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1.  Negative learning bias is associated with risk aversion in a genetic animal model of depression 
The lateral habenula (LHb) is activated by aversive stimuli and the omission of reward, inhibited by rewarding stimuli and is hyperactive in helpless rats—an animal model of depression. Here we test the hypothesis that congenital learned helpless (cLH) rats are more sensitive to decreases in reward size and/or less sensitive to increases in reward than wild-type (WT) control rats. Consistent with the hypothesis, we found that cLH rats were slower to switch preference between two responses after a small upshift in reward size on one of the responses but faster to switch their preference after a small downshift in reward size. cLH rats were also more risk-averse than WT rats—they chose a response delivering a constant amount of reward (“safe” response) more often than a response delivering a variable amount of reward (“risky” response) compared to WT rats. Interestingly, the level of bias toward negative events was associated with the rat's level of risk aversion when compared across individual rats. cLH rats also showed impaired appetitive Pavlovian conditioning but more accurate responding in a two-choice sensory discrimination task. These results are consistent with a negative learning bias and risk aversion in cLH rats, suggesting abnormal processing of rewarding and aversive events in the LHb of cLH rats.
doi:10.3389/fnhum.2014.00001
PMCID: PMC3893716  PMID: 24474914
lateral habenula; reinforcement learning; depression; helplessness; cLH; risk aversion; reward; behavior
2.  Reduced Frontal Brain Volume in Non-Treatment-Seeking Cocaine-Dependent Individuals: Exploring the Role of Impulsivity, Depression, and Smoking 
In cocaine-dependent patients, gray matter (GM) volume reductions have been observed in the frontal lobes that are associated with the duration of cocaine use. Studies are mostly restricted to treatment-seekers and studies in non-treatment-seeking cocaine abusers are sparse. Here, we assessed GM volume differences between 30 non-treatment-seeking cocaine-dependent individuals and 33 non-drug using controls using voxel-based morphometry. Additionally, within the group of non-treatment-seeking cocaine-dependent individuals, we explored the role of frequently co-occurring features such as trait impulsivity (Barratt Impulsivity Scale, BIS), smoking, and depressive symptoms (Beck Depression Inventory), as well as the role of cocaine use duration, on frontal GM volume. Smaller GM volumes in non-treatment-seeking cocaine-dependent individuals were observed in the left middle frontal gyrus. Moreover, within the group of cocaine users, trait impulsivity was associated with reduced GM volume in the right orbitofrontal cortex, the left precentral gyrus, and the right superior frontal gyrus, whereas no effect of smoking severity, depressive symptoms, or duration of cocaine use was observed on regional GM volumes. Our data show an important association between trait impulsivity and frontal GM volumes in cocaine-dependent individuals. In contrast to previous studies with treatment-seeking cocaine-dependent patients, no significant effects of smoking severity, depressive symptoms, or duration of cocaine use on frontal GM volume were observed. Reduced frontal GM volumes in non-treatment-seeking cocaine-dependent subjects are associated with trait impulsivity and are not associated with co-occurring nicotine dependence or depression.
doi:10.3389/fnhum.2014.00007
PMCID: PMC3894477  PMID: 24478673
cocaine dependence; drug abuse; voxel-based morphometry; frontal; depression; nicotine
3.  Visual Evoked Potential and Magnetic Resonance Imaging are More Effective Markers of Multiple Sclerosis Progression than Laser Polarimetry with Variable Corneal Compensation 
Background: The aim of our study was to assess the role of laser polarimetry and visual evoked potentials (VEP) as potential biomarkers of disease progression in multiple sclerosis (MS).
Participants: A total of 41 patients with MS (82 eyes) and 22 age-related healthy volunteers (44 eyes) completed the study. MS patients were divided into two groups, one (ON) with a history of optic neuritis (17 patients, 34 eyes) and another group (NON) without it (24 patients, 48 eyes). The MS patients and controls underwent laser polarimetry (GDx) examination of the retinal nerve fiber layer (RNFL). In the MS group, we also examined: Kurtzke “expanded disability status scale” (EDSS), the duration of the disorder, VEP – latency and amplitude, and conventional brain magnetic resonance imaging (MRI). Our results were statistically analyzed using ANOVA, Mann–Whitney, and Spearman correlation analyses.
