Have you been there before? Decoding recognition of spatial scenes from fMRI signals in precuneus.

One potential application of forensic "brain reading" is to test whether a suspect has previously experienced a crime scene. Here, we investigated whether it is possible to decode real life autobiographic exposure to spatial locations using fMRI. In the first session, participants visited four out of eight possible rooms on a university campus. During a subsequent scanning session, subjects passively viewed pictures and videos from these eight possible rooms (four old, four novel) without giving any responses. A multivariate searchlight analysis was employed that trained a classifier to distinguish between "seen" versus "unseen" stimuli from a subset of six rooms. We found that bilateral precuneus encoded information that can be used to distinguish between previously seen and unseen rooms and that also generalized to the two stimuli left out from training. We conclude that activity in bilateral precuneus is associated with the memory of previously visited rooms, irrespective of the identity of the room, thus supporting a parietal contribution to episodic memory for spatial locations. Importantly, we could decode whether a room was visited in real life without the need of explicit judgments about the rooms. This suggests that recognition is an automatic response that can be decoded from fMRI data, thus potentially supporting forensic applications of concealed information tests for crime scene recognition.

Encoding of continuous perceptual choices in human early visual cortex.

Introduction: Research on the neural mechanisms of perceptual decision-making has typically focused on simple categorical choices, say between two alternative motion directions. Studies on such discrete alternatives have often suggested that choices are encoded either in a motor-based or in an abstract, categorical format in regions beyond sensory cortex. Methods: In this study, we used motion stimuli that could vary anywhere between 0° and 360° to assess how the brain encodes choices for features that span the full sensory continuum. We employed a combination of neuroimaging and encoding models based on Gaussian process regression to assess how either stimuli or choices were encoded in brain responses. Results: We found that single-voxel tuning patterns could be used to reconstruct the trial-by-trial physical direction of motion as well as the participants' continuous choices. Importantly, these continuous choice signals were primarily observed in early visual areas. The tuning properties in this region generalized between choice encoding and stimulus encoding, even for reports that reflected pure guessing. Discussion: We found only little information related to the decision outcome in regions beyond visual cortex, such as parietal cortex, possibly because our task did not involve differential motor preparation. This could suggest that decisions for continuous stimuli take can place already in sensory brain regions, potentially using similar mechanisms to the sensory recruitment in visual working memory.

Clarifying the nature of stochastic fluctuations and accumulation processes in spontaneous movements.

Experiments on choice-predictive brain signals have played an important role in the debate on free will. In a seminal study, Benjamin Libet and colleagues found that a negative-going EEG signal, the readiness potential (RP), can be observed over motor-related brain regions even hundreds of ms before the time of the conscious decision to move. If the early onset of the readiness potential is taken as an indicator of the "brain's decision to move" this could mean that this decision is made early, by unconscious brain activity, rather than later, at the time when the subject believes to have decided. However, an alternative kind of interpretation, involving ongoing stochastic fluctuations, has recently been brought to light. This stochastic decision model (SDM) takes its inspiration from leaky accumulator models of perceptual decision making. It suggests that the RP originates from an accumulation of ongoing stochastic fluctuations. In this view, the decision happens only at a much later stage when an accumulated noisy signal (plus imperative) reaches a threshold. Here, we clarify a number of confusions regarding both the evidence for the stochastic decision model as well as the interpretation that it offers. We will explore several points that we feel are in need of clarification: (a) the empirical evidence for the role of stochastic fluctuations is so far only indirect; (b) the interpretation of animal studies is unclear; (c) a model that is deterministic during the accumulation stage can explain the data in a similar way; (d) the primary focus in the literature has been on the role of random fluctuations whereas the deterministic aspects of the model have been largely ignored; (e) contrary to the original interpretation, the deterministic component of the model is quantitatively the dominant input into the accumulator; and finally (f) there is confusion regarding the role of "imperative" in the SDM and its link to "evidence" in perceptual decision making. Our aim is not to rehabilitate the role of the RP in the free will debate. Rather we aim to address some confusions regarding the evidence for accumulators playing a role in these preparatory brain processes.

