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The availability of effective vaccines and a high vaccination rate allowed the recent mitigation, or even withdrawal, of many protective measures for containing the SARS CoV-2 pandemic. At the same time, new and highly mutated variants of the virus are found to have significantly higher transmissibility and reduced vaccine efficacy, thus causing high infection rates during the third year of the pandemic. The combination of reduced measures and increased infectivity poses a particular risk for unvaccinated individuals, including animals susceptible to the virus. Among the latter, non-human primates (NHPs) are particularly vulnerable. They serve as important models in various fields of biomedical research and because of their cognitive capabilities, they receive particular attention in animal welfare regulations around the world. Yet, although they played an extraordinarily important role for developing and testing vaccines against SARS-CoV-2, the protection of captive rhesus monkeys against Covid-19 has rarely been discussed. We here report upon twofold mRNA vaccination of a cohort of 19 elderly rhesus monkeys (Macaca mulatta) against infection by SARS-CoV-2. All animals were closely monitored on possible side effects of vaccination, and were tested for neutralising antibodies against the virus. The data show that vaccination of rhesus monkeys is a safe and reliable measure to protect these animals against SARS-CoV-2.

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The nonhuman primate (NHP) constitutes an extraordinarily important model in neuroscience research for understanding the neuronal underpinnings of perceptual, motor, cognitive, and executive functions of the primate brain, and to study the physiological causes, effects, and potential treatments of brain disorders. Because of their cognitive capabilities, NHPs receive special attention in animal welfare regulations around the world, and their well-being is a benchmark for the evaluation, monitoring, and refinement of experimental procedures. As a consequence, many typical neuroscientific procedures are considered only mildly severe by animal welfare boards. There is, however, an ongoing debate about possible long-term and cumulative effects. Because of a lack of longitudinal data, it is unclear whether mildly severe procedures may cause more significant harm on the long-term, and to what extent they may impact animal well-being and healthiness over time. We here make use of a database of blood samples drawn over a period of 15 years from 39 rhesus monkeys (Macaca mulatta) to address the issue of long-term, cumulative effects of neuroscientific procedures. A careful analysis of indicative primate blood markers for chronic inflammation, hydration status, and stress levels, their comparison to baseline values from both the same animals and the literature, and evaluation of additional hematologic, physiological, and behavioral parameters did not provide support for the notion of long-term, cumulative effects on the monkeys' healthiness and well-being. The results may serve the community as a reference for the severity assessment of neuroscientific experiments involving NHPs.

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Selective visual attention allows the brain to focus on behaviorally relevant information while ignoring irrelevant signals. As a possible mechanism, routing-by-synchronization was proposed: neural populations receiving attended signals align their gamma-rhythmic activity to that of the sending populations, such that incoming spikes arrive at excitability peaks of receiving populations, enhancing signal transfer. Conversely, non-attended signals arrive unaligned to the receiver's oscillation, reducing signal transfer. Therefore, visual signals should be transferred through gamma-rhythmic bursts of information, resulting in a modulation of the stimulus content within the receiving population's activity by its gamma phase and amplitude. To test this prediction, we quantified gamma-phase-dependent stimulus content within neural activity from area V4 of two male macaques performing a visual attention task. For the attended stimulus, we find highest stimulus information content near excitability peaks, an effect that increases with oscillation amplitude, establishing a functional link between selective processing and gamma-activity.SIGNIFICANCE STATEMENT The ability to focus on the behaviorally relevant signals is essential for the brain to cope with the continuous, high-dimensional stream of sensory information it receives. What are the neural mechanisms which allow such selective processing in the visual system? We analyzed data from area V4 and found that the amount of visual signal information content is tightly linked to the phase of local gamma-rhythmic activity, with maximal signal content occurring near peaks of neural excitability. Our investigations provide direct evidence that selective attention relies on rhythmic temporal coordination between visual areas, and establish novel methods for pinpointing pulsed transmission schemes in neural data.