Results: In the MS group, brain atrophy and new T2 brain lesions in MRI correlated with both VEP latencies and amplitudes. Separate comparisons revealed VEP latency testing to be less sensitive in ON than in NON-patients. In ON patients, VEP amplitudes correlated mildly with brain atrophy (r = −0.15) and strongly with brain new MRI lesions (r = −0.8). In NON-patients, highly significant correlation of new MRI brain lesions with VEP latencies (r = 0.63, r = 0.6) and amplitudes (r = −0.3, r = −4.2) was found. EDSS also correlated with brain atrophy in this group (r = 0.5). Our study did not find a correlation of GDx measures with MRI tests. The GDx method was not able to detect whole brain demyelinization and the degeneration process, but was only able to reveal the involvement of optic nerves in ON and NON-patients.
Conclusion: In our study, we found that both methods (VEP and GDx) can be used for the detection of optic nerve damage, but VEP was found to be superior in evaluating whole brain demyelinization and axonal degeneration. Both VEP and MRI, but not GDx, have an important role in monitoring disease progression in MS patients, independent of the ON history.
doi:10.3389/fnhum.2014.00010
PMCID: PMC3898055  PMID: 24478676
VEP; laser polarimetry of optic nerve; MRI; multiple sclerosis; demyelinization and axonal degeneration
4.  Multiple neural states of representation in short-term memory? It’s a matter of attention 
Short-term memory (STM) refers to the capacity-limited retention of information over a brief period of time, and working memory (WM) refers to the manipulation and use of that information to guide behavior. In recent years it has become apparent that STM and WM interact and overlap with other cognitive processes, including attention (the selection of a subset of information for further processing) and long-term memory (LTM—the encoding and retention of an effectively unlimited amount of information for a much longer period of time). Broadly speaking, there have been two classes of memory models: systems models, which posit distinct stores for STM and LTM (Atkinson and Shiffrin, 1968; Baddeley and Hitch, 1974); and state-based models, which posit a common store with different activation states corresponding to STM and LTM (Cowan, 1995; McElree, 1996; Oberauer, 2002). In this paper, we will focus on state-based accounts of STM. First, we will consider several theoretical models that postulate, based on considerable behavioral evidence, that information in STM can exist in multiple representational states. We will then consider how neural data from recent studies of STM can inform and constrain these theoretical models. In the process we will highlight the inferential advantage of multivariate, information-based analyses of neuroimaging data (fMRI and electroencephalography (EEG)) over conventional activation-based analysis approaches (Postle, in press). We will conclude by addressing lingering questions regarding the fractionation of STM, highlighting differences between the attention to information vs. the retention of information during brief memory delays.
doi:10.3389/fnhum.2014.00005
PMCID: PMC3899521  PMID: 24478671
short-term memory; attention; representational states; multivariate pattern analysis; unattended memory items
5.  Primacy and recency effects as indices of the focus of attention 
Ongoing debate surrounds the capacity and characteristics of the focus of attention. The present study investigates whether a pattern of larger recency effects and smaller primacy effects reported in previous working memory studies is specific to task conditions used in those studies, or generalizes across manipulations of task-demand. Two experiments varied task-demands by requiring participants to remember lists of letters and to then respond to a subsequent two-item probe by indicating either the item that was presented later in the list (judgment of recency) or the item was presented earlier (judgment of primacy). Analyses tested the prediction that a WM task emphasizing later items in a list (judgment of recency) would encourage exaggerated recency effects and attenuated primacy effects, while a task emphasizing earlier items (judgment of primacy) would encourage exaggerated primacy effects and attenuated recency effects. Behavioral results from two experiments confirmed this prediction. In contrast to past studies, fMRI contrasts revealed no brain regions where activity was significantly altered by the presence of recency items in the probe, for either task condition. However, presence of the primacy item in the probe significantly influenced activity in frontal lobe brain regions linked to active maintenance, but the location and direction of activation changes varied as a function of task instructions. In sum, two experiments demonstrate that the behavioral and neural signatures of WM, specifically related to primacy and recency effects, are dependent on task-demands. Findings are discussed as they inform models of the structure and capacity of WM.
