Jason D. Forte

Quantitative Review Finds No Evidence of Cognitive Effects in Healthy Populations From Single-session Transcranial Direct Current Stimulation (tDCS)

BACKGROUND Over the last 15-years, transcranial direct current stimulation (tDCS), a relatively novel form of neuromodulation, has seen a surge of popularity in both clinical and academic settings. Despite numerous claims suggesting that a single session of tDCS can modulate cognition in healthy adult populations (especially working memory and language production), the paradigms utilized and results reported in the literature are extremely variable. To address this, we conduct the largest quantitative review of the cognitive data to date. METHODS Single-session tDCS data in healthy adults (18-50) from every cognitive outcome measure reported by at least two different research groups in the literature was collected. Outcome measures were divided into 4 broad categories: executive function, language, memory, and miscellaneous. To account for the paradigmatic variability in the literature, we undertook a three-tier analysis system; each with less-stringent inclusion criteria than the prior. Standard mean difference values with 95% CIs were generated for included studies and pooled for each analysis. RESULTS Of the 59 analyses conducted, tDCS was found to not have a significant effect on any - regardless of inclusion laxity. This includes no effect on any working memory outcome or language production task. CONCLUSION Our quantitative review does not support the idea that tDCS generates a reliable effect on cognition in healthy adults. Reasons for and limitations of this finding are discussed. This work raises important questions regarding the efficacy of tDCS, state-dependency effects, and future directions for this tool in cognitive research.

Evidence that transcranial direct current stimulation (tDCS) generates little-to-no reliable neurophysiologic effect beyond MEP amplitude modulation in healthy human subjects: A systematic review

BACKGROUND Transcranial direct current stimulation (tDCS) is a form of neuromodulation that is increasingly being utilized to examine and modify a number of cognitive and behavioral measures. The theoretical mechanisms by which tDCS generates these changes are predicated upon a rather large neurophysiological literature. However, a robust systematic review of this neurophysiological data has not yet been undertaken. METHODS tDCS data in healthy adults (18-50) from every neurophysiological outcome measure reported by at least two different research groups in the literature was collected. When possible, data was pooled and quantitatively analyzed to assess significance. When pooling was not possible, data was qualitatively compared to assess reliability. RESULTS Of the 30 neurophysiological outcome measures reported by at least two different research groups, tDCS was found to have a reliable effect on only one: MEP amplitude. Interestingly, the magnitude of this effect has been significantly decreasing over the last 14 years. CONCLUSION Our systematic review does not support the idea that tDCS has a reliable neurophysiological effect beyond MEP amplitude modulation - though important limitations of this review (and conclusion) are discussed. This work raises questions concerning the mechanistic foundations and general efficacy of this device - the implications of which extend to the steadily increasing tDCS psychological literature.

Transcranial direct current stimulation: five important issues we aren't discussing (but probably should be)

Transcranial Direct Current Stimulation (tDCS) is a neuromodulatory device often publicized for its ability to enhance cognitive and behavioral performance. These enhancement claims, however, are predicated upon electrophysiological evidence and descriptions which are far from conclusive. In fact, a review of the literature reveals a number of important experimental and technical issues inherent with this device that are simply not being discussed in any meaningful manner. In this paper, we will consider five of these topics. The first, inter-subject variability, explores the extensive between- and within-group differences found within the tDCS literature and highlights the need to properly examine stimulatory response at the individual level. The second, intra-subject reliability, reviews the lack of data concerning tDCS response reliability over time and emphasizes the importance of this knowledge for appropriate stimulatory application. The third, sham stimulation and blinding, draws attention to the importance (yet relative lack) of proper control and blinding practices in the tDCS literature. The fourth, motor and cognitive interference, highlights the often overlooked body of research that suggests typical behaviors and cognitions undertaken during or following tDCS can impair or abolish the effects of stimulation. Finally, the fifth, electric current influences, underscores several largely ignored variables (such as hair thickness and electrode attachments methods) influential to tDCS electric current density and flow. Through this paper, we hope to increase awareness and start an ongoing dialog of these important issues which speak to the efficacy, reliability, and mechanistic foundations of tDCS.

Receptive field asymmetries produce color-dependent direction selectivity in primate lateral geniculate nucleus

Blue-on receptive fields recorded in primate retina and lateral geniculate nucleus are customarily described as showing overlapping blue-on and yellow-off receptive field components. However, the retinal pathways feeding the blue-on and yellow-off subfields arise from spatially discrete receptor populations, and recent studies have given contradictory accounts of receptive field structure of blue-on cells. Here we analyzed responses of blue-on cells to drifting gratings, in single-cell extracellular recordings from the dorsal lateral geniculate nucleus in marmosets. We show that most blue-on cells exhibit selectivity for the drift direction of achromatic gratings. The standard concentric difference-of-Gaussians (DOG) model thus cannot account for responses of these cells. We apply a simple, anatomically plausible, extension of the DOG model. The model incorporates temporally offset elliptical two-dimensional Gaussian subfields. The model can predict color-contingent direction and spatial tuning. Because direction tuning in blue-on cells depends on stimulus chromaticity, spatial frequency, and temporal frequency, this property is of little value as a general mechanism for image movement detection. It is possible that anatomical wiring for color selectivity has constrained the capacity of blue-on cells to contribute to spatial and motion vision.

