The Neural Basis of Mark Making: A Functional MRI Study of Drawing

Compared to most other forms of visually-guided motor activity, drawing is unique in that it “leaves a trail behind” in the form of the emanating image. We took advantage of an MRI-compatible drawing tablet in order to examine both the motor production and perceptual emanation of images. Subjects participated in a series of mark making tasks in which they were cued to draw geometric patterns on the tablet''s surface. The critical comparison was between when visual feedback was displayed (image generation) versus when it was not (no image generation). This contrast revealed an occipito-parietal stream involved in motion-based perception of the emerging image, including areas V5/MT+, LO, V3A, and the posterior part of the intraparietal sulcus. Interestingly, when subjects passively viewed animations of visual patterns emerging on the projected surface, all of the sensorimotor network involved in drawing was strongly activated, with the exception of the primary motor cortex. These results argue that the origin of the human capacity to draw and write involves not only motor skills for tool use but also motor-sensory links between drawing movements and the visual images that emanate from them in real time.     

Features of corticofugal effects of the motor area of the cortex on processing of heterogeneous afferentation in the parafascicular complex of the thalamus in the cat

Evoked potentials to somatic and visual stimuli were recorded in the parafascicular complex (parafascicular nuclei--centrum medianum--Pf--CM) of the thalamus of cats anaesthetized by nembutal. Cooling of the motor cortex was also used. The influence of the motor cortex on processing of the visual and somatic afferent signals at the thalamic level was found to be direct but different by its character. The motor cortex exerted unidirectional facilitatory modulatory action of a tonic type on the processing of visual afferentation and general facilitatory influence against the background of which particular excitatory and inhibitory effects were seen which it exerted on the processing of somatic signals. Episodically the motor cortex completely controlled the afferent inputs activated by somatic impulses. The motor cortical area non-equally influenced afferentation of the same modality forming the different components of the evoked potentials in Pf--CM. On the base of our present and earlier obtained data an idea has been formed of existence of a general principle of differentiated influences of polysensory areas on heterogeneous afferentation on nonspecific and association thalamic nuclei, and of realization of these influences through separation of functionally isolated subsystems in descending pathways. Each of the subsystems by closing separate thalamo-cortical circuits might transmit signals of a single modality.     

Can ambient odors influence the recognition of emotional words? A behavioral and event-related potentials study

Odor context can affect the recognition of facial expressions. However, there is no evidence to date that odor can regulate the processing of emotional words conveyed by visual words. An emotional word recognition task was combined with event-related potential technology. Briefly, 49 adults were randomly divided into three odor contexts (pleasant odor, unpleasant odor, and no odor) to judge the valence of emotional words (positive, negative, and neutral). Both behavioral and Electroencephalography (EEG) data were collected. Both the pleasant odor and unpleasant odor contexts shortened the response time of the subjects to emotional words. In addition, negative words induced greater amplitudes of early posterior negativity (EPN) and late positive potential (LPP) than the positive and neutral words. However, the neutral words induced a larger N400 amplitude than the positive and negative words. More importantly, the processing of emotional words was found to be modulated by external odor contexts. For example, during the earlier (P2) processing stages, pleasant and unpleasant odor contexts induced greater P2 amplitudes than the no odor context. In the unpleasant odor context, negative words with the same odor valence induced greater P2 amplitudes than the positive words. During the later (N400) stages, various regions of the brain regions exhibited different results. For example, in the left and right frontal areas of the brain, exposure to positive words in a pleasant odor context resulted in a smaller N400 amplitude than exposure to neutral words in the same context. Meanwhile, in the left and right central regions, emotional words with the same valence as pleasant or unpleasant odor contexts elicited the minimum N400 amplitude. Individuals are very sensitive to emotional information. With deeper processing, different cognitive processes are reflected and they can be modulated by external odors. In the early and late stages of word processing, both pleasant and unpleasant odor contexts exhibited an undifferentiated dominance effect and could specifically modulate affectively congruent words.     