doi:10.3389/fnhum.2014.00006
PMCID: PMC3900765  PMID: 24478672
working memory; focus of attention; primacy and recency effects; phonological rehearsal; medial temporal lobe
6.  Visual evoked potentials to change in coloration of a moving bar 
In our previous study we found that it takes less time to detect coloration change in a moving object compared to coloration change in a stationary one (Kreegipuu etal., 2006). Here, we replicated the experiment, but in addition to reaction times (RTs) we measured visual evoked potentials (VEPs), to see whether this effect of motion is revealed at the cortical level of information processing. We asked our subjects to detect changes in coloration of stationary (0°/s) and moving bars (4.4 and 17.6°/s). Psychophysical results replicate the findings from the previous study showing decreased RTs to coloration changes with increase of velocity of the color changing stimulus. The effect of velocity on VEPs was opposite to the one found on RTs. Except for component N1, the amplitudes of VEPs elicited by the coloration change of faster moving objects were reduced than those elicited by the coloration change of slower moving or stationary objects. The only significant effect of velocity on latency of peaks was found for P2 in frontal region. The results are discussed in the light of change-to-change interval and the two methods reflecting different processing mechanisms.
doi:10.3389/fnhum.2014.00019
PMCID: PMC3900876  PMID: 24478683
motion; velocity; color change; reaction time; visual evoked potentials
7.  Brain and intersubjectivity: a Hegelian hypothesis on the self-other neurodynamics 
doi:10.3389/fnhum.2014.00011
PMCID: PMC3901003  PMID: 24478677
intersubjectivity; self-consciousness; default mode network; mirror neuron system; self concept; dehumanization; Hegel
8.  Short-term retention of relational memory in amnesia revisited: accurate performance depends on hippocampal integrity 
Traditionally, it has been proposed that the hippocampus and adjacent medial temporal lobe cortical structures are selectively critical for long-term declarative memory, which entails memory for inter-item and item-context relationships. Whether the hippocampus might also contribute to short-term retention of relational memory representations has remained controversial. In two experiments, we revisit this question by testing memory for relationships among items embedded in scenes using a standard working memory trial structure in which a sample stimulus is followed by a brief delay and the corresponding test stimulus. In each experimental block, eight trials using different exemplars of the same scene were presented. The exemplars contained the same items but with different spatial relationships among them. By repeating the pictures across trials, any potential contributions of item or scene memory to performance were minimized, and relational memory could be assessed more directly than has been done previously. When test displays were presented, participants indicated whether any of the item-location relationships had changed. Then, regardless of their responses (and whether any item did change its location), participants indicated on a forced-choice test, which item might have moved, guessing if necessary. Amnesic patients were impaired on the change detection test, and were frequently unable to specify the change after having reported correctly that a change had taken place. Comparison participants, by contrast, frequently identified the change even when they failed to report the mismatch, an outcome that speaks to the sensitivity of the change specification measure. These results confirm past reports of hippocampal contributions to short-term retention of relational memory representations, and suggest that the role of the hippocampus in memory has more to do with relational memory requirements than the length of a retention interval.
doi:10.3389/fnhum.2014.00016
PMCID: PMC3901041  PMID: 24478681
amnesia; hippocampus; short-term memory; relational memory; delayed recognition
9.  Lexical selection in the semantically blocked cyclic naming task: the role of cognitive control and learning 
Studies of semantic interference in language production have provided evidence for a role of cognitive control mechanisms in regulating the activation of semantic competitors during naming. The present study investigated the relationship between individual differences in cognitive control abilities, for both younger and older adults, and the degree of semantic interference in a blocked cyclic naming task. We predicted that individuals with lower working memory capacity (as measured by word span), lesser ability to inhibit distracting responses (as measured by Stroop interference), and a lesser ability to resolve proactive interference (as measured by a recent negatives task) would show a greater increase in semantic interference in naming, with effects being larger for older adults. Instead, measures of cognitive control were found to relate to specific indices of semantic interference in the naming task, rather than overall degree of semantic interference, and few interactions with age were found, with younger and older adults performing similarly. The increase in naming latencies across naming trials within a cycle was negatively correlated with word span for both related and unrelated conditions, suggesting a strategy of narrowing response alternatives based upon memory for the set of item names. Evidence for a role of inhibition in response selection was obtained, as Stroop interference correlated positively with the change in naming latencies across cycles for the related, but not unrelated, condition. In contrast, recent negatives interference correlated negatively with the change in naming latencies across unrelated cycles, suggesting that individual differences in this tap the degree of strengthening of links in a lexical network based upon prior exposure. Results are discussed in terms of current models of lexical selection and consequences for word retrieval in more naturalistic production.