Instruction effects on randomness in sequence generation.

Randomness is a fundamental property of human behavior. It occurs both in the form of intrinsic random variability, say when repetitions of a task yield slightly different behavioral outcomes, or in the form of explicit randomness, say when a person tries to avoid being predicted in a game of rock, paper and scissors. Randomness has frequently been studied using random sequence generation tasks (RSG). A key finding has been that humans are poor at deliberately producing random behavior. At the same time, it has been shown that people might be better randomizers if randomness is only an implicit (rather than an explicit) requirement of the task. We therefore hypothesized that randomization performance might vary with the exact instructions with which randomness is elicited. To test this, we acquired data from a large online sample (n = 388), where every participant made 1,000 binary choices based on one of the following instructions: choose either randomly, freely, irregularly, according to an imaginary coin toss or perform a perceptual guessing task. Our results show significant differences in randomness between the conditions as quantified by conditional entropy and estimated Markov order. The randomization scores were highest in the conditions where people were asked to be irregular or mentally simulate a random event (coin toss) thus yielding recommendations for future studies on randomization behavior.

Psychophysics and computational modeling of feature-continuous motion perception.

Sensory decision-making is frequently studied using categorical tasks, even though the feature space of most stimuli is continuous. Recently, it has become more common to measure feature perception in a gradual fashion, say when studying motion perception across the full space of directions. However, continuous reports can be contaminated by perceptual or motor biases. Here, we examined such biases on perceptual reports by comparing two response methods. With the first method, participants reported motion direction in a motor reference frame by moving a trackball. With the second method, participants used a perceptual frame of reference with a perceptual comparison stimulus. We tested biases using three different versions of random dot kinematograms. We found strong and systematic biases in responses when reporting the direction in a motor frame of reference. For the perceptual frame of reference, these systematic biases were not evident. Independent of the response method, we also detected a systematic misperception where subjects sometimes confuse the physical stimulus direction with its opposite direction. This was confirmed using a von Mises mixture model that estimated the contribution of veridical perception, misperception, and guessing. Importantly, the more sensitive perceptual reporting method revealed that, with increasing levels of sensory evidence, perceptual performance increases not only in the form of higher detection probability, but under certain conditions also in the form of increased precision.

Sensory-Tactile Functional Mapping and Use-Associated Structural Variation of the Human Female Genital Representation Field.

The precise location of the human female genital representation field in the primary somatosensory cortex (S1) is controversial and its capacity for use-associated structural variation as a function of sexual behavior remains unknown. We used a functional magnetic resonance imaging (fMRI)-compatible sensory-tactile stimulation paradigm to functionally map the location of the female genital representation field in 20 adult women. Neural response to tactile stimulation of the clitoral region (vs right hand) identified individually-diverse focal bilateral activations in dorsolateral areas of S1 (BA1-BA3) in alignment with anatomic location. We next used cortical surface analyses to assess structural thickness across the 10 individually most activated vertices per hemisphere for each woman. We show that frequency of sexual intercourse within 12 months is correlated with structural thickness of the individually-mapped left genital field. Our results provide a precise functional localization of the female genital field and provide support for use-associated structural variation of the human genital cortex.SIGNIFICANCE STATEMENT We provide a precise location of the human female genital field in the somatosensory cortex and, for the first time, provide evidence in support of structural variation of the human genital field in association with frequency of genital contact. Our study represents a significant methodological advance by individually mapping genital fields for structural analyses. On a secondary level, our results suggest that any study investigating changes in the human genital field must map the field individually to achieve sufficient precision. Our results pave the way for future research into the plasticity of the human genital cortex as a function of normal or adverse experience as well as changes in pathologic conditions, i.e., sexual dysfunction, sexual deviation, or sexual risk-taking behavior.

Brains in dialogue: decoding neural preparation of speaking to a conversational partner.

