Overt Movement Research Articles

Sequential activation of premotor, primary somatosensory and primary motor areas in humans during cued finger movements

ObjectiveHuman voluntary movements are a final product of complex interactions between multiple sensory, cognitive and motor areas of central nervous system. The objective was to investigate temporal sequence of activation of premotor (PM), primary motor (M1) and somatosensory (S1) areas during cued finger movements. MethodsElectrocorticography (ECoG) was used to measure activation timing in human PM, S1, and M1 neurons in preparation for finger movements in 5 subjects with subdural grids for seizure localization. Cortical activation was determined by the onset of high gamma (HG) oscillation (70–150Hz). The three cortical regions were mapped anatomically using a common brain atlas and confirmed independently with direct electrical cortical stimulation, somatosensory evoked potentials and detection of HG response to tactile stimulation. Subjects were given visual cues to flex each finger or pinch the thumb and index finger. Movements were captured with a dataglove and time-locked with ECoG. A windowed covariance metric was used to identify the rising slope of HG power between two electrodes and compute time lag. Statistical constraints were applied to the time estimates to combat the noise. Rank sum testing was used to verify the sequential activation of cortical regions across 5 subjects. ResultsIn all 5 subjects, HG activation in PM preceded S1 by an average of 53±13ms (P=0.03), PM preceded M1 by 180±40ms (P=0.001) and S1 activation preceded M1 by 136±40ms (P=0.04). ConclusionsSequential HG activation of PM, S1 and M1 regions in preparation for movements is reported. Activity in S1 prior to any overt body movements supports the notion that these neurons may encode sensory information in anticipation of movements, i.e., an efference copy. Our analysis suggests that S1 modulation likely originates from PM. SignificanceFirst electrophysiological evidence of efference copy in humans.

Dual Textual Dynamics and Dual Readerly Dynamics: Double Narrative Movements in Mansfield's “Psychology”

In many fictional narratives, behind the plot development, there exists a powerful dynamic that runs, at a deeper level, throughout the text. This hidden dynamic paralleling the plot development is what I designate as textual (see Shen, Covert, Style). It involves different instabilities on the story level and different tensions on the discourse level (see Phelan, Narrative Progression, Narrative), thereby conveying contrasting or even opposing thematic significance, character images, and aesthetic values to those in the plot development, and arousing or having the potential to arouse contrasting or even opposing response from readers. In my previous work, I have distinguished covert from other types of meaning as investigated by various critical approaches since the nineteenth century, including the meanings revealed by New Critics and contemporary literary/narrative critics (see Shen, Style 7-12, Covert 148-52). Other concerns with meaning focus on deep or deeper levels of meaning of the plot development, especially ways in which latent meanings of the plot subvert or oppose its manifest ones. In contrast, my concern with progression is a concern with a hidden narrative movement paralleling the plot development. But previously I did not pay sufficient attention to the interaction and joint fonctioning of the two parallel narrative movements, especially in cases where the meanings of the progression subvert those of the plot development. This is the first time I try to offer theoretical models of the dual textual dynamics and the corresponding dual readerly dynamics, both to show and to call for attention to the joint functioning of the two parallel narrative movements. This may enable us to do better justice to the instability, tension, complexity, and otherwise self-contradictory nature of literary narratives. The present effort to theorize about the dual textual and readerly dynamics is based on the most telling case of Katherine Mansfield's Psychology. In this narrative, the progression takes on an opposite event structure, contrastive focalization, and different degrees of narratorial reliability. More specifically, while in the plot development the event structure is merely revelatory and things stay pretty much the same (Chatman 48), in the progression the event structure displays a progress towards a resolution; while in the plot the focalization keeps shifting, in the progression the focalization is quite fixed. Moreover, what appears to be reliable reporting by the narrator in the plot development frequently turns out to be merely character's illusion in the progression. This progression in Mansfield's Psychology very much relies on the ambiguity created by the use of free indirect discourse and point of view or focalization. These devices, among other modernist techniques, have attracted much critical attention in the investigation of Mansfield's fiction, including Psychology, since she is well known for her masterful use of such techniques. But the progression in Psychology has remained overlooked in existing literary criticism because ever since Aristotle, critical attention has focused on one narrative movement, the plot development. Indeed, the progression is a type of meaning that readers miss not because it's hidden but largely because their interpretive equipment won't allow them to see what is right there in plain sight (Abbott 560). As we will see below, unless we revise our interpretive framework and extend attention to another narrative movement behind the plot development, critical sophistication, acumen, and carefulness may not help much in discovering the double narrative movements as such. Significantly, the dual textual dynamics purposefully created by the author invites dual readerly dynamics. In more specific terms, the progression and the interaction between the and overt narrative movements invite the authorial audience to change the perception and judgment of various textual details in the overt plot, increasingly altering and complicating the understanding of the rhetorical purposes of the author and the thematic import, character relationship, and aesthetic effects of the text. …

