Department of Physiology and Neuroscience Program, Biomedicine Discovery Institute, Monash University, Clayton, VIC 3800, Australia.
RIKEN Center for Brain Science, RIKEN Institute, Wako-shi, 351-0198, Japan.
Brain. 2024 Nov 4;147(11):3751-3763. doi: 10.1093/brain/awae273.
Transcranial direct current stimulation (tDCS) has garnered significant interest for its potential to enhance cognitive functions and as a therapeutic intervention in various cognitive disorders. However, the clinical application of tDCS has been hampered by significant variability in its cognitive outcomes. Furthermore, the widespread use of tDCS has raised concerns regarding its safety and efficacy, particularly in light of our limited understanding of its underlying neural mechanisms at the cellular level. We still do not know 'where', 'when' and 'how' tDCS modulates information encoding by neurons, in order to lead to the observed changes in cognitive functions. Without elucidating these fundamental unknowns, the root causes of its outcome variability and long-term safety remain elusive, challenging the effective application of tDCS in clinical settings. Addressing this gap, our study investigates the effects of tDCS, applied over the dorsolateral prefrontal cortex, on cognitive abilities and individual neuron activity in macaque monkeys performing cognitive tasks. Like humans performing a delayed match-to-sample task, monkeys exhibited practice-related slowing in their responses (within-session behavioural adaptation). Concurrently, there were practice-related changes in simultaneously recorded activity of prefrontal neurons (within-session neuronal adaptation). Anodal tDCS attenuated both these behavioural and neuronal adaptations when compared with sham stimulation. Furthermore, tDCS abolished the correlation between response time of monkeys and neuronal firing rate. At a single-cell level, we also found that following tDCS, neuronal firing rate was more likely to exhibit task-specific modulation than after sham stimulation. These tDCS-induced changes in both behaviour and neuronal activity persisted even after the end of tDCS stimulation. Importantly, multiple applications of tDCS did not alter burst-like firing rates of individual neurons when compared with sham stimulation. This suggests that tDCS modulates neural activity without enhancing susceptibility to epileptiform activity, confirming a potential for safe use in clinical settings. Our research contributes unprecedented insights into the 'where', 'when' and 'how' of tDCS effects on neuronal activity and cognitive functions by showing that modulation of the behaviour of monkeys by the tDCS of the prefrontal cortex is accompanied by alterations in prefrontal cortical cell activity ('where') during distinct trial phases ('when'). Importantly, tDCS led to task-specific and state-dependent alterations in prefrontal cell activities ('how'). Our findings suggest a significant shift from the view that the effects of tDCS are merely attributable to polarity-specific shifts in cortical excitability and instead propose a more complex mechanism of action for tDCS that encompasses various aspects of cortical neuronal activity without increasing burst-like epileptiform susceptibility.
经颅直流电刺激 (tDCS) 因其增强认知功能的潜力以及作为各种认知障碍的治疗干预手段而受到广泛关注。然而,tDCS 的临床应用受到其认知结果显著变异性的阻碍。此外,tDCS 的广泛应用引发了对其安全性和有效性的担忧,特别是考虑到我们对其在细胞水平下的潜在神经机制的了解有限。我们仍然不知道 tDCS 如何通过神经元来调节信息编码,从而导致观察到的认知功能变化。在没有阐明这些基本未知因素的情况下,其结果变异性和长期安全性的根本原因仍然难以捉摸,这使得 tDCS 在临床环境中的有效应用具有挑战性。为了解决这一差距,我们的研究调查了经颅直流电刺激(应用于背外侧前额叶皮层)对执行认知任务的猕猴的认知能力和单个神经元活动的影响。就像人类执行延迟匹配样本任务一样,猕猴在反应中表现出与练习相关的减慢(会话内行为适应)。同时,在前额叶神经元的同时记录活动中也观察到与练习相关的变化(会话内神经元适应)。与假刺激相比,阳极 tDCS 减弱了这两种行为和神经元适应。此外,tDCS 消除了猴子的反应时间与神经元放电率之间的相关性。在单细胞水平上,我们还发现,tDCS 后神经元放电率更有可能表现出特定于任务的调制,而不是假刺激后。即使在 tDCS 刺激结束后,这些 tDCS 引起的行为和神经元活动的变化仍然持续存在。重要的是,与假刺激相比,多次 tDCS 应用不会改变单个神经元的爆发样放电率。这表明 tDCS 调节神经活动而不会增强癫痫样活动的易感性,从而确认了在临床环境中安全使用的潜力。我们的研究通过显示经颅直流电刺激对前额叶皮层的调制通过改变前额叶皮层细胞活动(“哪里”)在不同试验阶段(“何时”)来调节猴子的行为,为 tDCS 对神经元活动和认知功能的“哪里”、“何时”和“如何”影响提供了前所未有的见解。重要的是,tDCS 导致了前额叶细胞活动的任务特异性和状态依赖性改变(“如何”)。我们的研究结果表明,tDCS 的作用机制从仅仅归因于皮质兴奋性的极性特异性变化发生了重大转变,而是提出了一种更复杂的 tDCS 作用机制,包含了皮质神经元活动的各个方面,而不会增加爆发样癫痫样易感性。