In dialogue, language processing is adapted to the conversational partner. We hypothesize that the brain facilitates partner-adapted language processing through preparatory neural configurations (task sets) that are tailored to the conversational partner. In this experiment, we measured neural activity with functional magnetic resonance imaging (fMRI) while healthy participants in the scanner (a) engaged in a verbal communication task with a conversational partner outside of the scanner, or (b) spoke outside of a conversational context (to test the microphone). Using multivariate searchlight analysis, we identify cortical regions that represent information on whether speakers plan to speak to a conversational partner or without having a partner. Most notably a region that has been associated with processing social-affective information and perspective taking, the ventromedial prefrontal cortex, as well as regions that have been associated with prospective task representation, the bilateral ventral prefrontal cortex, are involved in encoding the speaking condition. Our results suggest that speakers prepare, in advance of speaking, for the social context in which they will speak.

Internal and external attention and the default mode network.

Focused attention meditations have been shown to improve psychological health and wellbeing and are nowadays an integral part of many psychotherapies. While research on the neural correlates of focused attention meditation is increasing, findings vary on whether meditations are associated with high or low activity in the default mode network (DMN). To clarify the relationship between focused attention meditation and the activity in DMN regions, it may be helpful to distinguish internal and external attention as well as different phases within one meditation: During focused attention meditation, the practitioner switches between mindful attention, mind-wandering and refocusing. Here, we employed a thought-probe paradigm to study the neural correlates of these different phases. Twenty healthy, meditation naïve participants were introduced to external (mindfulness of sound) and internal (mindfulness of breathing) attention meditation and then practiced the meditation at home for four consecutive days. They then performed the same focused attention meditations during fMRI scanning, in four runs alternating between internal and external attention. At pseudorandom intervals, participants were asked whether they had just been focused on the task (mindful attention) or had been distracted (mind-wandering). During mindful attention, brain regions typically associated with the DMN, such as the medial prefrontal cortex, posterior cingulate cortex and left temporoparietal junction showed significantly less neural activation compared to mind-wandering phases. Reduced activity of the DMN was found during both external and internal attention, with stronger deactivation in the posterior cingulate cortex during internal attention compared to external attention. Moreover, refocusing after mind-wandering was associated with activity in the left inferior frontal gyrus. Our results support the theory that mindful attention is associated with reduced DMN activity compared to mind-wandering, independent of the practitioner's attention focus (i.e., internal vs. external).

Default Network Activity Is Associated with Better Performance in a Vigilance Task.

When attention has to be maintained over prolonged periods performance slowly fluctuates and errors can occur. It has been shown that lapses of attention are correlated with BOLD signals in frontal and parietal cortex. This raises the question how attentional fluctuations are linked to the fronto-parietal default network. Because the attentional state fluctuates slowly we expect that potential links between attentional fluctuations and brain activity should be observable on longer time scales and importantly also before the execution of the task. In the present study we used fMRI to identify brain activity that is correlated with vigilance, defined as fluctuations of reaction times (RT) during a sustained attention task. We found that brain activity in visual cortex, parietal lobe (PL), inferior and superior frontal gyrus, and supplementary motor area (SMA) was higher when the subject had a relatively long RT. In contrast to our expectations, activity in the default network (DN) was higher when subjects had a relatively short RT, that means when the performance was improved. This modulation in the DN was present already several seconds before the task execution, thus pointing to activity in the DN as a potential cause of performance increases in simple repetitive tasks.

Am I seeing myself, my friend or a stranger? The role of personal familiarity in visual distinction of body identities in the human brain.

Several brain regions appear to play a role in representing different body identities. The specific contribution of each of these regions is still unclear, however. Here we investigated which brain areas enable the visual distinction between self and other bodies of different familiarity, and between familiar and unfamiliar other individuals, and moreover, where identity-specific information on the three individuals was encoded. Participants were confronted with standardized headless human body stimuli either showing the participant's own, a personally familiar or an unfamiliar other body, while performing a luminance discrimination task. Employing multivariate pattern analysis, we were able to identify areas that allowed for the distinction of self from personal familiar other bodies within the medial prefrontal cortex (mPFC) and posterior cingulate cortex/precuneus. Successful distinction of self from unfamiliar others was possible in the left middle frontal gyrus, the right inferior frontal gyrus, the left pre-supplementary motor area and the right putamen. Personally familiar others could be distinguished from unfamiliar others in the right temporoparietal junction (TPJ). An analysis of identity-specific information revealed a spatial gradient ranging from inferior posterior to superior anterior portions of the mPFC that was associated with encoding identity-related information for self via familiar to unfamiliar other bodies, respectively. Furthermore, several midline and frontal regions encoded information on more than one identity. The TPJ's role in deviance detection was underlined, as only identity-specific information on unfamiliar others was encoded here. Together, our findings suggest substantial spatial overlap in neural correlates of self and other body representation and thus, support the hypothesis of a socially-related representation of the self.