Pseudo Scope Marking Constructions in Blackfoot

This paper discusses and analyzes a kind of scope marking construction in Blackfoot (Algonquian) and adds to the diversity of such constructions found in languages around the world. This construction has many properties in common with other scope marking constructions but differs in some crucial ways. Hence, I use the label pseudo scope marking construction to describe this phenomenon. I show that this construction is actually derived by canonical wh-movement with one small difference. What undergoes overt wh-movement is not an XP but rather a feature. Overt feature movement is licensed by the morphology of the question words in this language. Specifically, there is a free morpheme tsa, which I propose corresponds only to a [wh] feature. As a free morpheme, it is able to undergo overt movement. The discussion relies on Lasnik’s Stray Affix Filter in addition to the particular morphological properties of question words in Blackfoot to account for these facts.

Embodiment effects and language comprehension in Alzheimer's disease.

It has been shown that when participants are asked to make sensibility judgments on sentences that describe a transfer of an object toward or away from their body, they are faster to respond when the response requires a movement in the same direction as the transfer described in the sentence. This phenomenon is known as the action compatibility effect (ACE). This study investigates whether the ACE exists for volunteers with Alzheimer's disease (AD), whether the ACE can facilitate language comprehension, and also whether the ACE can still be produced if the order of the two events is inverted, that is, whether overt movement can prime comprehension of transfer sentences. In Study 1, participants with AD, younger, and older adults were tested on an adaptation of the ACE Paradigm. In Study 2, the same paradigm was modified to include an arm movement that participants had to perform prior to sentence exposure on screen. In Study 1, young, older adults, and individuals with AD were faster to respond when the direction of the response movement matched the directionality implied by the sentence (ACE). In Study 2, no traditional ACE was found; participants were faster when the direction of the movement immediately preceding the sentence matched the directionality of the sentence. It was found that compatibility effects generated a relative advantage, that transfer schemata are easier to process, and that an ACE-like effect can be the result of mutual priming between language and movement. Results suggested preservation in AD of the neural systems for action engaged during language comprehension, and conditions under which comprehension in AD can be facilitated in real life may be identified.

Clinical evidence of the role of the cerebellum in the suppression of overt articulatory movements during reading. A study of reading in children and adolescents treated for cerebellar pilocytic astrocytoma.

It has been suggested that the cerebellum is involved in reading acquisition and in particular in the progression from automatic grapheme-phoneme conversion to the internalization of speech required for silent reading. This idea is in line with clinical and neuroimaging data showing a cerebellar role in subvocal rehearsal for printed verbalizable material and with computational "internal models" of the cerebellum suggesting its role in inner speech (i.e. covert speech without mouthing the words). However, studies examining a possible cerebellar role in the suppression of articulatory movements during silent reading acquisition in children are lacking. Here, we report clinical evidence that the cerebellum plays a part in this transition. Reading performances were compared between a group of 17 paediatric patients treated for benign cerebellar tumours and a group of controls matched for age, gender, and parental socio-educational level. The patients scored significantly lower on all reading, but the most striking difference concerned silent reading, perfectly acquired by almost all controls, contrasting with 41% of the patients who were unable to read any item silently. Silent reading was correlated with the Working Memory Index. The present findings converge with previous reports on an implication of the cerebellum in inner speech and in the automatization of reading. This cerebellar implication is probably not specific to reading, as it also seems to affect non-reading tasks such as counting.

