Hannah L. Filmer

Applications of transcranial direct current stimulation for understanding brain function

In recent years there has been an exponential rise in the number of studies employing transcranial direct current stimulation (tDCS) as a means of gaining a systems-level understanding of the cortical substrates underlying behaviour. These advances have allowed inferences to be made regarding the neural operations that shape perception, cognition, and action. Here we summarise how tDCS works, and show how research using this technique is expanding our understanding of the neural basis of cognitive and motor training. We also explain how oscillatory tDCS can elucidate the role of fluctuations in neural activity, in both frequency and phase, in perception, learning, and memory. Finally, we highlight some key methodological issues for tDCS and suggest how these can be addressed.

Improved multitasking following prefrontal tDCS

We have a limited capacity for mapping sensory information onto motor responses. This processing bottleneck is thought to be a key factor in determining our ability to make two decisions simultaneously - i.e., to multitask (Pashler, 1984, 1994; Welford, 1952). Previous functional imaging research (Dux, Ivanoff, Asplund, & Marois, 2006; Dux et al., 2009) has localised this bottleneck to the posterior lateral prefrontal cortex (pLPFC) of the left hemisphere. Currently, however, it is unknown whether this region is causally involved in multitasking performance. We investigated the role of the left pLPFC in multitasking using transcranial direct current stimulation (tDCS). The behavioural paradigm included single- and dual-task trials, each requiring a speeded discrimination of visual stimuli alone, auditory stimuli alone, or both visual and auditory stimuli. Reaction times for single- and dual-task trials were compared before, immediately after, and 20 min after anodal stimulation (excitatory), cathodal stimulation (inhibitory), or sham stimulation. The cost of responding to the two tasks (i.e., the reduction in performance for dual- vs single-task trials) was significantly reduced by cathodal stimulation, but not by anodal or sham stimulation. Overall, the results provide direct evidence that the left pLPFC is a key neural locus of the central bottleneck that limits an individuals ability to make two simple decisions simultaneously.

Size (mostly) doesn’t matter: the role of set size in object substitution masking

Conscious detection and discrimination of a visual target stimulus can be prevented by the presentation a spatially nonoverlapping, but temporally trailing, visual masking stimulus. This phenomenon, known as object substitution masking (OSM), has long been associated with spatial attention, with diffuse attention seemingly being key for the effect to be observed. Recently, this hypothesis has been questioned. We sought to provide a definitive test of the involvement of spatial attention in OSM by using an eight-alternative forced choice task under a range of mask durations, set sizes, and target/distractor spatial configurations. The results provide very little evidence that set size, and thus the distribution of spatial attention, interacts with masking magnitude. These findings have implications for understanding the mechanisms underlying OSM and the relationship between consciousness and attention.

TMS to V1 spares discrimination of emotive relative to neutral body postures.

This study used TMS to examine the role played by striate cortex (V1) in processing the emotional content of visual stimuli. Participants learned to discriminate two sets of body posture images. For half of each set, the postures emotional significance (threat versus pleasant) provided a redundant cue for the discrimination; the other half were emotionally neutral. An image was briefly presented at a lateral location in the visual field where a TMS pulse produced a phosphene, or at a control location in the opposite hemifield. A TMS pulse 70-140 ms after stimulus presentation at the phosphene location impaired discrimination of neutral stimuli with little effect on discrimination of emotional stimuli; the two classes of stimuli were equally discriminable when presented at the control location. The results are consistent with the proposal that emotionally salient patterns, such as social threat, can be discriminated independently of the geniculo-striate pathway.

Dissociable effects of anodal and cathodal tDCS reveal distinct functional roles for right parietal cortex in the detection of single and competing stimuli

Spatial attention can be used to direct neural processing resources to a subset of task-relevant or otherwise salient items within the environment. Such selective processes are particularly important for resolving competition between multiple stimuli. Deficits in processing single stimuli can arise after damage to parietal, frontal and temporal brain regions, as is typical in patients with contralesional spatial neglect. By contrast, deficits in processing multiple competing stimuli may arise specifically following lesions of the posterior parietal cortex (PPC), as occurs in the disorder of spatial extinction. It remains unclear, however, whether mechanisms involved in selecting single and competing stimuli reflect the same or dissociable neural operations within the PPC. To address this issue, in separate sessions, we applied transcranial direct current stimulation (tDCS) to the left or right PPC and measured the effect on detecting and discriminating single and competing visual stimulus events. Our results revealed reliable tDCS modulations of stimulus processing, specific to the right PPC, as well as a dissociation in the detection of single and competing stimuli. For the right PPC only, single stimuli presented to the left (contralateral) visual field were affected selectively by anodal tDCS, whereas competing stimuli across the two visual fields were affected by both anodal and cathodal tDCS. These contrasting effects of anodal and cathodal tDCS on perception of single and competing stimuli suggest dissociable neural coding properties within the right PPC.