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BACKGROUND: Electrical stimulation is a widely used method in the neurosciences with a variety of application fields. However, stimulation frequently induces large and long-lasting artifacts, which superimpose on the actual neuronal signal. Existing methods were developed for analyzing fast events such as spikes, but are not well suited for the restoration of LFP signals. NEW METHOD: We developed a method that extracts artifact components while also leaving the LFP components of the neuronal signal intact. We based it on an exponential fit of the average artifact shape, which is subsequently adapted to the individual artifacts amplitude and then subtracted. Importantly, we used for fitting of the individual artifact only a short initial time window, in which the artifact is dominating the superimposition with the neuronal signal. Using this short period ensures that LFP components are not part of the fit, which leaves them unaffected by the subsequent artifact removal. RESULTS: By using the method presented here, we could diminish the substantial distortions of neuronal signals caused by electrical stimulation to levels that were statistically indistinguishable from the original data. Furthermore, the effect of stimulation on the phases of γ- and ß- oscillations was reduced by 85 and 75 %, respectively. COMPARISON WITH EXISTING METHODS: This approach avoids signal loss as caused by methods cutting out artifacts and minimizes the distortion of the signal's temporal structure as compared to other approaches. CONCLUSION: The method presented here allows for a successful reconstruction of broad-band signals.

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Recordings of epidural field potentials (EFPs) allow neuronal activity to be acquired over a large region of cortical tissue with minimal invasiveness. Because electrodes are placed on top of the dura and do not enter the neuronal tissue, EFPs offer intriguing options for both clinical and basic science research. On the other hand, EFPs represent the integrated activity of larger neuronal populations and possess a higher trial-by-trial variability and a reduced signal-to-noise ratio due the additional barrier of the dura. It is thus unclear whether and to what extent EFPs have sufficient spatial selectivity to allow for conclusions about the underlying functional cortical architecture, and whether single EFP trials provide enough information on the short timescales relevant for many clinical and basic neuroscience purposes. We used the high spatial resolution of primary visual cortex to address these issues and investigated the extent to which very short EFP traces allow reliable decoding of spatial information. We briefly presented different visual objects at one of nine closely adjacent locations and recorded neuronal activity with a high-density epidural multielectrode array in three macaque monkeys. With the use of receiver operating characteristics (ROC) to identify the most informative data, machine-learning algorithms provided close-to-perfect classification rates for all 27 stimulus conditions. A binary classifier applying a simple max function on ROC-selected data further showed that single trials might be classified with 100% performance even without advanced offline classifiers. Thus, although highly variable, EFPs constitute an extremely valuable source of information and offer new perspectives for minimally invasive recording of large-scale networks.NEW & NOTEWORTHY Epidural field potential (EFP) recordings provide a minimally invasive approach to investigate large-scale neural networks, but little is known about whether they possess the required specificity for basic and clinical neuroscience. By making use of the spatial selectivity of primary visual cortex, we show that single-trial information can be decoded with close-to-perfect performance, even without using advanced classifiers and based on very few data. This labels EFPs as a highly attractive and widely usable signal.

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Neurophysiological data acquisition using multi-electrode arrays and/or (semi-) chronic recordings frequently has to deal with low signal-to-noise ratio (SNR) of neuronal responses and potential failure of detecting evoked responses within random background fluctuations. Conventional methods to extract action potentials (spikes) from background noise often apply thresholds to the recorded signal, usually allowing reliable detection of spikes when data exhibit a good SNR, but often failing when SNR is poor. We here investigate a threshold-independent, fast, and automated procedure for analysis of low SNR data, based on fullwave-rectification and low-pass filtering the signal as a measure of the entire spiking activity (ESA). We investigate the sensitivity and reliability of the ESA-signal for detecting evoked responses by applying an automated receptive field (RF) mapping procedure to semi-chronically recorded data from primary visual cortex (V1) of five macaque monkeys. For recording sites with low SNR, the usage of ESA improved the detection rate of RFs by a factor of 2.5 in comparison to MUA-based detection. For recording sites with medium and high SNR, ESA delivered 30% more RFs than MUA. This significantly higher yield of ESA-based RF-detection still hold true when using an iterative procedure for determining the optimal spike threshold for each MUA individually. Moreover, selectivity measures for ESA-based RFs were quite compatible with MUA-based RFs. Regarding RF size, ESA delivered larger RFs than thresholded MUA, but size difference was consistent over all SNR fractions. Regarding orientation selectivity, ESA delivered more sites with significant orientation-dependent responses but with somewhat lower orientation indexes than MUA. However, preferred orientations were similar for both signal types. The results suggest that ESA is a powerful signal for applications requiring automated, fast, and reliable response detection, as e.g., brain-computer interfaces and neuroprosthetics, due to its high sensitivity and its independence from user-dependent intervention. Because the full information of the spiking activity is preserved, ESA also constitutes a valuable alternative for offline analysis of data with limited SNR.

