The Spatial Reach of Neuronal Coherence and Spike-Field Coupling across the Human Neocortex.

Neuronal coherence is thought to be a fundamental mechanism of communication in the brain, where synchronized field potentials coordinate synaptic and spiking events to support plasticity and learning. Although the spread of field potentials has garnered great interest, little is known about the spatial reach of phase synchronization, or neuronal coherence. Functional connectivity between different brain regions is known to occur across long distances, but the locality of synchronization across the neocortex is understudied. Here we used simultaneous recordings from electrocorticography (ECoG) grids and high-density microelectrode arrays to estimate the spatial reach of neuronal coherence and spike-field coherence (SFC) across frontal, temporal, and occipital cortices during cognitive tasks in humans. We observed the strongest coherence within a 2-3 cm distance from the microelectrode arrays, potentially defining an effective range for local communication. This range was relatively consistent across brain regions, spectral frequencies, and cognitive tasks. The magnitude of coherence showed power law decay with increasing distance from the microelectrode arrays, where the highest coherence occurred between ECoG contacts, followed by coherence between ECoG and deep cortical local field potential (LFP), and then SFC (i.e., ECoG > LFP > SFC). The spectral frequency of coherence also affected its magnitude. Alpha coherence (8-14 Hz) was generally higher than other frequencies for signals nearest the microelectrode arrays, whereas delta coherence (1-3 Hz) was higher for signals that were farther away. Action potentials in all brain regions were most coherent with the phase of alpha oscillations, which suggests that alpha waves could play a larger, more spatially local role in spike timing than other frequencies. These findings provide a deeper understanding of the spatial and spectral dynamics of neuronal synchronization, further advancing knowledge about how activity propagates across the human brain.SIGNIFICANCE STATEMENT Coherence is theorized to facilitate information transfer across cerebral space by providing a convenient electrophysiological mechanism to modulate membrane potentials in spatiotemporally complex patterns. Our work uses a multiscale approach to evaluate the spatial reach of phase coherence and spike-field coherence during cognitive tasks in humans. Locally, coherence can reach up to 3 cm around a given area of neocortex. The spectral properties of coherence revealed that alpha phase-field and spike-field coherence were higher within ranges <2 cm, whereas lower-frequency delta coherence was higher for contacts farther away. Spatiotemporally shared information (i.e., coherence) across neocortex seems to reach farther than field potentials alone.

Pretrial predictors of conflict response efficacy in the human prefrontal cortex.

The ability to perform motor actions depends, in part, on the brain's initial state. We hypothesized that initial state dependence is a more general principle and applies to cognitive control. To test this idea, we examined human single units recorded from the dorsolateral prefrontal (dlPFC) cortex and dorsal anterior cingulate cortex (dACC) during a task that interleaves motor and perceptual conflict trials, the multisource interference task (MSIT). In both brain regions, variability in pre-trial firing rates predicted subsequent reaction time (RT) on conflict trials. In dlPFC, ensemble firing rate patterns suggested the existence of domain-specific initial states, while in dACC, firing patterns were more consistent with a domain-general initial state. The deployment of shared and independent factors that we observe for conflict resolution may allow for flexible and fast responses mediated by cognitive initial states. These results also support hypotheses that place dACC hierarchically earlier than dlPFC in proactive control.

A generalized magnitude system for space, time, and quantity? A cautionary note.

We investigated the claim that larger stimuli are perceived to last longer (Xuan, Zhang, He, & Chen, 2007). This claim, along with other similar claims of interactions between magnitude representations, is frequently used to support the generalized magnitude system hypothesis-the suggestion that the brain represents information about different magnitudes (e.g., time, space, and quantity) via a common mechanism. It is not clear, however, whether the size of a stimulus genuinely affects the perceived duration of the stimulus or simply biases decisions about duration. This was addressed using duration "equality judgments," which have been proposed to measure perceived duration unconfounded by decisional bias-in contrast to "comparative judgments," which are generally considered bias-prone. Using equality judgments, we failed to find support for the claim that larger stimuli are perceived to last longer, despite replicating the original effect reported by Xuan et al. (2007) using comparative judgments. Instead, unexpectedly, larger stimuli were judged-though not necessarily perceived-as shorter in duration. This result casts doubt on the conclusions of a significant body of behavioral interference studies using comparative judgments, which support a generalized magnitude system. We also identify a hitherto unrecognized potential source of decisional bias associated with equality judgments.

Near, yet so far: the effect of pictorial cues on spatial attention.

Distinct cognitive and neural mechanisms underlie perception and action in near (within-reach) and far (outside-reach) space. Objects in far space can be brought into the brain's near-space through tool-use. We determined whether a near object can be pushed into far space by changing the pictorial context in which it occurs. Participants (n = 372) made relative length judgements for lines presented in near space, but superimposed over photographs of near and far objects. The left segment of the line was overestimated in the baseline and near-context conditions whereas the right was overestimated in the far-context. The change from leftward to rightward overestimation is the same when lines are physically shifted from near to far space. Because participants did not have to do anything in relation to the photograph, the results suggest that simply viewing images with a near/far context can cause a shift of attention along the distal/proximal axis, which may reflect differential activation of the ventral/dorsal visual streams.