The role of executive attention in object substitution masking

It was long thought that a key characteristic of object substitution masking (OSM) was the requirement for spatial attention to be dispersed for the mask to impact visual sensitivity. However, recent studies have provided evidence questioning whether spatial attention interacts with OSM magnitude, suggesting that the previous reports reflect the impact of performance being at ceiling for the low attention load conditions. Another technique that has been employed to modulate attention in OSM paradigms involves presenting the target stimulus foveally, but with another demanding task shown immediately prior, and thus taxing executive/temporal attention. Under such conditions, when the two tasks occur in close temporal proximity relatively to greater temporal separation, masking is increased. However this effect could also be influenced by performance being at ceiling in some conditions. Here, we manipulated executive attention for a foveated target using a dual-task paradigm. Critically, ceiling performance was avoided by thresholding the target stimulus prior to it being presented under OSM conditions. We found no evidence for an interaction between executive attention load and masking. Collectively, along with the previous findings, our results provide compelling evidence that OSM as a phenomenon occurs independently of attention.

Anodal tDCS applied during multitasking training leads to transferable performance gains

Cognitive training can lead to performance improvements that are specific to the tasks trained. Recent research has suggested that transcranial direct current stimulation (tDCS) applied during training of a simple response-selection paradigm can broaden performance benefits to an untrained task. Here we assessed the impact of combined tDCS and training on multitasking, stimulus-response mapping specificity, response-inhibition, and spatial attention performance in a cohort of healthy adults. Participants trained over four days with concurrent tDCS – anodal, cathodal, or sham – applied to the left prefrontal cortex. Immediately prior to, 1 day after, and 2 weeks after training, performance was assessed on the trained multitasking paradigm, an untrained multitasking paradigm, a go/no-go inhibition task, and a visual search task. Training combined with anodal tDCS, compared with training plus cathodal or sham stimulation, enhanced performance for the untrained multitasking paradigm and visual search tasks. By contrast, there were no training benefits for the go/no-go task. Our findings demonstrate that anodal tDCS combined with multitasking training can extend to untrained multitasking paradigms as well as spatial attention, but with no extension to the domain of response inhibition.

Transcranial direct current stimulation of superior medial frontal cortex disrupts response selection during proactive response inhibition

Cognitive control is a vital executive process that is involved in selecting, generating, and maintaining appropriate, goal-directed behaviour. One operation that draws heavily on this resource is the mapping of sensory information to appropriate motor responses (i.e., response selection). Recently, a transcranial direct current stimulation (tDCS) study demonstrated that the left posterior lateral prefrontal cortex (pLPFC) is casually involved in response selection and response selection training. Correlational brain imaging evidence has also implicated the superior medial frontal cortex (SMFC) in response selection, and there is causal evidence that this brain region is involved in the proactive modulation of response tendencies when occasional stopping is required (response inhibition). However, to date there is only limited causal evidence that implicates the SMFC in response selection. Here, we investigated the role of SMFC in response selection, response selection training (Experiment 1) and response selection when occasional response inhibition is anticipated (Experiments 2 and 3) by employing anodal, cathodal, and sham tDCS. Cathodal stimulation of the SMFC modulated response selection by increasing reaction times in the context of proactive response inhibition. Our results suggest a context dependent role of the SMFC in response selection and hint that task set can influence the interaction between the brain and behaviour.

Dynamic, continuous multitasking training leads to task-specific improvements but does not transfer across action selection tasks

The ability to perform multiple tasks concurrently is an ever-increasing requirement in our information-rich world. Despite this, multitasking typically compromises performance due to the processing limitations associated with cognitive control and decision-making. While intensive dual-task training is known to improve multitasking performance, only limited evidence suggests that training-related performance benefits can transfer to untrained tasks that share overlapping processes. In the real world, however, coordinating and selecting several responses within close temporal proximity will often occur in high-interference environments. Over the last decade, there have been notable reports that training on video action games that require dynamic multitasking in a demanding environment can lead to transfer effects on aspects of cognition such as attention and working memory. Here, we asked whether continuous and dynamic multitasking training extends benefits to tasks that are theoretically related to the trained tasks. To examine this issue, we asked a group of participants to train on a combined continuous visuomotor tracking task and a perceptual discrimination task for six sessions, while an active control group practiced the component tasks in isolation. A battery of tests measuring response selection, response inhibition, and spatial attention was administered before and immediately after training to investigate transfer. Multitasking training resulted in substantial, task-specific gains in dual-task ability, but there was no evidence that these benefits generalized to other action control tasks. The findings suggest that training on a combined visuomotor tracking and discrimination task results in task-specific benefits but provides no additional value for untrained action selection tasks.Cognition: multitasking training boosts task-specific skills onlyTraining on a dynamic multitasking game leads to task-specific improvements but this performance gain does not transfer to novel tasks. Bender and colleagues from the University of Queensland, Australia, had one group of human participants train on a combined visuomotor tracking and shape discrimination task over several days, while another group trained on the two tasks in isolation (active control group). Following training, multitasking performance selectively improved for the multitasking group and not for the active control group. Critically, this training-related benefit did not generalize to a wide range of cognitive tasks that are theoretically linked to the current dual-task paradigm, indicating that repeated exposure to two concurrent tasks induces the learning of task-specific skills so that coordination between two specific tasks can be implemented more efficiently.