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Bidirectional neural interfaces for multi-channel, high-density recording and electrical stimulation of neural activity in the central nervous system are fundamental tools for neuroscience and medical applications. Especially for clinical use, these electrical interfaces must be stable over several years, which is still a major challenge due to the foreign body response of neural tissue. A feasible solution to reduce this inflammatory response is to enable a free-floating implantation of high-density, silicon-based neural probes to avoid mechanical coupling between the skull and the cortex during brain micromotion. This paper presents our latest development of a reproducible microfabrication process, which allows a monolithic integration of a highly-flexible, polyimide-based cable with a silicon-stiffened neural probe at a high resolution of 1 µm. For a precise and complete insertion of the free-floating probes into the cortex, a new silicon-based, vacuum-actuated insertion tool is presented, which can be attached to commercially available electrode drives. To reduce the electrode impedance and enable safe and stable microstimulation an additional coating with the electrical conductive polymer PEDOT:PSS is used. The long-term stability of the presented free-floating neural probes is demonstrated in vitro and in vivo. The promising results suggest the feasibility of these neural probes for chronic applications.

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The need for fast and dynamic processing of relevant information imposes high demands onto the flexibility and efficiency of the nervous system. A good example for such flexibility is the attention-dependent selection of relevant sensory information. Studies investigating attentional modulations of neuronal responses to simultaneously arriving input showed that neurons respond, as if only the attended stimulus would be present within their receptive fields (RF). However, attention also improves neuronal representation and behavioral performance, when only one stimulus is present. Thus, attention serves for selecting relevant input and changes the neuronal processing of signals representing selected stimuli, ultimately leading to a more efficient behavioral performance. Here, we tested the hypothesis that attention configures the strength of functional coupling between a local neuronal network's neurons specifically for effective processing of signals representing attended stimuli. This coupling is measured as the strength of γ-synchronization between these neurons. The hypothesis predicts that the pattern of synchronization in local networks should depend on which stimulus is attended. Furthermore, we expect this pattern to be similar for the attended stimulus presented alone or together with irrelevant stimuli in the RF. To test these predictions, we recorded spiking-activity and local field potentials (LFP) with closely spaced electrodes in area V4 of monkeys performing a demanding attention task. Our results show that the γ-band phase coherence (γ-PhC) between spiking-activity and the LFP, as well as the spiking-activity of two groups of neurons, strongly depended on which of the two stimuli in the RF was attended. The γ-PhC was almost identical for the attended stimulus presented either alone or together with a distractor. The functional relevance of dynamic γ-band synchronization is further supported by the observation of strongly degraded γ-PhC before behavioral errors, while firing rates were barely affected. These qualitatively different results point toward a failure of attention-dependent top-down mechanisms to correctly synchronize the local neuronal network in V4, even though this network receives the correctly selected input. These findings support the idea of a flexible, demand-dependent dynamic configuration of local neuronal networks, for performing different functions, even on the same sensory input.

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Selective attention allows focusing on only part of the incoming sensory information. Neurons in the extrastriate visual cortex reflect such selective processing when different stimuli are simultaneously present in their large receptive fields. Their spiking response then resembles the response to the attended stimulus when presented in isolation. Unclear is where in the neuronal pathway attention intervenes to achieve such selective signal routing and processing. To investigate this question, we tagged two equivalent visual stimuli by independent broadband luminance noise and used the spectral coherence of these behaviorally irrelevant signals with the field potential of a local neuronal population in male macaque monkeys' area V4 as a measure for their respective causal influences. This new experimental paradigm revealed that signal transmission was considerably weaker for the not-attended stimulus. Furthermore, our results show that attention does not need to modulate responses in the input populations sending signals to V4 to selectively represent a stimulus, nor do they suggest a change of the V4 neurons' output gain depending on their feature similarity with the stimuli. Our results rather imply that selective attention uses a gating mechanism comprising the synaptic "inputs" that transmit signals from upstream areas into the V4 neurons. A minimal model implementing attention-dependent routing by gamma-band synchrony replicated the attentional gating effect and the signals' spectral transfer characteristics. It supports the proposal that selective interareal gamma-band synchrony subserves signal routing and explains our experimental finding that attention selectively gates signals already at the level of afferent synaptic input.SIGNIFICANCE STATEMENT Depending on the behavioral context, the brain needs to channel the flow of information through its networks of massively interconnected neurons. We designed an experiment that allows to causally assess routing of information originating from an attended object. We found that attention "gates" signals at the interplay between afferent fibers and the local neurons. A minimal model demonstrated that coherent gamma-rhythmic activity (∼60 Hz) between local neurons and their afferent-providing input neurons can realize the gating. Importantly, the attended signals did not need to be amplified already in an earlier processing stage, nor did they get amplified by a simple output response modulation. The method provides a useful tool to study mechanisms of dynamic network configuration underlying cognitive processes.

