Department of Physiology, Feinberg School of Medicine, Northwestern University, Chicago, Illinois; and.
J Neurophysiol. 2013 Nov;110(10):2497-506. doi: 10.1152/jn.00177.2013. Epub 2013 Aug 21.
The propensity of a neuron to synchronize is captured by its infinitesimal phase response curve (iPRC). Determining whether an iPRC is biphasic, meaning that small depolarizing perturbations can actually delay the next spike, if delivered at appropriate phases, is a daunting experimental task because negative lobes in the iPRC (unlike positive ones) tend to be small and may be occluded by the normal discharge variability of a neuron. To circumvent this problem, iPRCs are commonly derived from numerical models of neurons. Here, we propose a novel and natural method to estimate the iPRC by direct estimation of its spectral modes. First, we show analytically that the spectral modes of the iPRC of an arbitrary oscillator are readily measured by applying weak harmonic perturbations. Next, applying this methodology to biophysical neuronal models, we show that a low-dimensional spectral reconstruction is sufficient to capture the structure of the iPRC. This structure was preserved reasonably well even with added physiological scale jitter in the neuronal models. To validate the methodology empirically, we applied it first to a low-noise electronic oscillator with a known design and then to cortical pyramidal neurons, recorded in whole cell configuration, that are known to possess a monophasic iPRC. Finally, using the methodology in conjunction with perforated-patch recordings from pallidal neurons, we show, in contrast to recent modeling studies, that these neurons have biphasic somatic iPRCs. Biphasic iPRCs would cause lateral somatically targeted pallidal inhibition to desynchronize pallidal neurons, providing a plausible explanation for their lack of synchrony in vivo.
神经元同步的倾向由其无穷小相位响应曲线(iPRC)捕获。确定 iPRC 是否为双相的,即小的去极化扰动如果在适当的相位下传递实际上可以延迟下一个尖峰,这是一项艰巨的实验任务,因为 iPRC 中的负瓣(与正瓣不同)往往很小,并且可能被神经元的正常放电变异性掩盖。为了规避此问题,通常从神经元的数值模型中推导出 iPRC。在这里,我们提出了一种新颖而自然的方法,通过直接估计其频谱模式来估计 iPRC。首先,我们从理论上证明,通过施加弱谐波扰动,很容易测量任意振荡器的 iPRC 的频谱模式。接下来,将此方法应用于生物物理神经元模型,我们表明,低维频谱重建足以捕获 iPRC 的结构。即使在神经元模型中添加了生理尺度抖动,该结构也能得到很好的保留。为了通过经验验证该方法,我们首先将其应用于具有已知设计的低噪声电子振荡器,然后将其应用于全细胞配置中记录的皮层锥体神经元,已知这些神经元具有单相 iPRC。最后,使用该方法与从苍白球神经元进行的穿孔贴片记录相结合,我们表明,与最近的建模研究相反,这些神经元具有双相的体相 iPRC。双相 iPRC 会导致侧向靶向苍白球的抑制使苍白球神经元去同步,为它们在体内缺乏同步性提供了合理的解释。