doi:10.3389/fnhum.2014.00009
PMCID: PMC3902204  PMID: 24478675
lexical selection; semantic blocking; aging; inhibition; individual differences
10.  Habenular expression of rare missense variants of the β4 nicotinic receptor subunit alters nicotine consumption 
The CHRNA5-CHRNA3-CHRNB4 gene cluster, encoding the α5, α3, and β4 nicotinic acetylcholine receptor (nAChR) subunits, has been linked to nicotine dependence. The habenulo-interpeduncular (Hb-IPN) tract is particularly enriched in α3β4 nAChRs. We recently showed that modulation of these receptors in the medial habenula (MHb) in mice altered nicotine consumption. Given that β4 is rate-limiting for receptor activity and that single nucleotide polymorphisms (SNPs) in CHRNB4 have been linked to altered risk of nicotine dependence in humans, we were interested in determining the contribution of allelic variants of β4 to nicotine receptor activity in the MHb. We screened for missense SNPs that had allele frequencies >0.0005 and introduced the corresponding substitutions in Chrnb4. Fourteen variants were analyzed by co-expression with α3. We found that β4A90I and β4T374I variants, previously shown to associate with reduced risk of smoking, and an additional variant β4D447Y, significantly increased nicotine-evoked current amplitudes, while β4R348C, the mutation most frequently encountered in sporadic amyotrophic lateral sclerosis (sALS), showed reduced nicotine currents. We employed lentiviruses to express β4 or β4 variants in the MHb. Immunoprecipitation studies confirmed that β4 lentiviral-mediated expression leads to specific upregulation of α3β4 but not β2 nAChRs in the Mhb. Mice injected with the β4-containing virus showed pronounced aversion to nicotine as previously observed in transgenic Tabac mice overexpressing Chrnb4 at endogenous sites including the MHb. Habenular expression of the β4 gain-of-function allele T374I also resulted in strong aversion, while transduction with the β4 loss-of function allele R348C failed to induce nicotine aversion. Altogether, these data confirm the critical role of habenular β4 in nicotine consumption, and identify specific SNPs in CHRNB4 that modify nicotine-elicited currents and alter nicotine consumption in mice.
doi:10.3389/fnhum.2014.00012
PMCID: PMC3902282  PMID: 24478678
medial habenula; nicotine consumption; SNP; lentivirus transduction; electrophysiological recordings; smoking dependence
11.  A call for an open, informed study of all aspects of consciousness 
doi:10.3389/fnhum.2014.00017
PMCID: PMC3902298  PMID: 24478682
consciousness; scientific method; parapsychology; PSI; psychical research
12.  Tai chi training reduces self-report of inattention in healthy young adults 
It is important to identify effective non-pharmacological alternatives to stimulant medications that reduce symptoms of attention deficit hyperactivity disorder (ADHD). In this study of healthy young adults, we measured the effects of training in tai chi, which involves mindful attention to the body during movement. Using a non-randomized, controlled, parallel design, students in a 15-week introductory tai chi course (n = 28) and control participants (n = 44) were tested for ADHD indicators and cognitive function at three points over the course of the 15-weeks. The tai chi students’ self-report of attention, but not hyperactivity–impulsivity, improved compared to controls. At baseline, inattention correlated positively with reaction time variability in an affective go/no-go task across all participants, and improvements in attention correlated with reductions in reaction time variability across the tai chi students. Affective bias changed in the tai chi students, as reaction times to positive- and negative-valenced words equalized over time. These results converge to suggest that tai chi training may help improve attention in healthy young adults. Further studies are needed to confirm these results and to evaluate tai chi as therapy for individuals with ADHD.
doi:10.3389/fnhum.2014.00013
PMCID: PMC3902356  PMID: 24478679
Tai chi; attention deficit disorder with hyperactivity; meditation; mindfulness; non-pharmacological intervention; college students; young adults
13.  Linking neuroscientific research on decision making to the educational context of novice students assigned to a multiple-choice scientific task involving common misconceptions about electrical circuits 
Functional magnetic resonance imaging was used to identify the brain-based mechanisms of uncertainty and certainty associated with answers to multiple-choice questions involving common misconceptions about electric circuits. Twenty-two scientifically novice participants (humanities and arts college students) were asked, in an fMRI study, whether or not they thought the light bulbs in images presenting electric circuits were lighted up correctly, and if they were certain or uncertain of their answers. When participants reported that they were unsure of their responses, analyses revealed significant activations in brain areas typically involved in uncertainty (anterior cingulate cortex, anterior insula cortex, and superior/dorsomedial frontal cortex) and in the left middle/superior temporal lobe. Certainty was associated with large bilateral activations in the occipital and parietal regions usually involved in visuospatial processing. Correct-and-certain answers were associated with activations that suggest a stronger mobilization of visual attention resources when compared to incorrect-and-certain answers. These findings provide insights into brain-based mechanisms of uncertainty that are activated when common misconceptions, identified as such by science education research literature, interfere in decision making in a school-like task. We also discuss the implications of these results from an educational perspective.
