The reentry hypothesis: The putative interaction of the frontal eye field, ventrolateral prefrontal cortex, and areas V4, IT for attention and eye movement

Attention is known to play a key role in perception, including action selection, object recognition and memory. Despite findings revealing competitive interactions among cell populations, attention remains difficult to explain. The central purpose of this paper is to link up a large number of findings in a single computational approach. Our simulation results suggest that attention can be well explained on a network level involving many areas of the brain. We argue that attention is an emergent phenomenon that arises from reentry and competitive interactions. We hypothesize that guided visual search requires the usage of an object-specific template in prefrontal cortex to sensitize V4 and IT cells whose preferred stimuli match the target template. This induces a feature-specific bias and provides guidance for eye movements. Prior to an eye movement, a spatially organized reentry from occulomotor centers, specifically the movement cells of the frontal eye field, occurs and modulates the gain of V4 and IT cells. The processes involved are elucidated by quantitatively comparing the time course of simulated neural activity with experimental data. Using visual search tasks as an example, we provide clear and empirically testable predictions for the participation of IT, V4 and the frontal eye field in attention. Finally, we explain a possible physiological mechanism that can lead to non-flat search slopes as the result of a slow, parallel discrimination process

Change blindness: eradication of gestalt strategies

Arrays of eight, texture-defined rectangles were used as stimuli in a one-shot change blindness (CB) task where there was a 50% chance that one rectangle would change orientation between two successive presentations separated by an interval. CB was eliminated by cueing the target rectangle in the first stimulus, reduced by cueing in the interval and unaffected by cueing in the second presentation. This supports the idea that a representation was formed that persisted through the interval before being 'overwritten' by the second presentation (Landman et al, 2003 Vision Research 43149–164]. Another possibility is that participants used some kind of grouping or Gestalt strategy. To test this we changed the spatial position of the rectangles in the second presentation by shifting them along imaginary spokes (by ±1 degree) emanating from the central fixation point. There was no significant difference seen in performance between this and the standard task [F(1,4)=2.565, p=0.185]. This may suggest two things: (i) Gestalt grouping is not used as a strategy in these tasks, and (ii) it gives further weight to the argument that objects may be stored and retrieved from a pre-attentional store during this task