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Nonspatially selective attention is based on the notion that specific features or objects in the visual environment are effectively prioritized in cortical visual processing. Feature-based attention (FBA), in particular, is a well-studied process that dynamically and selectively addresses neurons preferentially processing the attended feature attribute (e.g., leftward motion). In everyday life, however, behavior may require high sensitivity for an entire feature dimension (e.g., motion), but experimental evidence for a feature dimension-specific attentional modulation on a cellular level is lacking. Therefore, we investigated neuronal activity in macaque motion-selective mediotemporal area (MT) in an experimental setting requiring the monkeys to detect either a motion change or a color change. We hypothesized that neural activity in MT is enhanced when the task requires perceptual sensitivity to motion. In line with this, we found that mean firing rates were higher in the motion task and that response variability and latency were lower compared with values in the color task, despite identical visual stimulation. This task-specific, dimension-based modulation of motion processing emerged already in the absence of visual input, was independent of the relation between the attended and stimulating motion direction, and was accompanied by a spatially global reduction of neuronal variability. The results provide single-cell support for the hypothesis of a feature dimension-specific top-down signal emphasizing the processing of an entire feature class.NEW & NOTEWORTHY Cortical processing serving visual perception prioritizes information according to current task requirements. We provide evidence in favor of a dimension-based attentional mechanism addressing all neurons that process visual information in the task-relevant feature domain. Behavioral tasks required monkeys to attend either color or motion, causing modulations of response strength, variability, latency, and baseline activity of motion-selective monkey area MT neurons irrespective of the attended motion direction but specific to the attended feature dimension.

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High-density electrocortical (ECoG) microelectrode arrays are promising signal-acquisition platforms for brain-computer interfaces envisioned, e.g., as high-performance communication solutions for paralyzed persons. We propose a multi-channel microelectrode array capable of recording ECoG field potentials with high spatial resolution. The proposed array is of a 150 mm2 total recording area; it has 124 circular electrodes (100, 300 and 500 µm in diameter) situated on the edges of concentric hexagons (min. 0.8 mm interdistance) and a skull-facing reference electrode (2.5 mm2 surface area). The array is processed as a free-standing device to enable monolithic integration of a rigid interposer, designed for soldering of fine-pitch SMD-connectors on a minimal assembly area. Electrochemical characterization revealed distinct impedance spectral bands for the 100, 300 and 500 µm-type electrodes, and for the array's own reference. Epidural recordings from the primary visual cortex (V1) of an awake Rhesus macaque showed natural electrophysiological signals and clear responses to standard visual stimulation. The ECoG electrodes of larger surface area recorded signals with greater spectral power in the gamma band, while the skull-facing reference electrode provided higher average gamma power spectral density (γPSD) than the common average referencing technique.

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Neurons in the middle temporal area (MT) respond to motion onsets and speed changes with a transient-sustained firing pattern. The latency of the transient response has recently been shown to correlate with reaction time in a speed change detection task, but it is not known how the sign, the amplitude, and the latency of this response depend on the sign and the magnitude of a speed change, and whether these transients can be decoded to explain speed change detection behavior. To investigate this issue, we measured the neuronal representation of a wide range of positive and negative speed changes in area MT of fixating macaques and obtained three major findings. First, speed change transients not only reflect a neuron's absolute speed tuning but are shaped by an additional gain that scales the tuned response according to the magnitude of a relative speed change. Second, by means of a threshold model positive and negative population transients of a moderate number of MT neurons explain detection of both positive and negative speed changes, respectively, at a level comparable to human detection rates under identical visual stimulation. Third, like reaction times in a psychophysical model of velocity detection, speed change response latencies follow a power-law function of the absolute difference of a speed change. Both this neuronal representation and its close correlation with behavioral measures of speed change detection suggest that neuronal transients in area MT facilitate the detection of rapid changes in visual input.