Modulation of Arm Reaching Movements during Processing of Arm/Hand-Related Action Verbs with and without Emotional Connotation

The theory of embodied language states that language comprehension relies on an internal reenactment of the sensorimotor experience associated with the processed word or sentence. Most evidence in support of this hypothesis had been collected using linguistic material without any emotional connotation. For instance, it had been shown that processing of arm-related verbs, but not of those leg-related verbs, affects the planning and execution of reaching movements; however, at present it is unknown whether this effect is further modulated by verbs evoking an emotional experience. Showing such a modulation might shed light on a very debated issue, i.e. the way in which the emotional meaning of a word is processed. To this end, we assessed whether processing arm/hand-related verbs describing actions with negative connotations (e.g. to stab) affects reaching movements differently from arm/hand-related verbs describing actions with neutral connotation (e.g. to comb). We exploited a go/no-go paradigm in which healthy participants were required to perform arm-reaching movements toward a target when verbs expressing emotional hand actions, neutral hand actions or foot actions were shown, and to refrain from moving when no-effector-related verbs were presented. Reaction times and percentages of errors increased when the verb involved the same effector as used to give the response. However, we also found that the size of this interference decreased when the arm/hand-related verbs had a negative emotional connotation. Crucially, we show that such modulation only occurred when the verb semantics had to be retrieved. These results suggest that the comprehension of negatively valenced verbs might require the simultaneous reenactment of the neural circuitry associated with the processing of the emotion evoked by their meaning and of the neural circuitry associated with their motor features.     

The influence of emotional coloring of images on visual working memory in adults and adolescents

The influence of emotional valence (positive, negative or neutral) of realistic images on the functioning of visual working memory (WM) was studied in adults (n = 40) and adolescents (n = 17). In adults, emotional coloring of stimuli increased the reaction time and decreased the accuracy of WM task performance. This effect was more pronounced for negative than for positive valence: the minimal reaction time was observed for the neutral stimuli, the maximal for the negative emotional stimuli, and significant differences in the reaction time were found between all three types of images. The accuracy was lower for negative stimuli than for either positive or neutral stimuli. Compared with adults, adolescents of age 14–16 showed lower indices of the performance accuracy and rate with neutral and positive stimuli in the WM task. In this group, no significant influence of the emotional valence of visual stimuli on the accuracy of WM task performance was found.     

Valence-specific modulation in the accumulation of perceptual evidence prior to visual scene recognition

Visual scene recognition is a dynamic process through which incoming sensory information is iteratively compared with predictions regarding the most likely identity of the input stimulus. In this study, we used a novel progressive unfolding task to characterize the accumulation of perceptual evidence prior to scene recognition, and its potential modulation by the emotional valence of these scenes. Our results show that emotional (pleasant and unpleasant) scenes led to slower accumulation of evidence compared to neutral scenes. In addition, when controlling for the potential contribution of non-emotional factors (i.e., familiarity and complexity of the pictures), our results confirm a reliable shift in the accumulation of evidence for pleasant relative to neutral and unpleasant scenes, suggesting a valence-specific effect. These findings indicate that proactive iterations between sensory processing and top-down predictions during scene recognition are reliably influenced by the rapidly extracted (positive) emotional valence of the visual stimuli. We interpret these findings in accordance with the notion of a genuine positivity offset during emotional scene recognition.     

Lower visual field advantage for motion segmentation during high competition for selection

A series of visual enumeration tasks were conducted investigating the role of the dorsal visual stream in motion segmentation. Cortical areas representing the lower visual field have greater connections with the parietal cortex and should therefore show an advantage for processes driven by the dorsal stream (Previc, 1990). We looked for differences in processing displays in the upper versus lower visual field when targets required segmentation from distractors in an enumeration task. In a baseline condition, random configurations of moving and static items were presented briefly (200 ms) to the upper or lower visual field. Fast and efficient enumeration took place both for moving targets and for static targets presented alone; there was no effect of visual field. In contrast, for moving targets, a lower visual field advantage was found when the inclusion of static distractors demanded segmentation by motion. This disappeared at the smaller display sizes when the targets were presented in canonical patterns. The results are consistent with segmentation of moving targets from static distractors being mediated by dorsal regions of the visual cortex, particularly under conditions of high load (non-canonical patterns). These regions show greater sensitivity to the lower visual field and to magnocellular-based input.     