Nägemistaju automaatsete protsesside eksperimentaalne uurimine

Väitekirja elektrooniline versioon ei sisalda publikatsiooneVäitekiri keskendub nägemistaju protsesside eksperimentaalsele uurimisele, mis on suuremal või vähemal määral automaatsed. Uurimistöös on kasutatud erinevaid eksperimentaalseid katseparadigmasid ja katsestiimuleid ning nii käitumuslikke- kui ka ajukuvamismeetodeid. Esimesed kolm empiirilist uurimust käsitlevad liikumisinformatsiooni töötlust, mis on evolutsiooni käigus kujunenud üheks olulisemaks baasprotsessiks nägemistajus. Esmalt huvitas meid, kuidas avastatakse liikuva objekti suunamuutusi, kui samal ajal toimub ka taustal liikumine (Uurimus I). Nägemistaju uurijad on pikka aega arvanud, et liikumist arvutatakse alati mõne välise objekti või tausta suhtes. Meie uurimistulemused ei kinnitanud taolise suhtelise liikumise printsiibi paikapidavust ning toetavad pigem seisukohta, et eesmärkobjekti liikumisinformatsiooni töötlus on automaatne protsess, mis tuvastab silma põhjas toimuvaid nihkeid, ja taustal toimuv seda eriti ei mõjuta. Teise uurimuse tulemused (Uurimus II) näitasid, et nägemissüsteem töötleb väga edukalt ka seda liikumisinformatsiooni, millele vaatleja teadlikult tähelepanu ei pööra. See tähendab, et samal ajal, kui inimene on mõne tähelepanu hõlmava tegevusega ametis, suudab tema aju taustal toimuvaid sündmusi automaatselt registreerida. Igapäevaselt on inimese nägemisväljas alati palju erinevaid objekte, millel on erinevad omadused, mistõttu järgmiseks huvitas meid (Uurimus III), kuidas ühe tunnuse (antud juhul värvimuutuse) töötlemist mõjutab mõne teise tunnusega toimuv (antud juhul liikumiskiiruse) muutus. Näitasime, et objekti liikumine parandas sama objekti värvimuutuse avastamist, mis viitab, et nende kahe omaduse töötlemine ajus ei ole päris eraldiseisev protsess. Samuti tähendab taoline tulemus, et hoolimata ühele tunnusele keskendumisest ei suuda inimene ignoreerida teist tähelepanu tõmbavat tunnust (liikumine), mis viitab taas kord automaatsetele töötlusprotsessidele. Neljas uurimus keskendus emotsionaalsete näoväljenduste töötlusele, kuna need kannavad keskkonnas hakkamasaamiseks vajalikke sotsiaalseid signaale, mistõttu on alust arvata, et nende töötlus on kujunenud suuresti automaatseks protsessiks. Näitasime, et emotsiooni väljendavaid nägusid avastati kiiremini ja kergemini kui neutraalse ilmega nägusid ning et vihane nägu tõmbas rohkem tähelepanu kui rõõmus (Uurimus IV). Väitekirja viimane osa puudutab visuaalset lahknevusnegatiivsust (ingl Visual Mismatch Negativity ehk vMMN), mis näitab aju võimet avastada automaatselt erinevusi enda loodud mudelist ümbritseva keskkonna kohta. Selle automaatse erinevuse avastamise mehhanismi uurimisse andsid oma panuse nii Uurimus II kui Uurimus IV, mis mõlemad pakuvad välja tõendusi vMMN tekkimise kohta eri tingimustel ja katseparadigmades ning ka vajalikke metodoloogilisi täiendusi. Uurimus V on esimene kogu siiani ilmunud temaatilist teadustööd hõlmav ülevaateartikkel ja metaanalüüs visuaalsest lahknevusnegatiivsusest psühhiaatriliste ja neuroloogiliste haiguste korral, mis panustab oluliselt visuaalse lahknevusnegatiivsuse valdkonna arengusse.The research presented and discussed in the thesis is an experimental exploration of processes in visual perception, which all display a considerable amount of automaticity. These processes are targeted from different angles using different experimental paradigms and stimuli, and by measuring both behavioural and brain responses. In the first three empirical studies, the focus is on motion detection that is regarded one of the most basic processes shaped by evolution. Study I investigated how motion information of an object is processed in the presence of background motion. Although it is widely believed that no motion can be perceived without establishing a frame of reference with other objects or motion on the background, our results found no support for relative motion principle. This finding speaks in favour of a simple and automatic process of detecting motion, which is largely insensitive to the surrounding context. Study II shows that the visual system is built to automatically process motion information that is outside of our attentional focus. This means that even if we are concentrating on some task, our brain constantly monitors the surrounding environment. Study III addressed the question of what happens when multiple stimulus qualities (motion and colour) are present and varied, which is the everyday reality of our visual input. We showed that velocity facilitated the detection of colour changes, which suggests that processing motion and colour is not entirely isolated. These results also indicate that it is hard to ignore motion information, and processing it is rather automatically initiated. The fourth empirical study focusses on another example of visual input that is processed in a rather automatic way and carries high survival value – emotional expressions. In Study IV, participants detected emotional facial expressions faster and more easily compared with neutral facial expressions, with a tendency towards more automatic attention to angry faces. In addition, we investigated the emergence of visual mismatch negativity (vMMN) that is one of the most objective and efficient methods for analysing automatic processes in the brain. Study II and Study IV proposed several methodological gains for registering this automatic change-detection mechanism. Study V is an important contribution to the vMMN research field as it is the first comprehensive review and meta-analysis of the vMMN studies in psychiatric and neurological disorders