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BACKGROUND: Initially, human area MT+ was considered a visual area solely processing motion information but further research has shown that it is also involved in various different cognitive operations, such as working memory tasks requiring motion-related information to be maintained or cognitive tasks with implied or expected motion.In the present fMRI study in humans, we focused on MT+ modulation during working memory maintenance using a dynamic shape-tracking working memory task with no motion-related working memory content. Working memory load was systematically varied using complex and simple stimulus material and parametrically increasing retention periods. Activation patterns for the difference between retention of complex and simple memorized stimuli were examined in order to preclude that the reported effects are caused by differences in retrieval. RESULTS: Conjunction analysis over all delay durations for the maintenance of complex versus simple stimuli demonstrated a wide-spread activation pattern. Percent signal change (PSC) in area MT+ revealed a pattern with higher values for the maintenance of complex shapes compared to the retention of a simple circle and with higher values for increasing delay durations. CONCLUSIONS: The present data extend previous knowledge by demonstrating that visual area MT+ presents a brain activity pattern usually found in brain regions that are actively involved in working memory maintenance.

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Selective visual attention is known to be associated with characteristic modulations of neuronal activity in early visual cortex, but there is only rare evidence showing that these neuronal modulations are directly related to attention-dependent behavioral improvements. Here, we describe a strong, transient increase in the response of neurons in the mediotemporal (MT) area to behaviorally relevant speed changes that is not only modulated by attention but also highly correlated with the animal's performance. In trials with fast reaction time (RT), this transient component occurs with short latency, whereas latency increases monotonically with slower RT. Importantly, RTs are related not to the firing rate modulation during sustained attentive tracking of the target prior to the speed change but to the variability of the neuronal response. Our findings suggest a direct link between attention-dependent response modulations in early visual cortex and improved behavioral performance.

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Brain-computer interfaces have been proposed as a solution for paralyzed persons to communicate and interact with their environment. However, the neural signals used for controlling such prostheses are often noisy and unreliable, resulting in a low performance of real-world applications. Here we propose neural signatures of selective visual attention in epidural recordings as a fast, reliable, and high-performance control signal for brain prostheses. We recorded epidural field potentials with chronically implanted electrode arrays from two macaque monkeys engaged in a shape-tracking task. For single trials, we classified the direction of attention to one of two visual stimuli based on spectral amplitude, coherence, and phase difference in time windows fixed relative to stimulus onset. Classification performances reached up to 99.9%, and the information about attentional states could be transferred at rates exceeding 580 bits/min. Good classification can already be achieved in time windows as short as 200 ms. The classification performance changed dynamically over the trial and modulated with the task's varying demands for attention. For all three signal features, the information about the direction of attention was contained in the γ-band. The most informative feature was spectral amplitude. Together, these findings establish a novel paradigm for constructing brain prostheses as, for example, virtual spelling boards, promising a major gain in performance and robustness for human brain-computer interfaces.

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Receptive fields (RFs) of cortical sensory neurons increase in size along consecutive processing stages. When multiple stimuli are present in a large visual RF, a neuron typically responds to an attended stimulus as if only that stimulus were present. However, the mechanism by which a neuron selectively responds to a subset of its inputs while discarding all others is unknown. Here, we show that neurons can switch between subsets of their afferent inputs by highly specific modulations of interareal gamma-band phase-coherence (PC). We measured local field potentials, single- and multi-unit activity in two male macaque monkeys (Macaca mulatta) performing an attention task. Two small stimuli were placed on a screen; the stimuli were driving separate local V1 populations, while both were driving the same local V4 population. In each trial, we cued one of the two stimuli to be attended. We found that gamma-band PC of the local V4 population with multiple subpopulations of its V1 input was differentially modulated. It was high with the input subpopulation representing the attended stimulus, while simultaneously it was very low between the same V4 population and the other input-providing subpopulation representing the irrelevant stimulus. These differential modulations, which depend on stimulus relevance, were also found in the locking of spikes from V4 neurons to the gamma-band oscillations of the V1 input subpopulations. This rapid, highly specific interareal locking provides neurons with a powerful dynamic routing mechanism to select and process only the currently relevant signals.