Attentional Demands Influence Vocal Compensations to Pitch Errors Heard in Auditory Feedback

Auditory feedback is required to maintain fluent speech. At present, it is unclear how attention modulates auditory feedback processing during ongoing speech. In this event-related potential (ERP) study, participants vocalized/a/, while they heard their vocal pitch suddenly shifted downward a ½ semitone in both single and dual-task conditions. During the single-task condition participants passively viewed a visual stream for cues to start and stop vocalizing. In the dual-task condition, participants vocalized while they identified target stimuli in a visual stream of letters. The presentation rate of the visual stimuli was manipulated in the dual-task condition in order to produce a low, intermediate, and high attentional load. Visual target identification accuracy was lowest in the high attentional load condition, indicating that attentional load was successfully manipulated. Results further showed that participants who were exposed to the single-task condition, prior to the dual-task condition, produced larger vocal compensations during the single-task condition. Thus, when participants’ attention was divided, less attention was available for the monitoring of their auditory feedback, resulting in smaller compensatory vocal responses. However, P1-N1-P2 ERP responses were not affected by divided attention, suggesting that the effect of attentional load was not on the auditory processing of pitch altered feedback, but instead it interfered with the integration of auditory and motor information, or motor control itself.     

Emotional Picture and Word Processing: An fMRI Study on Effects of Stimulus Complexity

Neuroscientific investigations regarding aspects of emotional experiences usually focus on one stimulus modality (e.g., pictorial or verbal). Similarities and differences in the processing between the different modalities have rarely been studied directly. The comparison of verbal and pictorial emotional stimuli often reveals a processing advantage of emotional pictures in terms of larger or more pronounced emotion effects evoked by pictorial stimuli. In this study, we examined whether this picture advantage refers to general processing differences or whether it might partly be attributed to differences in visual complexity between pictures and words. We first developed a new stimulus database comprising valence and arousal ratings for more than 200 concrete objects representable in different modalities including different levels of complexity: words, phrases, pictograms, and photographs. Using fMRI we then studied the neural correlates of the processing of these emotional stimuli in a valence judgment task, in which the stimulus material was controlled for differences in emotional arousal. No superiority for the pictorial stimuli was found in terms of emotional information processing with differences between modalities being revealed mainly in perceptual processing regions. While visual complexity might partly account for previously found differences in emotional stimulus processing, the main existing processing differences are probably due to enhanced processing in modality specific perceptual regions. We would suggest that both pictures and words elicit emotional responses with no general superiority for either stimulus modality, while emotional responses to pictures are modulated by perceptual stimulus features, such as picture complexity.     

Fixational Eye Movements in the Earliest Stage of Metazoan Evolution

All known photoreceptor cells adapt to constant light stimuli, fading the retinal image when exposed to an immobile visual scene. Counter strategies are therefore necessary to prevent blindness, and in mammals this is accomplished by fixational eye movements. Cubomedusae occupy a key position for understanding the evolution of complex visual systems and their eyes are assumedly subject to the same adaptive problems as the vertebrate eye, but lack motor control of their visual system. The morphology of the visual system of cubomedusae ensures a constant orientation of the eyes and a clear division of the visual field, but thereby also a constant retinal image when exposed to stationary visual scenes. Here we show that bell contractions used for swimming in the medusae refresh the retinal image in the upper lens eye of Tripedalia cystophora. This strongly suggests that strategies comparable to fixational eye movements have evolved at the earliest metazoan stage to compensate for the intrinsic property of the photoreceptors. Since the timing and amplitude of the rhopalial movements concur with the spatial and temporal resolution of the eye it circumvents the need for post processing in the central nervous system to remove image blur.     