Widespread temporal coding of cognitive control in the human prefrontal cortex.

When making decisions we often face the need to adjudicate between conflicting strategies or courses of action. Our ability to understand the neuronal processes underlying conflict processing is limited on the one hand by the spatiotemporal resolution of functional MRI and, on the other hand, by imperfect cross-species homologies in animal model systems. Here we examine the responses of single neurons and local field potentials in human neurosurgical patients in two prefrontal regions critical to controlled decision-making, the dorsal anterior cingulate cortex (dACC) and dorsolateral prefrontal cortex (dlPFC). While we observe typical modest conflict-related firing rate effects, we find a widespread effect of conflict on spike-phase coupling in the dACC and on driving spike-field coherence in the dlPFC. These results support the hypothesis that a cross-areal rhythmic neuronal coordination is intrinsic to cognitive control in response to conflict, and provide new evidence to support the hypothesis that conflict processing involves modulation of the dlPFC by the dACC.

A new means of measuring index/ring finger (2D:4D) ratio and its association with gender and hand preference.

Relative finger length can predict a person's gender or their hand preference. We measured finger length using a new "tubes" test, which required participants to slide a clear plastic tube over their fingers and read the length from an attached mm scale. Data collected from 600 students demonstrated that the right fingers are longer than the left for dextrals, but not for non-dextrals. Examination of the relative length of the index and ring fingers revealed a clear gender effect. There was also an effect of hand preference on index/ring finger ratio whereby non-dextrals showed a more masculine pattern compared to dextrals. For non-dextrals, prenatal exposure to high testosterone levels may have caused both a shift away from dextrality and a more masculine pattern of finger ratio. In the second experiment, finger length was measured by the tubes test and by photocopying the hands in 124 undergraduates. The tubes test yielded a longer estimate of ring finger length compared to the photocopy method. Despite this, there was a strong correlation between the tests and both showed an association with gender. Finally, test-retest scores for 45 participants showed a high level of reliability for absolute and relative finger measures. We conclude that the tubes technique provides an effective and easy-to-use means of measuring finger length, which can be administered in a classroom setting.

Non-symbolic magnitudes are represented spatially: Evidence from a non-symbolic SNARC task.

A core proposition in numerical cognition is numbers are represented spatially. Evidence for this proposition comes from the "spatial numerical association of response codes" effect (SNARC) in which faster responses are made by the left/right hand judging whether one of a pair of Arabic digits is smaller/larger than the other. Less is known if a similar SNARC effect exists for non-symbolic magnitudes; and research that has been conducted used stimuli which could be translated into symbolic terms. To overcome this limitation, we employed a referent-to-target judgment paradigm in which a referent dot array (n = 30 dots) was follow by a second array of dots (e.g., n = 45 or 15 dots)-participants judged if the second array contained fewer or more dots than the referent array. Dot arrays with fewer dots were judged more quickly with the left hand compared to the right hand (i.e., a SNARC effect). Not all participants demonstrated a SNARC effect, however. Neither visuospatial working memory nor math ability was associated with the presence/absence of a non-symbolic SNARC effect. Implications of the non-symbolic SNARC effect for accounts of numerical cognition are discussed.

Neuronal activity in human anterior cingulate cortex modulates with internal cognitive state during multi-source interference task.

The dorsal anterior cingulate cortex (dACC) is thought to be essential for normal adaptation of one's behavior to difficult decisions, errors, and reinforcement. Here we examine single neurons from the human dACC in the context of a statistical model, including a cognitive state that varies with changes in cognitive interference induced by a Stroop-like task. We then include this cognitive state in point process models of single unit activity and subject reaction time. These results suggest that consideration of a latent cognitive state can explain additional variance in neural and behavioral dynamics.

Asymmetric frontal cortical activity predicts effort expenditure for reward.

An extensive literature shows that greater left, relative to right, frontal cortical activity (LFA) is involved in approach-motivated affective states and reflects stable individual differences in approach motivation. However, relatively few studies have linked LFA to behavioral indices of approach motivation. In this study, we examine the relation between LFA and effort expenditure for reward, a behavioral index of approach motivation. LFA was calculated for 51 right-handed participants (55% female) using power spectral analysis of electroencephalogram recorded at rest. Participants also completed the effort expenditure for rewards task (EEfRT), which presents a series of trials requiring a choice between a low-reward low-effort task and a high-reward high-effort task. We found that individuals with greater resting LFA were more willing to expend greater effort in the pursuit of larger rewards, particularly when reward delivery was less likely. Our findings offer a more nuanced understanding of the motivational significance of LFA, in terms of processes that mitigate the effort- and uncertainty-related costs of pursuing rewarding goals.

The highs and lows of the interaction between word meaning and space.