Incidental sequence learning in a motion coherence discrimination task: How response learning affects perception.

The serial reaction time task (SRTT) is a standard task used to investigate incidental sequence learning. Whereas incidental learning of motor sequences is well-established, few and disputed results support learning of perceptual sequences. Here we adapt a motion coherence discrimination task (Newsome & Paré, 1988) to the sequence learning paradigm. The new task has 2 advantages: (a) the stimulus is presented at fixation, thereby obviating overt eye movements, and (b) by varying coherence a perceptual threshold measure is available in addition to the performance measure of RT. Results from 3 experiments show that action relevance of the sequence is necessary for sequence learning to occur, that the amount of sequence knowledge varies with the ease of encoding the motor sequence, and that sequence knowledge, once acquired, has the ability to modify perceptual thresholds.

Examining intention in simulated actions: are children and young adults different?

Previous work with adults provides evidence that 'intention' used in processing simulated actions is similar to that used in planning and processing overt movements. The present study compared young adults and children on their ability to estimate distance reachability using a NOGO/GO paradigm in conditions of imagery only (IO) and imagery with actual execution (IE). Our initial thoughts were that whereas intention is associated with motivation and commitment to act, age-related differences could impact planning. Results indicated no difference in overall accuracy by condition within groups, and as expected adults were more accurate. These findings support an increasing body of evidence suggesting that the neurocognitive processes (in this case, intention) driving motor imagery and overt actions are similar, and as evidenced here, functioning by age 7.

The Body Action Coding System II: muscle activations during the perception and expression of emotion.

Research into the expression and perception of emotions has mostly focused on facial expressions. Recently, body postures have become increasingly important in research, but knowledge on muscle activity during the perception or expression of emotion is lacking. The current study continues the development of a Body Action Coding System (BACS), which was initiated in a previous study, and described the involvement of muscles in the neck, shoulders and arms during expression of fear and anger. The current study expands the BACS by assessing the activity patterns of three additional muscles. Surface electromyography of muscles in the neck (upper trapezius descendens), forearms (extensor carpi ulnaris), lower back (erector spinae longissimus) and calves (peroneus longus) were measured during active expression and passive viewing of fearful and angry body expressions. The muscles in the forearm were strongly active for anger expression and to a lesser extent for fear expression. In contrast, muscles in the calves were recruited slightly more for fearful expressions. It was also found that muscles automatically responded to the perception of emotion, without any overt movement. The observer's forearms responded to the perception of fear, while the muscles used for leaning backwards were activated when faced with an angry adversary. Lastly, the calf responded immediately when a fearful person was seen, but responded slower to anger. There is increasing interest in developing systems that are able to create or recognize emotional body language for the development of avatars, robots, and online environments. To that end, multiple coding systems have been developed that can either interpret or create bodily expressions based on static postures, motion capture data or videos. However, the BACS is the first coding system based on muscle activity.

On the asynchronously continuous control of mobile robot movement by motor cortical spiking activity.

This paper presents an asynchronously intracortical brain-computer interface (BCI) which allows the subject to continuously drive a mobile robot. This system has a great implication for disabled patients to move around. By carefully designing a multiclass support vector machine (SVM), the subject's self-paced instantaneous movement intents are continuously decoded to control the mobile robot. In particular, we studied the stability of the neural representation of the movement directions. Experimental results on the nonhuman primate showed that the overt movement directions were stably represented in ensemble of recorded units, and our SVM classifier could successfully decode such movements continuously along the desired movement path. However, the neural representation of the stop state for the self-paced control was not stably represented and could drift.

