezoelectric layer, then substituted the expression of magnetoelectic (ME) constant into this expression, which can convert the electric field tuning on the laminated magnetoelectric materials into the equivalent magnetic field tuning on the ferrite layer. Secondly, we built a numerical analysis model for tunable magnetoelectric microwave device which is formed by the laminated magnetoelectric material and microstrip line, and put the equivalent magnetic field into this numerical model, then the numerical analysis model of novel tunable magentoelectric microwave device is completed. Taking a tunable ME microwave resonator for example, the S parameters calculated by the numerical model in this paper are in good agreement with experimental results both qualitatively and quantitatively. Considering the demand for the designing and applying of the microwave devices, we studied the effect of geometrical dimensions of the ferrite layer (YIG/GGG) on the return loss and the resonance frequency.

Octopaminergic modulation of contrast gain adaptation in fly visual motion‐sensitive neurons

Locomotor activity like walking or flying has recently been shown to alter visual processing in several species. In insects, the neuromodulator octopamine is thought to play an important role in mediating state changes during locomotion of the animal [K.D. Longden & H.G. Krapp (2009) J. Neurophysiol., 102, 3606-3618; (2010) Front. Syst. Neurosci., 4, 153; S.N. Jung et al. (2011)J. Neurosci., 31, 9231-9237]. Here, we used the octopamine agonist chlordimeform (CDM) to mimic effects of behavioural state changes on visual motion processing. We recorded from identified motion-sensitive visual interneurons in the lobula plate of the blowfly Calliphora vicina. In these neurons, which are thought to be involved in visual guidance of locomotion, motion adaptation leads to a prominent attenuation of contrast sensitivity. Following CDM application, the neurons maintained high contrast sensitivity in the adapted state. This modulation of contrast gain adaptation was independent of the activity of the recorded neurons, because it was also present after stimulation with visual motion that did not result in deviations from the neurons' resting activity. We conclude that CDM affects presynaptic inputs of the recorded neurons. Accordingly, the effect of CDM was weak when adapting and test stimuli were presented in different parts of the receptive field, stimulating separate populations of local presynaptic neurons. In the peripheral visual system adaptation depends on the temporal frequency of the stimulus pattern and is therefore related to pattern velocity. Contrast gain adaptation could therefore be the basis for a shift in the velocity tuning that was previously suggested to contribute to state-dependent processing of visual motion information in the lobula plate interneurons.

Activity of the same motor cortex neurons during repeated experience with perturbed movement dynamics

Neurons in the primary motor cortex (M1) have been shown to have persistent, memory-like activity following adaptation to altered movement dynamics. However, the techniques used to study these memory traces limited recordings to only single sessions lasting no more than a few hours. Here, chronically implanted microelectrode arrays were used to study the long-term neuronal responses to repeated experience with perturbing, velocity-dependent force fields. Force-field-related neuronal activity within each session was similar to that found previously. That is, the directional tuning curves of the M1 neurons shifted in a manner appropriate to compensate for the forces. Next, the across-session behavior was examined. Long-term learning was evident in the performance improvements across multiple force-field sessions. Correlated with this change, the neuronal population had smaller within-session spike rate changes as experience with the force field increased. The smaller within-session changes were a result of persistent across-session shifts in directional tuning. The results extend the observation of memory traces of newly learned dynamics and provide further evidence for the role of M1 in early motor memory formation.

The influence of surround suppression on adaptation effects in primary visual cortex

Adaptation, the prolonged presentation of stimuli, has been used to probe mechanisms of visual processing in physiological, imaging, and perceptual studies. Previous neurophysiological studies have measured adaptation effects by using stimuli tailored to evoke robust responses in individual neurons. This approach provides an incomplete view of how an adapter alters the representation of sensory stimuli by a population of neurons with diverse functional properties. We implanted microelectrode arrays in primary visual cortex (V1) of macaque monkeys and measured orientation tuning and contrast sensitivity in populations of neurons before and after prolonged adaptation. Whereas previous studies in V1 have reported that adaptation causes stimulus-specific suppression of responsivity and repulsive shifts in tuning preference, we have found that adaptation can also lead to response facilitation and shifts in tuning toward the adapter. To explain this range of effects, we have proposed and tested a simple model that employs stimulus-specific suppression in both the receptive field and the spatial surround. The predicted effects on tuning depend on the relative drive provided by the adapter to these two receptive field components. Our data reveal that adaptation can have a much richer repertoire of effects on neuronal responsivity and tuning than previously considered and suggest an intimate mechanistic relationship between spatial and temporal contextual effects.