We examined whether the processing of words associated with distinct spatial locations automatically biases behavior toward these locations in space. In four experiments (Ns = 30, 34, 32, 32), participants were shown stimuli denoting objects typically associated with the upper and lower regions of visual space. In Experiment 1, words were categorized as man-made or natural by pressing one of two vertically arranged keys. Reaction times were faster for trials in which response locations were congruent with the stimulus-associated locations. Experiment 2 replicated the stimulus-response congruency effect when the stimuli were presented in a pictorial format. Stimuli-space interactions therefore seem to be driven by an automatic activation of the spatial attributes associated with the stimuli, irrespective of input format. In Experiments 3 and 4, a target detection task involving only one response button was employed to examine whether the effects observed in the first two experiments were due to attentional shifts, independent of response selection processes. In both experiments, the previously observed congruence effect between words and space either diminished or vanished completely. Consequently, the results of the four experiments in the current study point to a dominant role of response-selection processes in the genesis of space-object word interactions.

The relationship between vertical stimulation and horizontal attentional asymmetries.

The original aim was to examine the effect of perceived distance, induced by the Ponzo illusion, on left/right asymmetries for line bisection. In Experiment 1, university students (n = 29) made left/right bisection judgements for lines presented in the lower or upper half of the screen against backgrounds of the Ponzo stimuli, or a baseline. While the Ponzo illusion had relatively little effect on line bisection, elevation in the baseline condition had a strong effect, whereby the leftward bias was increased for upper lines. Experiment 2 (n = 17) eliminated the effect of elevation by presenting the line in the middle and moving the Ponzo stimuli relative to the line. Despite this change, the leftward bias was still stronger in the upper condition in the baseline condition. The final experiment (n = 17) investigated whether upper/lower visual stimulation, which was irrelevant to the task, affected asymmetries for line bisection. The results revealed that a rectangle presented in the upper half of the screen increased the leftward line bisection bias relative to a baseline and lower stimulation condition. These results corroborate neuroimaging research, showing increased right parietal activation associated with shifts of attention into the upper hemispace. This increased right parietal activation may increase the leftward attentional bias-resulting in a stronger leftward bias for line bisection.

Somatosensory prior entry assessed with temporal order judgments and simultaneity judgments.

Attention is central to perception, yet a clear understanding of how attention influences the latency of perception has proven surprisingly elusive. Recent research has indicated that spatially attended stimuli are perceived earlier than unattended stimuli across a range of sensory modalities-an effect termed prior entry. However, the method commonly used to measure this, the temporal order judgment (TOJ) task, has been criticized as susceptible to response bias, despite deliberate attempts to minimize such bias. A preferred alternative is the simultaneity judgment (SJ) task. We tested the prior-entry hypothesis for somatosensory stimuli using both a TOJ task (replicating an earlier experiment) and an SJ task. Prior-entry effects were found for both, though the effect was reduced in the SJ task. Additional experiments (TOJ and SJ) using visual cues established that the earlier perception of cued tactile targets does not result from intramodal sensory interactions between tactile cues and targets.

The importance of response type to the relationship between temporal order and numerical magnitude.

Time, number, and space may be represented in the brain by a common set of cognitive/neural mechanisms. In support of this conjecture, Schwarz and Eiselt (Journal of Experimental Psychology: Human Perception and Performance, 35, 989-1004, 2009) found that numerically smaller digits were perceived to occur earlier than larger digits, and they concluded that this difference reflected faster processing of smaller numbers. This difference, however, could have been related to a response bias, whereby participants map responses of "which first" onto the "first" number along the mental number line. In Experiment 1, participants made temporal order judgements between digits presented to the left or to the right. The point of subjective simultaneity was shifted so that the 9 had to be presented before the 2 in order for simultaneity to be perceived. This difference could reflect either faster processing of the 2 or a response bias. Experiments 2a and 2b eliminated response biases by using simultaneity judgements, which have no logical stimulus mapping. Both of the latter experiments established that the 2 was not processed faster than the 9. Although the present results relate specifically to numerical magnitude and temporal order associations, they also have broader implications. Other studies have reported associations between dimensions such as size, duration, and number and have attributed these to parietal mechanisms. Such associations, however, may also be an artefact of response biases.

Somatosensory prior entry.

The perceived timing of sensory events does not necessarily match the actual timing. In the present study, we investigated the effect of location of attention on the perceived timing of somatosensory stimuli. Participants judged the temporal order of two taps, delivered one to each hand, with taps presented at different elevations (upper and lower) on the opposing hands. The task was to discriminate the elevation of the tap presented first (or second). This vertical discrimination task was orthogonal to the horizontal attentional cuing manipulation, removing the response bias confound that has undermined earlier studies investigating the impact of attention on the perceived timing of sensory stimuli. We manipulated spatial attention either (1) exogenously (Experiment 1) in 13 participants, using brief taps to either the left or right hand, or (2) endogenously (Experiment 2) in 22 participants, using centrally presented symbolic cues. The results supported the hypothesis that attended stimuli are perceived more rapidly than unattended stimuli. This effect was larger when attention was exogenously manipulated. Previous research has demonstrated a similar effect for visual stimuli. The present study, which extends this result to somatosensory perception, indicates that the phenomenon may represent a more global feature of the perceptual system, which is possibly mediated by a common modality-independent mechanism.