Differential contribution of sensorimotor cortex and subthalamic nucleus to unimanual and bimanual hand movements.

Why does unilateral deep brain stimulation improve motor function bilaterally? To address this clinical observation, we collected parallel neural recordings from sensorimotor cortex (SMC) and the subthalamic nucleus (STN) during repetitive ipsilateral, contralateral, and bilateral hand movements in patients with Parkinson's disease. We used a cross-validated electrode-wise encoding model to map electromyography data to the neural signals. Electrodes in the STN encoded movement at a comparable level for both hands, whereas SMC electrodes displayed a strong contralateral bias. To examine representational overlap across the two hands, we trained the model with data from one condition (contralateral hand) and used the trained weights to predict neural activity for movements produced with the other hand (ipsilateral hand). Overall, between-hand generalization was poor, and this limitation was evident in both regions. A similar method was used to probe representational overlap across different task contexts (unimanual vs. bimanual). Task context was more important for the STN compared to the SMC indicating that neural activity in the STN showed greater divergence between the unimanual and bimanual conditions. These results indicate that SMC activity is strongly lateralized and relatively context-free, whereas the STN integrates contextual information with the ongoing behavior.

Left hemisphere dominance for bilateral kinematic encoding in the human brain.

Neurophysiological studies in humans and nonhuman primates have revealed movement representations in both the contralateral and ipsilateral hemispheres. Inspired by clinical observations, we ask if this bilateral representation differs for the left and right hemispheres. Electrocorticography was recorded in human participants during an instructed-delay reaching task, with movements produced with either the contralateral or ipsilateral arm. Using a cross-validated kinematic encoding model, we found stronger bilateral encoding in the left hemisphere, an effect that was present during preparation and was amplified during execution. Consistent with this asymmetry, we also observed better across-arm generalization in the left hemisphere, indicating similar neural representations for right and left arm movements. Notably, these left hemisphere electrodes were centered over premotor and parietal regions. The more extensive bilateral encoding in the left hemisphere adds a new perspective to the pervasive neuropsychological finding that the left hemisphere plays a dominant role in praxis.

The brain is split into two hemispheres, each playing the leading role in coordinating movement for the opposite side of the body: lesions on the left hemisphere therefore often result in difficulties moving the right arm or leg, and vice versa. In fact, very few anatomical connections exist between a given hemisphere and the body parts on the same (or 'ipsilateral') side. Yet, movements produced with only one limb still engage both sides of the brain, with the hemisphere which does not control the action production, still encoding the direction and speed of the movement. Previous evidence also indicate that the two hemispheres may not have equal roles when coordinating ipsilateral movements. Merrick et al. aimed to shed light on these processes; to do so, they measured electrical activity from the surface of the brain of six patients as they moved their arms to reach a screen. The results revealed that, while the right hemisphere only encoded information about the opposite arm, the left hemisphere contained information about both arms. Finer analyses showed that, for both hemispheres, moving the opposite arm was strongly associated with activity in the primary motor cortex, a region which helps to execute movements. However, in the left hemisphere, movements from the ipsilateral arm were related to activity in brain areas involved in planning and integrating different types of sensory information. These findings contribute to a better understanding of how the motor system works, which could ultimately help with the development of brain-machine interfaces for patients who need a neuroprosthetic limb.

Hybrid dedicated and distributed coding in PMd/M1 provides separation and interaction of bilateral arm signals.

Pronounced activity is observed in both hemispheres of the motor cortex during preparation and execution of unimanual movements. The organizational principles of bi-hemispheric signals and the functions they serve throughout motor planning remain unclear. Using an instructed-delay reaching task in monkeys, we identified two components in population responses spanning PMd and M1. A "dedicated" component, which segregated activity at the level of individual units, emerged in PMd during preparation. It was most prominent following movement when M1 became strongly engaged, and principally involved the contralateral hemisphere. In contrast to recent reports, these dedicated signals solely accounted for divergence of arm-specific neural subspaces. The other "distributed" component mixed signals for each arm within units, and the subspace containing it did not discriminate between arms at any stage. The statistics of the population response suggest two functional aspects of the cortical network: one that spans both hemispheres for supporting preparatory and ongoing processes, and another that is predominantly housed in the contralateral hemisphere and specifies unilateral output.

Involuntary Entry Into Consciousness From the Activation of Sets: Object Counting and Color Naming.