Neural coding of assessing another person's knowledge based on nonverbal cues.

For successful communication, conversational partners need to estimate each other's current knowledge state. Nonverbal facial and bodily cues can reveal relevant information about how confident a speaker is about what they are saying. Using functional magnetic resonance imaging, we aimed to identify brain regions that encode how confident a speaker is perceived to be. Participants viewed videos of people answering general knowledge questions and judged each respondent's confidence in their answer. Our results suggest a distinct role of two neural networks known to support social inferences, the so-called mentalizing and the mirroring network. While activation in both networks underlies the processing of nonverbal cues, only activity in the mentalizing network, most notably the medial prefrontal cortex and the bilateral temporoparietal junction, is modulated by how confident the respondent is judged to be. Our results support an integrative account of the mirroring and mentalizing network, in which the two systems support each other in aiding pragmatic processing.

Decoding vigilance with NIRS.

Sustained, long-term cognitive workload is associated with variations and decrements in performance. Such fluctuations in vigilance can be a risk factor especially during dangerous attention demanding activities. Functional MRI studies have shown that attentional performance is correlated with BOLD-signals, especially in parietal and prefrontal cortical regions. An interesting question is whether these BOLD-signals could be measured in real-world scenarios, say to warn in a dangerous workplace whenever a subjects' vigilance is low. Because fMRI lacks the mobility needed for such applications, we tested whether the monitoring of vigilance might be possible using Near-Infrared Spectroscopy (NIRS). NIRS is a highly mobile technique that measures hemodynamics in the surface of the brain. We demonstrate that non-invasive NIRS signals correlate with vigilance. These signals carry enough information to decode subjects' reaction times at a single trial level.

Dissociation between saliency signals and activity in early visual cortex.

Saliency is a measure that describes how attention is guided by local stimulus properties. Some hypotheses assign its computation to specific topographically organized areas of early human visual cortex. However, in most stimuli, saliency is correlated with luminance contrast, which in turn is known to correlate with activity in these early areas. Thus, any observed correlation of local activity with saliency might be due to the area encoding luminance contrast. Here we disentangle encoding of local luminance contrast and saliency by using stimuli where the two properties are uncorrelated. First, we conducted an eye-tracking study to verify that both negative and positive contrast modifications located in individual quadrants of the visual field increase saliency. Second, subjects viewed identical stimuli while fMRI signals were recorded. We find that positive contrast modifications induce a robust increase of activity in V1-V3 and hV4. However, negative contrast modifications lead to a reduced (V1, V2) or comparable (V3, hV4) activity level compared to unmodified quadrants. Furthermore, even with linear multivariate pattern-classification techniques, it is not possible to decode the location of the salient quadrant independent of the type of the contrast modification. Instead, decoding of the contrast-modified location is only possible separately for the two modification types in V1-V3. These findings suggest that the BOLD activity in V1-V3 is dominated by contrast-dependent processes and does not include the contrast invariance necessary for the computation of feature-invariant saliency.

Orientation pop-out processing in human visual cortex.

Visual stimuli can "pop out" if they are different to their background. There has been considerable debate as to the role of primary visual cortex (V1) versus higher visual areas (esp. V4) in pop-out processing. Here we parametrically modulated the relative orientation of stimuli and their backgrounds to investigate the neural correlates of pop-out in visual cortex while subjects were performing a demanding fixation task in a scanner. Whole brain and region of interest analyses confirmed a representation of orientation contrast in extrastriate visual cortex (V4), but not in striate visual cortex (V1). Thus, although previous studies have shown that human V1 can be involved in orientation pop-out, our findings demonstrate that there are cases where V1 is "blind" and pop-out detection is restricted to higher visual areas. Pop-out processing is presumably a distributed process across multiple visual regions.