Dynamics of spatial frequency tuning in mouse visual cortex

Neuronal spatial frequency tuning in primary visual cortex (V1) substantially changes over time. In both primates and cats, a shift of the neuron's preferred spatial frequency has been observed from low frequencies early in the response to higher frequencies later in the response. In most cases, this shift is accompanied by a decreased tuning bandwidth. Recently, the mouse has gained attention as a suitable animal model to study the basic mechanisms of visual information processing, demonstrating similarities in basic neuronal response properties between rodents and highly visual mammals. Here we report the results of extracellular single-unit recordings in the anesthetized mouse where we analyzed the dynamics of spatial frequency tuning in V1 and the lateromedial area LM within the lateral extrastriate area V2L. We used a reverse-correlation technique to demonstrate that, as in monkeys and cats, the preferred spatial frequency of mouse V1 neurons shifted from low to higher frequencies later in the response. However, this was not correlated with a clear selectivity increase or enhanced suppression of responses to low spatial frequencies. These results suggest that the neuronal connections responsible for the temporal shift in spatial frequency tuning may considerably differ between mice and monkeys.

Modeling short-term adaptation processes of visual motion detectors

In the brain, both neural processing dynamics as well as the perceptual interpretation of a stimulus can depend on sensory history. The underlying principle is a sensory adaptation to the statistics of the input collected over some timespan, allowing the system to tune its detectors, e.g. by better sampling the input space and adjusting the response. Here, we show how a model for adaptation in visual motion processing can be set up from first principles using a generative formulation and casting the problem of adaptation in terms of optimal estimation over time. The model leads to an online adaptation of velocity tuning curves, inducing shifts in the velocity tuning and changes in the tuning curve widths that are compatible with observations from physiological experiments on macaque MT neurons. We also show how such an adaptation leads to a greater computational efficiency by a better sampling of the velocity space, requiring less motion detectors to achieve a desired level of estimation accuracy.

Spatial Shifts of Audio-Visual Interactions by Perceptual Learning are Specific to the Trained Orientation and Eye

A large proportion of the human cortex is devoted to visual processing. Contrary to the traditional belief that multimodal integration takes place in multimodal processing areas separate from visual cortex, several studies have found that sounds may directly alter processing in visual brain areas. Furthermore, recent findings show that perceptual learning can change the perceptual mechanisms that relate auditory and visual senses. However, there is still a debate about the systems involved in cross-modal learning. Here, we investigated the specificity of audio-visual perceptual learning. Audio-visual cuing effects were tested on a Gabor orientation task and an object discrimination task in the presence of lateralised sound cues before and after eight-days of cross-modal task-irrelevant perceptual learning. During training, the sound cues were paired with visual stimuli that were misaligned at a proximal (trained) visual field location relative to the sound. Training was performed with one eye patched and with only one Gabor orientation. Consistent with previous findings we found that cross-modal perceptual training shifted the audio-visual cueing effect towards the trained retinotopic location. However, this shift in audio-visual tuning was only observed for the trained stimulus (Gabors), at the trained orientation, and in the trained eye. This specificity suggests that multimodal interactions resulting from cross-modal (audio-visual) task-irrelevant perceptual learning involves so-called unisensory visual processing areas in humans. Our findings provide further support for recent anatomical and physiological findings that suggest relatively early interactions in cross-modal processing.

The opsin shift and mechanism of spectral tuning in rhodopsin.

Molecular dynamics simulations and combined quantum mechanical and molecular mechanical calculations have been performed to investigate the mechanism of the opsin shift and spectral tuning in rhodopsin. A red shift of -980 cm(-1) was estimated in the transfer of the chromophore from methanol solution environment to the protonated Schiff base (PSB)-binding site of the opsin. The conformational change from a 6-s-cis-all-trans configuration in solution to the 6-s-cis-11-cis conformer contributes additional -200 cm(-1), and the remaining effects were attributed to dispersion interactions with the aromatic residues in the binding site. An opsin shift of 2100 cm(-1) was obtained, in reasonable accord with experiment (2730 cm(-1)). Dynamics simulations revealed that the 6-s-cis bond can occupy two main conformations for the β-ionone ring, resulting in a weighted average dihedral angle of about -50°, which may be compared with the experimental estimate of -28° from solid-state NMR and Raman data. We investigated a series of four single mutations, including E113D, A292S, T118A, and A269T, which are located near the PSB, along the polyene chain of retinal and close to the ionone ring. The computational results on absorption energy shift provided insights into the mechanism of spectral tuning, which involves all means of electronic structural effects, including the stabilization or destabilization of either the ground or the electronically excited state of the retinal PSB.