doi:10.3389/fnhum.2014.00014
PMCID: PMC3902357  PMID: 24478680
uncertainty; certainty; misconception; electricity; fMRI; decision making; education
14.  The perception of peripersonal space in right and left brain damage hemiplegic patients 
Peripersonal space, as opposed to extrapersonal space, is the space that contains reachable objects and in which multisensory and sensorimotor integration is enhanced. Thus, the perception of peripersonal space requires combining information on the spatial properties of the environment with information on the current capacity to act. In support of this, recent studies have provided converging evidences that perceiving objects in peripersonal space activates a neural network overlapping with that subtending voluntary motor action and motor imagery. Other studies have also underlined the dominant role of the right hemisphere (RH) in motor planning and of the left hemisphere (LH) in on-line motor guiding, respectively. In the present study, we investigated the effect of a right or left hemiplegia in the perception of peripersonal space. 16 hemiplegic patients with brain damage to the left (LH) or right (RH) hemisphere and eight matched healthy controls performed a color discrimination, a motor imagery and a reachability judgment task. Analyses of response times and accuracy revealed no variation among the three groups in the color discrimination task, suggesting the absence of any specific perceptual or decisional deficits in the patient groups. In contrast, the patient groups revealed longer response times in the motor imagery task when performed in reference to the hemiplegic arm (RH and LH) or to the healthy arm (RH). Moreover, RH group showed longer response times in the reachability judgment task, but only for stimuli located at the boundary of peripersonal space, which was furthermore significantly reduced in size. Considered together, these results confirm the crucial role of the motor system in motor imagery task and the perception of peripersonal space. They also revealed that RH damage has a more detrimental effect on reachability estimates, suggesting that motor planning processes contribute specifically to the perception of peripersonal space.
doi:10.3389/fnhum.2014.00003
PMCID: PMC3902828  PMID: 24478670
perception and action; spatial vision; peripersonal space; brain damage; right left hemispheres
15.  Hyperschematia after right brain damage: a meaningful entity? 
In recent years we reported three right-brain-damaged patients, who exhibited a left-sided disprortionate expansion of drawings, both by copying and from memory, contralateral to the side of the hemispheric lesion (Neurology, 67: 1801, 2006, Neurocase 14: 369, 2008). We proposed the term “hyperschematia” for such an expansion, with reference to an interpretation in terms of a lateral leftward distortion of the representation of extra-personal space, with a leftward anisometric expansion (relaxation) of the spatial medium. The symptom-complex shown by right-brain-damaged patients with “hyperschematia” includes: (1) a disproportionate leftward expansion of drawings (with possible addition of details), by copy and from memory (also in clay modeling, in one patient); (2) an overestimation of left lateral extent, when a leftward movement is required, associated in some patients with a perceptual underestimation; (3) unawareness of the disorder; (4) no unilateral spatial neglect. In most right-brain-damaged patients, left “hyperschematia” involves extra-personal space. In one patient the deficit was confined to a body part (left half-face: personal “hyperschematia”). The neural underpinnings of the disorder include damage to the fronto-temporo-parietal cortices, and subcortical structures in the right cerebral hemisphere, in the vascular territory of the middle cerebral artery. Here, four novel additional patients are reported. Finally, “hypeschematia” is reconsidered, in its clinical components, the underlying pathological mechanisms, as well as its neural underpinnings.
doi:10.3389/fnhum.2014.00008
PMCID: PMC3904079  PMID: 24478674
hyperschematia; extra-personal space; representation; anisometry
16.  When memory leads the brain to take scenes at face value: face areas are reactivated at test by scenes that were paired with faces at study 
In the first use of the event-related optical signal as a brain imaging tool for the study of long-term memory, we examined relational or associative aspects of memory, widely presumed to involve the interplay among multiple brain regions in representing and reactivating different elements of a given event. Here, we found that a brain region known to be involved in face processing (the posterior superior temporal sulcus) was active not only when viewing faces during the study phase but also when viewing scenes at test that, through prior learning, were associated with specific faces. These findings, demonstrating the activation of stimulus-specific cortical regions in the absence of stimuli of that type, based on learned relations, reveal cortical substrates of the reactivation of relational memories.