Similar neural mechanisms for perceptual guesses and free decisions.

When facing perceptual choices under challenging conditions we might believe to be purely guessing. But which brain regions determine the outcome of our guesses? One possibility is that higher-level, domain-general brain regions might help break the symmetry between equal-appearing choices. Here we directly investigated whether perceptual guesses share brain networks with other types of free decisions. We trained an fMRI-based pattern classifier to distinguish between two perceptual guesses and tested whether it was able to predict the outcome of similar non-perceptual choices, as in conventional free choice tasks. Activation patterns in the medial posterior parietal cortex cross-predicted free decisions from perceptual guesses and vice versa. This inter-changeability strongly speaks for a similar neural code for both types of decisions. The posterior parietal cortex might be part of a domain-general system that helps resolve decision conflicts when no choice option is more or less likely or valuable, thus preventing behavioural stalemate.

The neural encoding of guesses in the human brain.

Human perception depends heavily on the quality of sensory information. When objects are hard to see we often believe ourselves to be purely guessing. Here we investigated whether such guesses use brain networks involved in perceptual decision making or independent networks. We used a combination of fMRI and pattern classification to test how visibility affects the signals, which determine choices. We found that decisions regarding clearly visible objects are predicted by signals in sensory brain regions, whereas different regions in parietal cortex became predictive when subjects were shown invisible objects and believed themselves to be purely guessing. This parietal network was highly overlapping with regions, which have previously been shown to encode free decisions. Thus, the brain might use a dedicated network for determining choices when insufficient sensory information is available.

Decoding successive computational stages of saliency processing.

An important requirement for vision is to identify interesting and relevant regions of the environment for further processing. Some models assume that salient locations from a visual scene are encoded in a dedicated spatial saliency map [1, 2]. Then, a winner-take-all (WTA) mechanism [1, 2] is often believed to threshold the graded saliency representation and identify the most salient position in the visual field. Here we aimed to assess whether neural representations of graded saliency and the subsequent WTA mechanism can be dissociated. We presented images of natural scenes while subjects were in a scanner performing a demanding fixation task, and thus their attention was directed away. Signals in early visual cortex and posterior intraparietal sulcus (IPS) correlated with graded saliency as defined by a computational saliency model. Multivariate pattern classification [3, 4] revealed that the most salient position in the visual field was encoded in anterior IPS and frontal eye fields (FEF), thus reflecting a potential WTA stage. Our results thus confirm that graded saliency and WTA-thresholded saliency are encoded in distinct neural structures. This could provide the neural representation required for rapid and automatic orientation toward salient events in natural environments.

The extended language network: a meta-analysis of neuroimaging studies on text comprehension.

Language processing in context requires more than merely comprehending words and sentences. Important subprocesses are inferences for bridging successive utterances, the use of background knowledge and discourse context, and pragmatic interpretations. The functional neuroanatomy of these text comprehension processes has only recently been investigated. Although there is evidence for right-hemisphere contributions, reviews have implicated the left lateral prefrontal cortex, left temporal regions beyond Wernicke's area, and the left dorso-medial prefrontal cortex (dmPFC) for text comprehension. To objectively confirm this extended language network and to evaluate the respective contribution of right hemisphere regions, meta-analyses of 23 neuroimaging studies are reported here. The analyses used replicator dynamics based on activation likelihood estimates. Independent of the baseline, the anterior temporal lobes (aTL) were active bilaterally. In addition, processing of coherent compared with incoherent text engaged the dmPFC and the posterior cingulate cortex. Right hemisphere activations were seen most notably in the analysis of contrasts testing specific subprocesses, such as metaphor comprehension. These results suggest task dependent contributions for the lateral PFC and the right hemisphere. Most importantly, they confirm the role of the aTL and the fronto-medial cortex for language processing in context.