Retinotopic organization of the posterior suprasylvian area of the feline cerebral cortex

Representation of the visual field was investigated in the feline posterior suprasylvian area (PSA) using electrophysiological mapping techniques. The PSA is one of the extrastriatal visual structures of the cerebral cortex. The PSA retinotopic organization pattern was also studied. Neuronal receptive fields (RF) were mainly located in the upper contralateral quadrant and just a small number in the lower contralateral quadrant of the visual field. Approximately 10% of RF were located in the upper ipsilateral quadrant. The central area of the visual field extending in a radius of 20–30° from the area centralis was mainly represented in the upper section of the PSA (areas 21a and 21b). The RP of neurons located more peripherally to the area centralis are found in the lower portion of the PSA (areas 20a and 20b); these occupy a correspondingly greater area. Experimental finding did not confirm any substantial differences in the retinotopic organization of areas 21a, 21b, 20a, and 20b comprising the PSA. Data obtained would tend to indicate that the PSA consists of two areas, 21a and 21b, which do not appear to be subdivided, with more densely distributed visual neurons in the former than in the latter.Institute of Experimental Biology of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 23, No. 3, pp. 290–296, May–June, 1991.     

The Role of the Caudal Superior Parietal Lobule in Updating Hand Location in Peripheral Vision: Further Evidence from Optic Ataxia

Patients with optic ataxia (OA), who are missing the caudal portion of their superior parietal lobule (SPL), have difficulty performing visually-guided reaches towards extra-foveal targets. Such gaze and hand decoupling also occurs in commonly performed non-standard visuomotor transformations such as the use of a computer mouse. In this study, we test two unilateral OA patients in conditions of 1) a change in the physical location of the visual stimulus relative to the plane of the limb movement, 2) a cue that signals a required limb movement 180° opposite to the cued visual target location, or 3) both of these situations combined. In these non-standard visuomotor transformations, the OA deficit is not observed as the well-documented field-dependent misreach. Instead, OA patients make additional eye movements to update hand and goal location during motor execution in order to complete these slow movements. Overall, the OA patients struggled when having to guide centrifugal movements in peripheral vision, even when they were instructed from visual stimuli that could be foveated. We propose that an intact caudal SPL is crucial for any visuomotor control that involves updating ongoing hand location in space without foveating it, i.e. from peripheral vision, proprioceptive or predictive information.     

Parkinson-Related Changes of Activation in Visuomotor Brain Regions during Perceived Forward Self-Motion

Radial expanding optic flow is a visual consequence of forward locomotion. Presented on screen, it generates illusionary forward self-motion, pointing at a close vision-gait interrelation. As particularly parkinsonian gait is vulnerable to external stimuli, effects of optic flow on motor-related cerebral circuitry were explored with functional magnetic resonance imaging in healthy controls (HC) and patients with Parkinson’s disease (PD). Fifteen HC and 22 PD patients, of which 7 experienced freezing of gait (FOG), watched wide-field flow, interruptions by narrowing or deceleration and equivalent control conditions with static dots. Statistical parametric mapping revealed that wide-field flow interruption evoked activation of the (pre-)supplementary motor area (SMA) in HC, which was decreased in PD. During wide-field flow, dorsal occipito-parietal activations were reduced in PD relative to HC, with stronger functional connectivity between right visual motion area V5, pre-SMA and cerebellum (in PD without FOG). Non-specific ‘changes’ in stimulus patterns activated dorsolateral fronto-parietal regions and the fusiform gyrus. This attention-associated network was stronger activated in HC than in PD. PD patients thus appeared compromised in recruiting medial frontal regions facilitating internally generated virtual locomotion when visual motion support falls away. Reduced dorsal visual and parietal activations during wide-field optic flow in PD were explained by impaired feedforward visual and visuomotor processing within a magnocellular (visual motion) functional chain. Compensation of impaired feedforward processing by distant fronto-cerebellar circuitry in PD is consistent with motor responses to visual motion stimuli being either too strong or too weak. The ‘change’-related activations pointed at covert (stimulus-driven) attention.     