doi:10.3389/fnhum.2014.00018
PMCID: PMC3905208  PMID: 24523688
relational memory; cortical reactivation; face processing; scene processing; brain imaging; event-related optical signal (EROS)
17.  Time-course of cortical networks involved in working memory 
Working memory (WM) is one of the most studied cognitive constructs. Although many neuroimaging studies have identified brain networks involved in WM, the time course of these networks remains unclear. In this paper we use dense-array electroencephalography (dEEG) to capture neural signals during performance of a standard WM task, the n-back task, and a blend of principal components analysis and independent components analysis (PCA/ICA) to statistically identify networks of WM and their time courses. Results reveal a visual cortex centric network, that also includes the posterior cingulate cortex, that is active prior to stimulus onset and that appears to reflect anticipatory, attention-related processes. After stimulus onset, the ventromedial prefrontal cortex, lateral prefrontal prefrontal cortex, and temporal poles become associated with the prestimulus network. This second network appears to reflect executive control processes. Following activation of the second network, the cortices of the temporo-parietal junction with the temporal lobe structures seen in the first and second networks re-engage. This third network appears to reflect activity of the ventral attention network involved in control of attentional reorientation. The results point to important temporal features of network dynamics that integrate multiple subsystems of the ventral attention network with the default mode network in the performance of working memory tasks.
doi:10.3389/fnhum.2014.00004
PMCID: PMC3905217  PMID: 24523686
dense-array EEG; working memory; attention; temporal parietal junction; frontal lobe
18.  Cognitive style, cortical stimulation, and the conversion hypothesis 
What does it mean to have a “verbal cognitive style?” We adopt the view that a cognitive style represents a cognitive strategy, and we posit the conversion hypothesis – the notion that individuals with a proclivity for the verbal cognitive style tend to code nonverbal information into the verbal domain. Here we used repetitive transcranial magnetic stimulation (rTMS) to disrupt this hypothesized verbal conversion strategy. Following our previous research implicating left supramarginal gyrus (SMG) in the verbal cognitive style, we used an fMRI paradigm to localize left SMG activity for each subject, then these functional peaks became rTMS targets. Left SMG stimulation impaired performance during a task requiring conversion from pictures to verbal labels. The magnitude of this effect was predicted by individuals’ level of verbal cognitive style, supporting the hypothesized role of left SMG in the verbal labeling strategy, and more generally supporting the conversion hypothesis for cognitive styles.
doi:10.3389/fnhum.2014.00015
PMCID: PMC3905265  PMID: 24523687
cognitive styles; rTMS; conversion hypothesis; verbalizer; fMRI
19.  N1 enhancement in synesthesia during visual and audio–visual perception in semantic cross-modal conflict situations: an ERP study 
Synesthesia entails a special kind of sensory perception, where stimulation in one sensory modality leads to an internally generated perceptual experience of another, not stimulated sensory modality. This phenomenon can be viewed as an abnormal multisensory integration process as here the synesthetic percept is aberrantly fused with the stimulated modality. Indeed, recent synesthesia research has focused on multimodal processing even outside of the specific synesthesia-inducing context and has revealed changed multimodal integration, thus suggesting perceptual alterations at a global level. Here, we focused on audio–visual processing in synesthesia using a semantic classification task in combination with visually or auditory–visually presented animated and in animated objects in an audio–visual congruent and incongruent manner. Fourteen subjects with auditory-visual and/or grapheme-color synesthesia and 14 control subjects participated in the experiment. During presentation of the stimuli, event-related potentials were recorded from 32 electrodes. The analysis of reaction times and error rates revealed no group differences with best performance for audio-visually congruent stimulation indicating the well-known multimodal facilitation effect. We found enhanced amplitude of the N1 component over occipital electrode sites for synesthetes compared to controls. The differences occurred irrespective of the experimental condition and therefore suggest a global influence on early sensory processing in synesthetes.
doi:10.3389/fnhum.2014.00021
PMCID: PMC3906591  PMID: 24523689
synesthesia; multimodal; EEG; N1; integration
20.  Effects of arousal on cognitive control: empirical tests of the conflict-modulated Hebbian-learning hypothesis 
An increasing number of empirical phenomena that were previously interpreted as a result of cognitive control, turn out to reflect (in part) simple associative-learning effects. A prime example is the proportion congruency effect, the finding that interference effects (such as the Stroop effect) decrease as the proportion of incongruent stimuli increases. While this was previously regarded as strong evidence for a global conflict monitoring-cognitive control loop, recent evidence has shown that the proportion congruency effect is largely item-specific and hence must be due to associative learning. The goal of our research was to test a recent hypothesis about the mechanism underlying such associative-learning effects, the conflict-modulated Hebbian-learning hypothesis, which proposes that the effect of conflict on associative learning is mediated by phasic arousal responses. In Experiment 1, we examined in detail the relationship between the item-specific proportion congruency effect and an autonomic measure of phasic arousal: task-evoked pupillary responses. In Experiment 2, we used a task-irrelevant phasic arousal manipulation and examined the effect on item-specific learning of incongruent stimulus–response associations. The results provide little evidence for the conflict-modulated Hebbian-learning hypothesis, which requires additional empirical support to remain tenable.