Neocortical Interneurons: From Diversity, Strength

Interneurons in the neocortex of the brain are small, locally projecting inhibitory GABAergic cells with a broad array of anatomical and physiological properties. The diversity of interneurons is believed to be crucial for regulating myriad operations in the neocortex. Here, we describe current theories about how interneuron diversity may support distinct neocortical processes that underlie perception.

What and where: A Bayesian inference theory of attention

In the theoretical framework of this paper, attention is part of the inference process that solves the visual recognition problem of what is where. The theory proposes a computational role for attention and leads to a model that predicts some of its main properties at the level of psychophysics and physiology. In our approach, the main goal of the visual system is to infer the identity and the position of objects in visual scenes: spatial attention emerges as a strategy to reduce the uncertainty in shape information while feature-based attention reduces the uncertainty in spatial information. Featural and spatial attention represent two distinct modes of a computational process solving the problem of recognizing and localizing objects, especially in difficult recognition tasks such as in cluttered natural scenes.We describe a specific computational model and relate it to the known functional anatomy of attention. We show that several well-known attentional phenomena – including bottom-up pop-out effects, multiplicative modulation of neuronal tuning curves and shift in contrast responses – all emerge naturally as predictions of the model. We also show that the Bayesian model predicts well human eye fixations (considered as a proxy for shifts of attention) in natural scenes.

Large dielectric tuning and microwave phase shift at low electric field in epitaxial Ba0.5Sr0.5TiO3 on SrTiO3

Dielectric properties of annealed and as-grown ferroelectric Ba0.5Sr0.5TiO3 (BST) grown by pulsed laser deposition on sputtered BST seed layers on strontium titanate (STO) substrates were investigated at microwave frequencies in the realm of tunability of its dielectric constant as well as phase shifters based on this material. The as-grown layers were nearly fully relaxed with measured lattice parameters nearly identical to those of bulk BST. The tuning of the relative dielectric constant (∼1750 at zero bias at 10 GHz) of the annealed BST was found to be as high as 59% and 56% at 10 and 19 GHz, respectively. The analysis of the loss in the BST results in a measured tan δ of 0.02 for the annealed as well as the unannealed films at a frequency of 18 GHz. Phase shifters also exhibited high tuning with differential phase shift figures of merit of 35 and 55°/dB at a field of 60 kV/cm at 10 and 19 GHz, respectively. Serendipitously, most of the tuning occurs at low fields, and thus we propose a new figure of merit, taking into account the amplitude of applied electric field in order to achieve the phase shift. In this new realm we achieved the values of this overall figure of merit of 1.2 and 1.8° cm/dB kV at 10 and 19 GHz, respectively, using an applied electric field of only 10 kV/cm.

Representational gain in cortical area underlies increase of memory strength

Neuronal plasticity that develops in the cortex during learning is assumed to represent memory content, but the functions of such plasticity are actually unknown. The shift in spectral tuning in primary auditory cortex (A1) to the frequency of a tone signal is a compelling candidate for a substrate of memory because it has all of the cardinal attributes of associative memory: associativity, specificity, rapid induction, consolidation, and long-term retention. Tuning shifts increase the representational area of the signal in A1, as an increasing function of performance level, suggesting that area encodes the magnitude of acquired stimulus significance. The present study addresses the question of the specific function of learning-induced associative representational plasticity. We tested the hypothesis that specific increases in A1 representational area for an auditory signal serve the mnemonic function of enhancing memory strength for that signal. Rats were trained to bar-press for reward contingent on the presence of a signal tone (5.0 kHz), and assessed for memory strength during extinction. The amount of representational area gain for the signal frequency band was significantly positively correlated with resistance to extinction to the signal frequency in two studies that spanned the range of task difficulty. These findings indicate that specific gain in cortical representational area underlies the strength of the behaviorally-relevant contents of memory. Thus, mnemonic functions of cortical plasticity are determinable.