Altered negative unconscious processing in major depressive disorder: an exploratory neuropsychological study

Objective

Major depressive disorder (MDD) has been characterized by abnormalities in emotional processing. However, what remains unclear is whether MDD also shows deficits in the unconscious processing of either positive or negative emotions. We conducted a psychological study in healthy and MDD subjects to investigate unconscious emotion processing and its valence-specific alterations in MDD patients.

Methods

We combined a well established paradigm for unconscious visual processing, the continuous flash suppression, with positive and negative emotional valences to detect the attentional preference evoked by the invisible emotional facial expressions.

Results

Healthy subjects showed an attentional bias for negative emotions in the unconscious condition while this valence bias remained absent in MDD patients. In contrast, this attentional bias diminished in the conscious condition for both healthy subjects and MDD.

Conclusion

Our findings demonstrate for the first time valence-specific deficits specifically in the unconscious processing of emotions in MDD; this may have major implications for subsequent neurobiological investigations as well as for clinical diagnosis and therapy.     

Links among emotional awareness,somatic awareness and autonomic homeostatic processing

Emotional awareness and somatic interoceptive awareness are essential processes for human psychosomatic health. A typical trait of lacking emotional awareness related to psychosomatic symptoms is alexithymia. In contrast, alexisomia refers to the trait of lacking somatic awareness. Links between emotional and somatic awareness and homeostatic processing are also significant for the psychosomatic health. The purpose of the present paper is to review the links among emotional awareness, somatic interoceptive awareness and autonomic homeostatic processing. On the basis of the collected evidence, the following arguments were presented¹: (1) The main subcortical neural substrates for these processes are limbic-related systems, which are also responsible for autonomic functions for optimization of homeostatic efficiency. (2) Considerable studies have shown that autonomic activity and/or reactivity to stress correlate with both emotional and interoceptive awareness. A hypothesis was advocated about the links between the two types of awareness and autonomic function: Autonomic dysfunction, especially high sympathetic tone at baseline and/or attenuated reactivity or variability to stress, appears to be involved in disturbance of emotional and interoceptive awareness. (3) Several studies suggest that a link or a cooperative relationship exists between emotional and somatic awareness, and that somatic awareness is the more fundamental of the two types of awareness. Emotional awareness, somatic awareness and autonomic homeostatic processing generally occur in parallel or concurrently. However, some complex features of pathologies include coexistence of reduced interoceptive awareness and somatosensory amplification. The autonomic homeostatic process is fundamentally involved in emotional and somatic awareness. Investigation of these types of awareness with both neuroimaging evaluations and estimation of peripheral autonomic function are required as next steps for exploration of the relationship between awareness and human somatic states including somatic symptoms as well as general psychosomatic health.     

Sex differences in the cerebral lateralization of a cichlid fish when detouring to view emotionally conditioned stimuli

The lateralization of emotion has been described in a variety of animals. The right hemisphere has been implicated in the processing of negative emotions while positive emotions are processed in the left. Most animal studies of this phenomenon to date have used intrinsically emotionally arousing stimuli and there are few examples of lateralized responses to learned emotional triggers. It is known that males and females may demonstrate different patterns of lateralization, and that these sex differences may interact with other variables. We investigated the lateralized response of male and female convict cichlids to emotionally conditioned stimuli. One stimulus was given an appetitive (positive emotional valence) association by pairing with food. The other stimulus was given an aversive (negative emotional valence) association by pairing with a chemical alarm signal. We found that males tend to be more strongly lateralized to aversive stimuli while females are more strongly lateralized when responding to appetitive stimuli.     