Predicting Perceptual Decision Biases from Early Brain Activity

Perceptual decision making is believed to be driven by the accumulation of sensory evidence following stimulus encoding. More controversially, some studies report that neural activity preceding the stimulus also affects the decision process. We used a multivariate pattern classification approach for the analysis of the human electroencephalogram (EEG) to decode choice outcomes in a perceptual decision task from spatially and temporally distributed patterns of brain signals. When stimuli provided discriminative information, choice outcomes were predicted by neural activity following stimulus encoding; when stimuli provided no discriminative information, choice outcomes were predicted by neural activity preceding the stimulus. Moreover, in the absence of discriminative information, the recent choice history primed the choices on subsequent trials. A diffusion model fitted to the choice probabilities and response time distributions showed that the starting point of the evidence accumulation process was shifted toward the previous choice, consistent with the hypothesis that choice priming biases the accumulation process toward a decision boundary. This bias is reflected in prestimulus brain activity, which, in turn, becomes predictive of future decisions. Our results provide a model of how non-stimulus-driven decision making in humans could be accomplished on a neural level.

Spatial limitations of temporal segmentation

We investigated the spatial parameters that permit temporal phase segmentation. Subjects identified a stimulus quadrant which was modulated 180 degrees out of phase with the rest of the stimulus at temporal frequencies between 2 and 30 Hz. We determined the modulation sensitivity for regular square lattices of Gaussian spots and a stimulus made from solid quadrants with varying separation. Sensitivity declined rapidly when spatial separation of the modulating areas was approximately 0.4 degree, but was relatively unchanged by further spatial separations. The results suggest that there are two systems that can detect temporal phase differences. The first is a segregation process that operates below 10 Hz, where phase can be consciously followed and compared across large retinal distances. The second system is a segmentation mechanism that operates at higher temporal frequencies but only over a short range.

Summation of Visual Motion across Eye Movements Reflects a Nonspatial Decision Mechanism

Human vision remains perceptually stable even though retinal inputs change rapidly with each eye movement. Although the neural basis of visual stability remains unknown, a recent psychophysical study pointed to the existence of visual feature-representations anchored in environmental rather than retinal coordinates (e.g., “spatiotopic” receptive fields; Melcher and Morrone, 2003). In that study, sensitivity to a moving stimulus presented after a saccadic eye movement was enhanced when preceded by another moving stimulus at the same spatial location before the saccade. The finding is consistent with spatiotopic sensory integration, but it could also have arisen from a probabilistic improvement in performance due to the presence of more than one motion signal for the perceptual decision. Here we show that this statistical advantage accounts completely for summation effects in this task. We first demonstrate that measurements of summation are confounded by noise related to an observers uncertainty about motion onset times. When this uncertainty is minimized, comparable summation is observed regardless of whether two motion signals occupy the same or different locations in space, and whether they contain the same or opposite directions of motion. These results are incompatible with the tuning properties of motion-sensitive sensory neurons and provide no evidence for a spatiotopic representation of visual motion. Instead, summation in this context reflects a decision mechanism that uses abstract representations of sensory events to optimize choice behavior.

Onset Rivalry: The Initial Dominance Phase Is Independent Of Ongoing Perceptual Alternations

Binocular rivalry has been used to study a wide range of visual processes, from the integration of low-level features to the selection of signals that reach awareness. However, many of these studies do not distinguish between early and late phases of rivalry. There is clear evidence that the “onset” stage of rivalry is characterized by stable, yet idiosyncratic biases that are not evident in the average dominance of sustained rivalry viewing. Low-level stimulus features also have robust effects in the onset phase that are not seen in sustained rivalry, suggesting these phases may be driven at least partly by different neural mechanisms. The effects of high-level cognitive and affective factors at onset are less clear but also show differences from their effects in sustained viewing. These findings have important implications for the interpretation of any rivalry experiments using brief presentation paradigms and for understanding how the brain copes with binocular discrepancies in natural viewing conditions in which our eyes constantly move around an ever-changing environment. This review will summarize current research and explore the factors influencing this “onset” stage.

Spatial coding and response redundancy in parallel visual pathways of the marmoset Callithrix jacchus.

Many neurons in the primary visual cortex (area V1) show pronounced selectivity for the orientation and spatial frequency of visual stimuli, whereas most neurons in subcortical afferent streams show little selectivity for these stimulus attributes. It has been suggested that this transformation is a functional sign of increased coding efficiency, whereby the redundancy (or overlap in response properties) is reduced at consecutive levels of visual processing. Here we compared experimentally the response redundancy in area V1 with that in the three main dorsal thalamic afferent streams, the parvocellular (PC), koniocellular (KC), and magnocellular (MC) divisions of the dorsal lateral geniculate nucleus (LGN) in marmosets. The spatial frequency and orientation tuning of single cells in the LGN and area V1 were measured, using luminance contrast sine-wave gratings. A joint spatial frequency-orientation response selectivity profile was calculated for each cell. Response redundancy for each population was estimated by cross-multiplication of the joint selectivity profiles for pairs of cells. We show that when estimated in this way, redundancy in LGN neurons is approximately double that of neurons in cortical area V1. However, there are differences between LGN subdivisions, such that the KC pathway has a spatial representation that lies between the redundant code of the PC and MC pathways and the more efficient sparse spatial code of area V1.

Transmission of colour and acuity signals by parvocellular cells in marmoset monkeys

Non‐technical summary  Colour gets a free ride, according to our study of visual nerve cell responses in marmoset monkeys. All male marmosets are red–green colour‐blind (dichromatic), but most female marmosets have normal trichromatic colour vision. It is known that signals for high‐acuity daytime vision are carried in the parvocellular (P) pathway, and the P pathway also carries signals for red–green colour vision in trichromats. Here we compared P cell responses with patterned stimuli in dichromatic and trichromatic marmosets, and found no detectable difference in resolving power for fine patterns. These results indicate that red–green colour vision does not come at a cost for spatial vision. The ‘piggyback ride’ for colour signals in the P pathway may have encouraged the evolution of full colour vision in primates, including great apes, monkeys and humans.