How active perception and attractor dynamics shape perceptual categorization: A computational model

We propose a computational model of perceptual categorization that fuses elements of grounded and sensorimotor theories of cognition with dynamic models of decision-making. We assume that category information consists in anticipated patterns of agent–environment interactions that can be elicited through overt or covert (simulated) eye movements, object manipulation, etc. This information is firstly encoded when category information is acquired, and then re-enacted during perceptual categorization. The perceptual categorization consists in a dynamic competition between attractors that encode the sensorimotor patterns typical of each category; action prediction success counts as “evidence” for a given category and contributes to falling into the corresponding attractor. The evidence accumulation process is guided by an active perception loop, and the active exploration of objects (e.g., visual exploration) aims at eliciting expected sensorimotor patterns that count as evidence for the object category. We present a computational model incorporating these elements and describing action prediction, active perception, and attractor dynamics as key elements of perceptual categorizations. We test the model in three simulated perceptual categorization tasks, and we discuss its relevance for grounded and sensorimotor theories of cognition.

Dopamine precursors depletion impairs impulse control in healthy volunteers.

The aim of the present study was to decipher the role of the dopamine system in impulse control. Impulsive actions entail (i) activation of the motor system by an impulse, which is an urge to act and (ii) a failure to suppress that impulse, when inappropriate, in order to prevent an error. These two aspects of action impulsivity can be experimentally disentangled in conflict reaction time tasks such as the Simon task, which measures susceptibility to acting on spontaneous impulses (as well as the proficiency of suppressing these impulses). In 12 healthy volunteers performing a Simon task, dopamine availability was reduced with an amino acid drink deficient in the dopamine precursors, phenylalanine and tyrosine. Classic behavioral measures were augmented with an analysis of the electromyographic activity of the response effectors. Electromyography allows one to detect covert activations undetectable with strictly behavioral measures and further reveals the participants' ability to quickly suppress covert activations before they result in an overt movement. Following dopamine depletion, compared with a placebo condition, participants displayed comparable impulse activation but were less proficient at suppressing the interference from this activation. These results provide evidence that the dopamine system is directly involved in the suppression of maladaptive response impulses.

Identification of a Brainstem Circuit Regulating Visual Cortical State in Parallel with Locomotion

Sensory processing is dependent upon behavioral state. In mice, locomotion is accompanied by changes in cortical state and enhanced visual responses. Although recent studies have begun to elucidate intrinsic cortical mechanisms underlying this effect, the neural circuits that initially couple locomotion to cortical processing are unknown. The mesencephalic locomotor region (MLR) has been shown to be capable of initiating running and is associated with the ascending reticular activating system. Here, we find that optogenetic stimulation of the MLR in awake, head-fixed mice can induce both locomotion and increases in the gain of cortical responses. MLR stimulation below the threshold for overt movement similarly changed cortical processing, revealing that MLR's effects on cortex are dissociable from locomotion. Likewise, stimulation of MLR projections to the basal forebrain also enhanced cortical responses, suggesting a pathway linking the MLR to cortex. These studies demonstrate that the MLR regulates cortical state in parallel with locomotion.

Reorganization of large-scale cognitive networks during automation of imagination of a complex sequential movement

We investigated the functional reconfiguration of the cerebral networks involved in imagination of sequential movements of the left foot, both performed at regular and fast speed after mental imagery training. Thirty-five volunteers were scanned with a 3T MRI while they imagined a sequence of ankle movements (dorsiflexion, plantar flexion, varus and valgus) before and after mental practice. Subjects were distributed in two groups: the first group executed regular movements whereas the second group made fast movements. We applied the general linear model (GLM) and model-free, exploratory tensorial independent component analytic (TICA) approaches to identify plastic post-training effects on brain activation. GLM showed that post-training imagination of movement was accompanied by a dual effect: a specific recruitment of a medial prefronto-cingulo-parietal circuit reminiscent of the default-mode network, with the left putamen, and a decreased activity of a lateral fronto-parietal network. Training-related subcortical changes only consisted in an increased activity in the left striatum. Unexpectedly, no difference was observed in the cerebellum. TICA also revealed involvement of the left executive network, and of the dorsal control executive network but no significant differences were found between pre- and post-training phases. Therefore, repetitive motor mental imagery induced specific putamen (motor rehearsal) recruitment that one previously observed during learning of overt movements, and, simultaneously, a specific shift of activity from the dorsolateral prefrontal cortex (attention, working memory) to the medial posterior parietal and cingulate cortices (mental imagery and memory rehearsal). Our data complement and confirm the notion that differential and coupled recruitment of cognitive networks can constitute a neural marker of training effects.