Weakly Supervised Action Localization by Sparse Temporal Pooling Network

We propose a weakly supervised temporal action localization algorithm on untrimmed videos using convolutional neural networks. Our algorithm learns from video-level class labels and predicts temporal intervals of human actions with no requirement of temporal localization annotations. We design our network to identify a sparse subset of key segments associated with target actions in a video using an attention module and fuse the key segments through adaptive temporal pooling. Our loss function is comprised of two terms that minimize the video-level action classification error and enforce the sparsity of the segment selection. At inference time, we extract and score temporal proposals using temporal class activations and class-agnostic attentions to estimate the time intervals that correspond to target actions. The proposed algorithm attains state-of-the-art results on the THUMOS14 dataset and outstanding performance on ActivityNet1.3 even with its weak supervision.Comment: Accepted to CVPR 201

Salience-based selection: attentional capture by distractors less salient than the target

Current accounts of attentional capture predict the most salient stimulus to be invariably selected first. However, existing salience and visual search models assume noise in the map computation or selection process. Consequently, they predict the first selection to be stochastically dependent on salience, implying that attention could even be captured first by the second most salient (instead of the most salient) stimulus in the field. Yet, capture by less salient distractors has not been reported and salience-based selection accounts claim that the distractor has to be more salient in order to capture attention. We tested this prediction using an empirical and modeling approach of the visual search distractor paradigm. For the empirical part, we manipulated salience of target and distractor parametrically and measured reaction time interference when a distractor was present compared to absent. Reaction time interference was strongly correlated with distractor salience relative to the target. Moreover, even distractors less salient than the target captured attention, as measured by reaction time interference and oculomotor capture. In the modeling part, we simulated first selection in the distractor paradigm using behavioral measures of salience and considering the time course of selection including noise. We were able to replicate the result pattern we obtained in the empirical part. We conclude that each salience value follows a specific selection time distribution and attentional capture occurs when the selection time distributions of target and distractor overlap. Hence, selection is stochastic in nature and attentional capture occurs with a certain probability depending on relative salience

The iso-response method

Throughout the nervous system, neurons integrate high-dimensional input streams and transform them into an output of their own. This integration of incoming signals involves filtering processes and complex non-linear operations. The shapes of these filters and non-linearities determine the computational features of single neurons and their functional roles within larger networks. A detailed characterization of signal integration is thus a central ingredient to understanding information processing in neural circuits. Conventional methods for measuring single-neuron response properties, such as reverse correlation, however, are often limited by the implicit assumption that stimulus integration occurs in a linear fashion. Here, we review a conceptual and experimental alternative that is based on exploring the space of those sensory stimuli that result in the same neural output. As demonstrated by recent results in the auditory and visual system, such iso-response stimuli can be used to identify the non-linearities relevant for stimulus integration, disentangle consecutive neural processing steps, and determine their characteristics with unprecedented precision. Automated closed-loop experiments are crucial for this advance, allowing rapid search strategies for identifying iso-response stimuli during experiments. Prime targets for the method are feed-forward neural signaling chains in sensory systems, but the method has also been successfully applied to feedback systems. Depending on the specific question, “iso-response” may refer to a predefined firing rate, single-spike probability, first-spike latency, or other output measures. Examples from different studies show that substantial progress in understanding neural dynamics and coding can be achieved once rapid online data analysis and stimulus generation, adaptive sampling, and computational modeling are tightly integrated into experiments

Top-down effects on early visual processing in humans: a predictive coding framework

An increasing number of human electroencephalography (EEG) studies examining the earliest component of the visual evoked potential, the so-called C1, have cast doubts on the previously prevalent notion that this component is impermeable to top-down effects. This article reviews the original studies that (i) described the C1, (ii) linked it to primary visual cortex (V1) activity, and (iii) suggested that its electrophysiological characteristics are exclusively determined by low-level stimulus attributes, particularly the spatial position of the stimulus within the visual field. We then describe conflicting evidence from animal studies and human neuroimaging experiments and provide an overview of recent EEG and magnetoencephalography (MEG) work showing that initial V1 activity in humans may be strongly modulated by higher-level cognitive factors. Finally, we formulate a theoretical framework for understanding top-down effects on early visual processing in terms of predictive coding