doi:10.3389/fnhum.2014.00023
PMCID: PMC3906597  PMID: 24523690
conflict; Stroop effect; interference; arousal; pupillometry; accessory stimulus
21.  Chronic traumatic encephalopathy and other neurodegenerative proteinopathies 
“Chronic traumatic encephalopathy” (CTE) is described as a slowly progressive neurodegenerative disease believed to result from multiple concussions. Traditionally, concussions were considered benign events and although most people recover fully, about 10% develop a post-concussive syndrome with persisting neurological, cognitive and neuropsychiatric symptoms. CTE was once thought to be unique to boxers, but it has now been observed in many different athletes having suffered multiple concussions as well as in military personal after repeated blast injuries. Much remains unknown about the development of CTE but its pathological substrate is usually tau, similar to that seen in Alzheimer's disease (AD) and frontotemporal lobar degeneration (FTLD). The aim of this “perspective” is to compare and contrast clinical and pathological CTE with the other neurodegenerative proteinopathies and highlight that there is an urgent need for understanding the relationship between concussion and the development of CTE as it may provide a window into the development of a proteinopathy and thus new avenues for treatment.
doi:10.3389/fnhum.2014.00030
PMCID: PMC3907709  PMID: 24550810
concussions; chronic traumatic encephalopathy; neurodegenerative disease; Alzheimer's disease; frontotemporal lobar degeneration; tau
22.  Sensory-to-motor integration during auditory repetition: a combined fMRI and lesion study 
The aim of this paper was to investigate the neurological underpinnings of auditory-to-motor translation during auditory repetition of unfamiliar pseudowords. We tested two different hypotheses. First we used functional magnetic resonance imaging in 25 healthy subjects to determine whether a functionally defined area in the left temporo-parietal junction (TPJ), referred to as Sylvian-parietal-temporal region (Spt), reflected the demands on auditory-to-motor integration during the repetition of pseudowords relative to a semantically mediated nonverbal sound-naming task. The experiment also allowed us to test alternative accounts of Spt function, namely that Spt is involved in subvocal articulation or auditory processing that can be driven either bottom-up or top-down. The results did not provide convincing evidence that activation increased in either Spt or any other cortical area when non-semantic auditory inputs were being translated into motor outputs. Instead, the results were most consistent with Spt responding to bottom up or top down auditory processing, independent of the demands on auditory-to-motor integration. Second, we investigated the lesion sites in eight patients who had selective difficulties repeating heard words but with preserved word comprehension, picture naming and verbal fluency (i.e., conduction aphasia). All eight patients had white-matter tract damage in the vicinity of the arcuate fasciculus and only one of the eight patients had additional damage to the Spt region, defined functionally in our fMRI data. Our results are therefore most consistent with the neurological tradition that emphasizes the importance of the arcuate fasciculus in the non-semantic integration of auditory and motor speech processing.
doi:10.3389/fnhum.2014.00024
PMCID: PMC3908611  PMID: 24550807
fMRI; lesions; language; speech; aphasia
23.  Increased metabolic activity in the septum and habenula during stress is linked to subsequent expression of learned helplessness behavior 
Uncontrollable stress can have a profound effect on an organism's ability to respond effectively to future stressful situations. Behavior subsequent to uncontrollable stress can vary greatly between individuals, falling on a spectrum between healthy resilience and maladaptive learned helplessness. It is unclear whether dysfunctional brain activity during uncontrollable stress is associated with vulnerability to learned helplessness; therefore, we measured metabolic activity during uncontrollable stress that correlated with ensuing inability to escape future stressors. We took advantage of small animal positron emission tomography (PET) and 2-deoxy-2[18F]fluoro-D-glucose (18FDG) to probe in vivo metabolic activity in wild type Sprague Dawley rats during uncontrollable, inescapable, unpredictable foot-shock stress, and subsequently tested the animals response to controllable, escapable, predictable foot-shock stress. When we correlated metabolic activity during the uncontrollable stress with consequent behavioral outcomes, we found that the degree to which animals failed to escape the foot-shock correlated with increased metabolic activity in the lateral septum and habenula. When used a seed region, metabolic activity in the habenula correlated with activity in the lateral septum, hypothalamus, medial thalamus, mammillary nuclei, ventral tegmental area, central gray, interpeduncular nuclei, periaqueductal gray, dorsal raphe, and rostromedial tegmental nucleus, caudal linear raphe, and subiculum transition area. Furthermore, the lateral septum correlated with metabolic activity in the preoptic area, medial thalamus, habenula, interpeduncular nuclei, periaqueductal gray, dorsal raphe, and caudal linear raphe. Together, our data suggest a group of brain regions involved in sensitivity to uncontrollable stress involving the lateral septum and habenula.