Humans rely on the same rules to assess emotional valence and intensity in conspecific and dog vocalizations

Humans excel at assessing conspecific emotional valence and intensity, based solely on non-verbal vocal bursts that are also common in other mammals. It is not known, however, whether human listeners rely on similar acoustic cues to assess emotional content in conspecific and heterospecific vocalizations, and which acoustical parameters affect their performance. Here, for the first time, we directly compared the emotional valence and intensity perception of dog and human non-verbal vocalizations. We revealed similar relationships between acoustic features and emotional valence and intensity ratings of human and dog vocalizations: those with shorter call lengths were rated as more positive, whereas those with a higher pitch were rated as more intense. Our findings demonstrate that humans rate conspecific emotional vocalizations along basic acoustic rules, and that they apply similar rules when processing dog vocal expressions. This suggests that humans may utilize similar mental mechanisms for recognizing human and heterospecific vocal emotions.     

Mental representations of movements. Brain potentials associated with imagination of hand movements

The present study was designed in order to contribute towards the understanding of the physiology of motor imagery. DC potentials were recorded when subjects either imagined or executed a sequence of unilateral or bilateral hand movements. The sequence consisted of hand movements in 4 directions, forwards, backwards, to the right and to the left, and varied from trial to trial. The sequence had been cued by visual targets on a computer screen and had to be memorized before the trial was initiated. Changes of DC potentials between task execution and imagination were localized in central recordings (C3, Cz, C4) with larger amplitudes when executing the task than when imagining to do so. Stimulation of peripheral receptors associated with task execution or a different level of activation of the cortico-motoneural system could account for this finding. The main result of the present study was that with unilateral performance, the side of the performing hand (right, left) had localized effects in recordings over the sensorimotor hand area (C3, C4) which were qualitatively the same with imagination and execution and quantitatively similar (i.e., without significant difference). Performance of the right hand augmented negative DC potentials in C3, performance of the left hand augmented amplitudes in C4. This result is consistent with the assumption that the primary motor cortex is active with motor imagery. Finally, the question has been addressed whether motor imagery may involve the left hemisphere to a larger extent than the execution of the movement. It is shown that a particular contribution of the left hemisphere associated with motor imagery may only show up under strictly controlled conditions.     

It’s All in the Eyes: Subcortical and Cortical Activation during Grotesqueness Perception in Autism

Atypical face processing plays a key role in social interaction difficulties encountered by individuals with autism. In the current fMRI study, the Thatcher illusion was used to investigate several aspects of face processing in 20 young adults with high-functioning autism spectrum disorder (ASD) and 20 matched neurotypical controls. “Thatcherized” stimuli were modified at either the eyes or the mouth and participants discriminated between pairs of faces while cued to attend to either of these features in upright and inverted orientation. Behavioral data confirmed sensitivity to the illusion and intact configural processing in ASD. Directing attention towards the eyes vs. the mouth in upright faces in ASD led to (1) improved discrimination accuracy; (2) increased activation in areas involved in social and emotional processing; (3) increased activation in subcortical face-processing areas. Our findings show that when explicitly cued to attend to the eyes, activation of cortical areas involved in face processing, including its social and emotional aspects, can be enhanced in autism. This suggests that impairments in face processing in autism may be caused by a deficit in social attention, and that giving specific cues to attend to the eye-region when performing behavioral therapies aimed at improving social skills may result in a better outcome.     

Brain activity elicited by positive and negative feedback in preschool-aged children

To investigate the processing of positive vs. negative feedback in children aged 4-5 years, we devised a prize-guessing game that is analogous to gambling tasks used to measure feedback-related brain responses in adult studies. Unlike adult studies, the feedback-related negativity (FRN) elicited by positive feedback was as large as that elicited by negative feedback, suggesting that the neural system underlying the FRN may not process feedback valence in early childhood. In addition, positive feedback, compared with negative feedback, evoked a larger P1 over the occipital scalp area and a larger positive slow wave (PSW) over the right central-parietal scalp area. We believe that the PSW is related to emotional arousal and the intensive focus on positive feedback that is present in the preschool and early school years has adaptive significance for both cognitive and emotional development during this period.