Department of Neurology and Stroke and Hertie Institute for Clinical Brain Research, Eberhard Karls University of Tübingen, 72076 Tübingen, Germany,
Institute for Medical Psychology and Behavioral Neurobiology, Eberhard Karls University of Tübingen, 72076 Tübingen, Germany, and.
J Neurosci. 2019 Dec 11;39(50):10034-10043. doi: 10.1523/JNEUROSCI.1730-19.2019. Epub 2019 Nov 4.
Alpha oscillations (8-14 Hz) are assumed to gate information flow in the brain by means of pulsed inhibition; that is, the phasic suppression of cortical excitability and information processing once per alpha cycle, resulting in stronger net suppression for larger alpha amplitudes due to the assumed amplitude asymmetry of the oscillation. While there is evidence for this hypothesis regarding occipital alpha oscillations, it is less clear for the central sensorimotor μ-alpha rhythm. Probing corticospinal excitability via transcranial magnetic stimulation (TMS) of the primary motor cortex and the measurement of motor evoked potentials (MEPs), we have previously demonstrated that corticospinal excitability is modulated by both amplitude and phase of the sensorimotor μ-alpha rhythm. However, the direction of this modulation, its proposed asymmetry, and its underlying mechanisms remained unclear. We therefore used real-time EEG-triggered single- and paired-pulse TMS in healthy humans of both sexes to assess corticospinal excitability and GABA-A-receptor mediated short-latency intracortical inhibition (SICI) at rest during spontaneous high amplitude μ-alpha waves at different phase angles (peaks, troughs, rising and falling flanks) and compared them to periods of low amplitude (desynchronized) μ-alpha. MEP amplitude was facilitated during troughs and rising flanks, but no phasic suppression was observed at any time, nor any modulation of SICI. These results are best compatible with sensorimotor μ-alpha reflecting asymmetric pulsed facilitation but not pulsed inhibition of motor cortical excitability. The asymmetric excitability with respect to rising and falling flanks of the μ-alpha cycle further reveals that voltage differences alone cannot explain the impact of phase. The pulsed inhibition hypothesis, which assumes that alpha oscillations actively inhibit neuronal processing in a phasic manner, is highly influential and has substantially shaped our understanding of these oscillations. However, some of its basic assumptions, in particular its asymmetry and inhibitory nature, have rarely been tested directly. Here, we explicitly investigated the asymmetry of modulation and its direction for the human sensorimotor μ-alpha rhythm. We found clear evidence of pulsed facilitation, but not inhibition, in the human motor cortex, challenging the generalizability of the pulsed inhibition hypothesis and advising caution when interpreting sensorimotor μ-alpha changes in the sensorimotor system. This study also demonstrates how specific assumptions about the neurophysiological underpinnings of cortical oscillations can be experimentally tested noninvasively in humans.
阿尔法振荡(8-14Hz)被认为通过脉冲抑制来调节大脑中的信息流;也就是说,皮质兴奋性和信息处理每隔一个阿尔法周期进行一次相位抑制,从而导致更大的阿尔法幅度产生更强的净抑制,因为振荡的幅度不对称假设。虽然有证据支持这种假设,即关于枕叶阿尔法振荡,但对于中央感觉运动μ-阿尔法节律,情况就不那么清楚了。通过对初级运动皮层进行经颅磁刺激(TMS)和测量运动诱发电位(MEP)来探测皮质脊髓兴奋性,我们之前已经证明,皮质脊髓兴奋性受到感觉运动μ-阿尔法节律的幅度和相位的调节。然而,这种调节的方向、它的拟议的不对称性及其潜在机制仍不清楚。因此,我们在健康的男女被试中使用实时 EEG 触发的单脉冲和双脉冲 TMS,评估皮质脊髓兴奋性和 GABA-A 受体介导的短潜伏期皮质内抑制(SICI),在不同相位角(波峰、波谷、上升和下降边缘)的自发高幅度μ-阿尔法波期间处于休息状态,并将其与低幅度(去同步)μ-阿尔法波期间进行比较。MEP 幅度在波谷和上升边缘时得到促进,但在任何时候都没有观察到相位抑制,也没有 SICI 的调制。这些结果与感觉运动μ-阿尔法反射出不对称脉冲促进但不是运动皮质兴奋性的脉冲抑制的假设最相符。相对于μ-阿尔法周期的上升和下降边缘的不对称兴奋性进一步表明,仅电压差异不能解释相位的影响。脉冲抑制假说假设α 振荡以相位方式主动抑制神经元处理,具有高度影响力,并极大地影响了我们对这些振荡的理解。然而,其一些基本假设,特别是其不对称性和抑制性,很少被直接测试。在这里,我们明确地研究了人类感觉运动μ-阿尔法节律的调制的不对称性及其方向。我们在人类运动皮层中发现了明显的脉冲促进证据,但没有抑制证据,这对脉冲抑制假说的普遍性提出了挑战,并建议在解释感觉运动系统中的感觉运动μ-阿尔法变化时要谨慎。这项研究还展示了如何在人类中非侵入性地实验测试关于皮质振荡神经生理学基础的具体假设。
Brain Stimul. 2018-6-19
IEEE Trans Biomed Eng. 2025-7-16
NPJ Parkinsons Dis. 2025-4-5
J Neuroeng Rehabil. 2025-3-27
Front Syst Neurosci. 2024-12-4
Brain Stimul. 2019-7-28
Cereb Cortex. 2019-8-14