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For processing and segmenting visual scenes, the brain is required to combine a multitude of features and sensory channels. It is neither known if these complex tasks involve optimal integration of information, nor according to which objectives computations might be performed. Here, we investigate if optimal inference can explain contour integration in human subjects. We performed experiments where observers detected contours of curvilinearly aligned edge configurations embedded into randomly oriented distractors. The key feature of our framework is to use a generative process for creating the contours, for which it is possible to derive a class of ideal detection models. This allowed us to compare human detection for contours with different statistical properties to the corresponding ideal detection models for the same stimuli. We then subjected the detection models to realistic constraints and required them to reproduce human decisions for every stimulus as well as possible. By independently varying the four model parameters, we identify a single detection model which quantitatively captures all correlations of human decision behaviour for more than 2000 stimuli from 42 contour ensembles with greatly varying statistical properties. This model reveals specific interactions between edges closely matching independent findings from physiology and psychophysics. These interactions imply a statistics of contours for which edge stimuli are indeed optimally integrated by the visual system, with the objective of inferring the presence of contours in cluttered scenes. The recurrent algorithm of our model makes testable predictions about the temporal dynamics of neuronal populations engaged in contour integration, and it suggests a strong directionality of the underlying functional anatomy.

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In monkeys, long-term recordings with chronically implanted microelectrodes frequently suffer from a continuously decreasing probability to record single units or even small multiunit clusters. This problem is associated with two technical limitations of the available devices: first, restrictions for electrode movement, and second, absent possibility to exchange electrodes easily on a regular basis. Permitting to adjust the recording site and to use new recording tracks with proper electrodes may avoid these problems and make chronic more similar to acute recordings. Here, we describe a novel type of implant tackling this issue. It consists of a new type of recording chamber combined with an exchangeable multielectrode array that precisely fits into it. The multielectrode array is reversibly fixed to the chamber, and within a minute it can be exchanged against another array equipped with new electrodes at the awake animal. The array allows for bidirectional movement of six electrodes for a distance of up to 12 mm. The recording chamber enables hermetical isolation of the intracranial space, resulting in long-lasting aseptic conditions and reducing dural thickening to a minimum, as confirmed by microbiological and histopathological analysis. The device has a simple design and is both easy to produce and low in cost. Functionality has been tested in primary and secondary visual cortex of three macaque monkeys over a period of up to 15 mo. The results show that even after more than a year, single and multiunit responses can be obtained with high incidence.

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Selective attention improves perception and modulates neuronal responses, but how attention-dependent changes of cortical activity improve the processing of attended objects is an open question. Changes in total signal strength or enhancements in signal-to-noise ratio have been proposed as putative mechanisms. However, it is still not clear whether, and to what extent, these processes contribute to the large perceptual improvements. We studied the ability to discriminate states of activity in visual cortex evoked by differently shaped objects depending on selective attention in monkeys. We found that gamma-band activity from V4 and V1 contains a high amount of information about stimulus shape, which increases for V4 recordings considerably with attention in successful trials, but not in case of behavioral errors. This effect resulted from enhanced differences between the stimulus-specific distributions of power spectral amplitudes. It could be explained neither by enhancements of signal-to-noise ratios, nor by changes in total signal power. Instead our results indicate that attention causes underlying cortical network states to become more distinct for different stimuli, providing a new neurophysiological explanation for improvements of behavioral performance by attention. The absence of the enhancement in discriminability in trials with behavioral errors demonstrates the relevance of this novel neural mechanism for perception.

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Feature-directed attention has been recently studied in various psychophysical, electrophysiological, and imaging studies. Convincing evidence has been obtained for its global effectiveness, but there is a debate about the processing fate of non-attended target features. A number of studies demonstrated feature-directed attention being associated with co-selection of non-relevant object features, thus resulting in selection of the entire object, whereas most other studies did not examine the extent to which processing of non-attended features was affected. Here, we present the results of two psychophysical experiments consisting of a Posner-like paradigm in which subjects were cued either to an individual feature or the entire object. We measured reaction times to changes in speed or colour of one of two simultaneously presented gratings. Our results strongly support the view that feature-based selection is a unique selection process different from object-based selection in that it can be associated with active suppression of non-relevant features.

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