High-level cognitions can enter consciousness through the activation of certain action sets and the presentation of external stimuli ("set-based entry," for short). Set-based entry arises in a manner that is involuntary and systematic. In the Reflexive Imagery Task, for example, subjects are presented with visual objects and instructed to not think of the names of the objects. Involuntary subvocalizations arise on roughly 80% of the trials. We examined whether or not set-based entry can also occur in the case of involuntary counting. Subjects in Experiment 1A were instructed to not count the number of objects presented in an array. Involuntary counting arose on a high proportion of the trials (a mean proportion of ∼0.90) for stimulus arrays having 2-5 objects, and such counting arose less frequently across trials when the array consisted of 6-10 objects (a mean proportion of ∼0.21). The data from Experiment 1B revealed that, when people choose to perform Set X, they also experience thoughts about an unselected Set (Set Y). Subjects were trained on one set (e.g., to "color name") and then, when presented with stimuli, were given the choice to perform the trained set or a novel set. Consistent with theories proposing that the conscious contents represent several potential action plans, subjects were equally likely to experience set-related imagery or set-unrelated imagery. Our findings regarding set-based entry are relevant to many subfields of psychology and neuroscience (e.g., the study of high-level mental processes, attention, imagery, and action control).

Ironic effects as reflexive responses: Evidence from word frequency effects on involuntary subvocalizations.

In ironic processing, one is more likely to think about something (e.g., white bears) when instructed to not think about that thing. To further investigate this phenomenon involving cognitive control, in the Reflexive Imagery Task (RIT), participants are instructed to not subvocalize the names of visual objects. On the majority of the trials, participants fail to suppress such subvocalizations. This finding supports theorizing that conscious thoughts can be triggered by external stimuli in a manner that is nontrivial, involuntary, and, importantly, reflex-like. These conclusions challenge intuitions that consciousness is unpredictable, whimsical, and somewhat insulated from external control. Perhaps these thoughts arise, not in a reflex-like manner, but from experimental demand or other high-level, strategic processes. This prevalent criticism would be inconsistent with the observation that the RIT effect is influenced by a stimulus parameter such as word frequency. Regarding demand characteristics, such an artifact would require participants to have a theory regarding how word frequency should influence responses. We introduce evidence that stimuli associated with high frequency names are more likely to yield involuntary subvocalizations than stimuli associated with low frequency names. These theoretically-relevant data suggest that ironic effects in paradigms such as the RIT resemble reflex-like processes.

External control of the stream of consciousness: Stimulus-based effects on involuntary thought sequences.

The stream of consciousness often appears whimsical and free from external control. Recent advances, however, reveal that the stream is more susceptible to external influence than previously assumed. Thoughts can be triggered by external stimuli in a manner that is involuntary, systematic, and nontrivial. Based on these advances, our experimental manipulation systematically triggered a sequence of, not one, but two involuntary thoughts. Participants were instructed to (a) not subvocalize the name of visual objects and (b) not count the number of letters comprising object names. On a substantial proportion of trials, participants experienced both kinds of involuntary thoughts. Each thought arose from distinct, high-level processes (naming versus counting). This is the first demonstration of the induction of two involuntary thoughts into the stream of consciousness. Stimulus word length influenced dependent measures systematically. Our findings are relevant to many fields associated with the study of consciousness, including attention, imagery, and action control.

Subjective aspects of working memory performance: memoranda-related imagery.

Although it is well accepted that working memory (WM) is intimately related to consciousness, little research has illuminated the liaison between the two phenomena. To investigate this under-explored nexus, we used an imagery monitoring task to investigate the subjective aspects of WM performance. Specifically, in two experiments, we examined the effects on consciousness of (a) holding in mind information having a low versus high memory load, and (b) holding memoranda in mind during the presentation of distractors (e.g., visual stimuli associated with a response incompatible with that of the memoranda). Higher rates of rehearsal (conscious imagery) occurred in the high load and distractor conditions than in comparable control conditions. Examination of the temporal properties of the rehearsal-based imagery revealed that, across subjects, imagery events occurred evenly throughout the delay. We hope that future variants of this new imagery monitoring task will reveal additional insights about WM, consciousness, and action control.

The olfactory system as the gateway to the neural correlates of consciousness.

How consciousness is generated by the nervous system remains one of the greatest mysteries in science. Investigators from diverse fields have begun to unravel this puzzle by contrasting conscious and unconscious processes. In this way, it has been revealed that the two kinds of processes differ in terms of the underlying neural events and associated cognitive mechanisms. We propose that, for several reasons, the olfactory system provides a unique portal through which to examine this contrast. For this purpose, the olfactory system is beneficial in terms of its (a) neuroanatomical aspects, (b) phenomenological and cognitive/mechanistic properties, and (c) neurodynamic (e.g., brain oscillations) properties. In this review, we discuss how each of these properties and aspects of the olfactory system can illuminate the contrast between conscious and unconscious processing in the brain. We conclude by delineating the most fruitful avenues of research and by entertaining hypotheses that, in order for an olfactory content to be conscious, that content must participate in a network that is large-scale, both in terms of the neural systems involved and the scope of information integration.