Wh-Fronting in a two-probe system

Prior work on wh-movement has distinguished among several types of wh-fronting languages that permit distinct patterns of overt and covert movement, instantiated for example by the Slavic languages, English, and German. This paper extends the cross-linguistic typology of multiple questions by arguing that Hebrew instantiates a new kind of wh-fronting language, unlike any that are discussed in the current literature. It will show that Hebrew distinguishes between two kinds of interrogative phrases: those that are headed by a wh-word (wh-headed phrases: what, who, [DP which X], where, how…) and those that contain a wh-word but are headed by some other element (wh-containing phrases: [NP N of wh], [PP P wh]). We observe the special status of wh-headed phrases when one occurs structurally lower in a question than a wh-containing phrase. In that case, the wh-headed phrase can be targeted by an Agree/Attract operation that ignores the presence of the c-commanding wh-containing phrase. The paper develops an account of the sensitivity of interrogative probing operations to the head of the interrogative phrase within Cable’s (2010) Q-particle theory. It proposes that the Hebrew Q has an EPP feature which can trigger head-movement of wh to Q and that a wh-probe exists alongside the more familiar Q-probe, and shows how these two modifications to the theory can account for the intricate dataset that emerges from the paper. The emerging picture is one in which interrogative probing does not occur wholesale but rather can be sensitive to particular interrogative features on potential goals.

The evolutionary neuroscience of musical beat perception: the Action Simulation for Auditory Prediction (ASAP) hypothesis.

Every human culture has some form of music with a beat: a perceived periodic pulse that structures the perception of musical rhythm and which serves as a framework for synchronized movement to music. What are the neural mechanisms of musical beat perception, and how did they evolve? One view, which dates back to Darwin and implicitly informs some current models of beat perception, is that the relevant neural mechanisms are relatively general and are widespread among animal species. On the basis of recent neural and cross-species data on musical beat processing, this paper argues for a different view. Here we argue that beat perception is a complex brain function involving temporally-precise communication between auditory regions and motor planning regions of the cortex (even in the absence of overt movement). More specifically, we propose that simulation of periodic movement in motor planning regions provides a neural signal that helps the auditory system predict the timing of upcoming beats. This “action simulation for auditory prediction” (ASAP) hypothesis leads to testable predictions. We further suggest that ASAP relies on dorsal auditory pathway connections between auditory regions and motor planning regions via the parietal cortex, and suggest that these connections may be stronger in humans than in non-human primates due to the evolution of vocal learning in our lineage. This suggestion motivates cross-species research to determine which species are capable of human-like beat perception, i.e., beat perception that involves accurate temporal prediction of beat times across a fairly broad range of tempi.

A Fully Automated Rodent Conditioning Protocol for Sensorimotor Integration and Cognitive Control Experiments

Rodents have been traditionally used as a standard animal model in laboratory experiments involving a myriad of sensory, cognitive, and motor tasks. Higher cognitive functions that require precise control over sensorimotor responses such as decision-making and attentional modulation, however, are typically assessed in nonhuman primates. Despite the richness of primate behavior that allows multiple variants of these functions to be studied, the rodent model remains an attractive, cost-effective alternative to primate models. Furthermore, the ability to fully automate operant conditioning in rodents adds unique advantages over the labor intensive training of nonhuman primates while studying a broad range of these complex functions. Here, we introduce a protocol for operantly conditioning rats on performing working memory tasks. During critical epochs of the task, the protocol ensures that the animal's overt movement is minimized by requiring the animal to 'fixate' until a Go cue is delivered, akin to nonhuman primate experimental design. A simple two alternative forced choice task is implemented to demonstrate the performance. We discuss the application of this paradigm to other tasks.

A fully automated rodent conditioning protocol for sensorimotor integration and cognitive control experiments.