doi:10.3389/fnhum.2014.00029
PMCID: PMC3909949  PMID: 24550809
metabolism; whole-brain; imaging; FDG; PET; circuitry; depression; DBS
24.  The entropic brain: a theory of conscious states informed by neuroimaging research with psychedelic drugs 
Entropy is a dimensionless quantity that is used for measuring uncertainty about the state of a system but it can also imply physical qualities, where high entropy is synonymous with high disorder. Entropy is applied here in the context of states of consciousness and their associated neurodynamics, with a particular focus on the psychedelic state. The psychedelic state is considered an exemplar of a primitive or primary state of consciousness that preceded the development of modern, adult, human, normal waking consciousness. Based on neuroimaging data with psilocybin, a classic psychedelic drug, it is argued that the defining feature of “primary states” is elevated entropy in certain aspects of brain function, such as the repertoire of functional connectivity motifs that form and fragment across time. Indeed, since there is a greater repertoire of connectivity motifs in the psychedelic state than in normal waking consciousness, this implies that primary states may exhibit “criticality,” i.e., the property of being poised at a “critical” point in a transition zone between order and disorder where certain phenomena such as power-law scaling appear. Moreover, if primary states are critical, then this suggests that entropy is suppressed in normal waking consciousness, meaning that the brain operates just below criticality. It is argued that this entropy suppression furnishes normal waking consciousness with a constrained quality and associated metacognitive functions, including reality-testing and self-awareness. It is also proposed that entry into primary states depends on a collapse of the normally highly organized activity within the default-mode network (DMN) and a decoupling between the DMN and the medial temporal lobes (which are normally significantly coupled). These hypotheses can be tested by examining brain activity and associated cognition in other candidate primary states such as rapid eye movement (REM) sleep and early psychosis and comparing these with non-primary states such as normal waking consciousness and the anaesthetized state.
doi:10.3389/fnhum.2014.00020
PMCID: PMC3909994  PMID: 24550805
serotonin; default mode network; criticality; entropy; 5-HT2A receptor; metastability; consciousness; REM sleep
25.  Broad-band Gaussian noise is most effective in improving motor performance and is most pleasant 
Modern attempts to improve human performance focus on stochastic resonance (SR). SR is a phenomenon in non-linear systems characterized by a response increase of the system induced by a particular level of input noise. Recently, we reported that an optimum level of 0–15 Hz Gaussian noise applied to the human index finger improved static isometric force compensation. A possible explanation was a better sensorimotor integration caused by increase in sensitivity of peripheral receptors and/or of internal SR. The present study in 10 subjects compares SR effects in the performance of the same motor task and on pleasantness, by applying three Gaussian noises chosen on the sensitivity of the fingertip receptors (0–15 Hz mostly for Merkel receptors, 250–300 Hz for Pacini corpuscles and 0–300 Hz for all). We document that only the 0–300 Hz noise induced SR effect during the transitory phase of the task. In contrast, the motor performance was improved during the stationary phase for all three noise frequency bandwidths. This improvement was stronger for 0–300 Hz and 250–300 Hz than for 0–15 Hz noise. Further, we found higher degree of pleasantness for 0–300 Hz and 250–300 Hz noise bandwidths than for 0–15 Hz. Thus, we show that the most appropriate Gaussian noise that could be used in haptic gloves is the 0–300 Hz, as it improved motor performance during both stationary and transitory phases. In addition, this noise had the highest degree of pleasantness and thus reveals that the glabrous skin can also forward pleasant sensations.
doi:10.3389/fnhum.2014.00022
PMCID: PMC3910318  PMID: 24550806
noise; frequency; stochastic resonance; finger; motor; force; humans

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