Rodents have been traditionally used as a standard animal model in laboratory experiments involving a myriad of sensory, cognitive, and motor tasks. Higher cognitive functions that require precise control over sensorimotor responses such as decision-making and attentional modulation, however, are typically assessed in nonhuman primates. Despite the richness of primate behavior that allows multiple variants of these functions to be studied, the rodent model remains an attractive, cost-effective alternative to primate models. Furthermore, the ability to fully automate operant conditioning in rodents adds unique advantages over the labor intensive training of nonhuman primates while studying a broad range of these complex functions. Here, we introduce a protocol for operantly conditioning rats on performing working memory tasks. During critical epochs of the task, the protocol ensures that the animal's overt movement is minimized by requiring the animal to 'fixate' until a Go cue is delivered, akin to nonhuman primate experimental design. A simple two alternative forced choice task is implemented to demonstrate the performance. We discuss the application of this paradigm to other tasks.

Differential Contribution of Bilateral Supplementary Motor Area to the Effective Connectivity Networks Induced by Task Conditions Using Dynamic Causal Modeling

Functional imaging studies have indicated hemispheric asymmetry of activation in bilateral supplementary motor area (SMA) during unimanual motor tasks. However, the hemispherically special roles of bilateral SMAs on primary motor cortex (M1) in the effective connectivity networks (ECN) during lateralized tasks remain unclear. Aiming to study the differential contribution of bilateral SMAs during the motor execution and motor imagery tasks, and the hemispherically asymmetric patterns of ECN among regions involved, the present study used dynamic causal modeling to analyze the functional magnetic resonance imaging data of the unimanual motor execution/imagery tasks in 12 right-handed subjects. Our results demonstrated that distributions of network parameters underlying motor execution and motor imagery were significantly different. The variation was mainly induced by task condition modulations of intrinsic coupling. Particularly, regardless of the performing hand, the task input modulations of intrinsic coupling from the contralateral SMA to contralateral M1 were positive during motor execution, while varied to be negative during motor imagery. The results suggested that the inhibitive modulation suppressed the overt movement during motor imagery. In addition, the left SMA also helped accomplishing left hand tasks through task input modulation of left SMA→right SMA connection, implying that hemispheric recruitment occurred when performing nondominant hand tasks. The results specified differential and altered contributions of bilateral SMAs to the ECN during unimanual motor execution and motor imagery, and highlighted the contributions induced by the task input of motor execution/imagery.

Fox Squirrels Match Food Assessment and Cache Effort to Value and Scarcity

Scatter hoarders must allocate time to assess items for caching, and to carry and bury each cache. Such decisions should be driven by economic variables, such as the value of the individual food items, the scarcity of these items, competition for food items and risk of pilferage by conspecifics. The fox squirrel, an obligate scatter-hoarder, assesses cacheable food items using two overt movements, head flicks and paw manipulations. These behaviors allow an examination of squirrel decision processes when storing food for winter survival. We measured wild squirrels' time allocations and frequencies of assessment and investment behaviors during periods of food scarcity (summer) and abundance (fall), giving the squirrels a series of 15 items (alternating five hazelnuts and five peanuts). Assessment and investment per cache increased when resource value was higher (hazelnuts) or resources were scarcer (summer), but decreased as scarcity declined (end of sessions). This is the first study to show that assessment behaviors change in response to factors that indicate daily and seasonal resource abundance, and that these factors may interact in complex ways to affect food storing decisions. Food-storing tree squirrels may be a useful and important model species to understand the complex economic decisions made under natural conditions.

Overt head movements moderate the effect of depressive symptoms on mood regulation

A dysfunction in the regulation of negative mood states is one of the core symptoms of depression. Research has found that levels of depression are associated with the intensity of the mood-regulation deficit. The present study aimed to explore the role the body plays in mood-regulation processes. More specifically, we studied whether head movements can influence mood persistence in dysphoric states. Subsequent to a sad-mood induction, participants were presented with a set of positive pictures immediately after performing either vertical (i.e., nodding) or lateral (i.e., shaking) head movements. We considered changes in mood from before to after the experimental task as an index of the effectiveness of mood regulation. As expected, the results showed that higher initial levels of depressive symptoms were associated with greater persistence of sad mood. More importantly, this association was present in participants who shook their heads, but not in those who nodded. These results show that body movements can contribute to mood-regulation processes, thus expanding our knowledge of the psychopathology of mood disorders.