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== Intracellular measurement of the excitatory and inhibitory synaptic inputs 兴奋性和抑制性突触输入的细胞内测量 ==
== Intracellular measurement of the excitatory and inhibitory synaptic inputs 兴奋性和抑制性突触输入的细胞内测量 ==
[[Image:Balance_of_inhibition_and_excitation_figure1.png|thumb|400px|right|F1|Computation of synaptic conductance evoked by sensory stimulus. The average response to whisker deflection in a spiny stellate neuron in layer IV of the rat primary somatosensory cortex is recorded in current-clamp mode while injecting 4 different currents (left panel). In addition, neuron’s capacitance and leak conductance are measured (not shown). By fitting the responses to equation (1) the average excitatory and inhibitory synaptic conductances evoked by the stimulus are recovered (right panel). Adapted from (Heiss et al., 2008).
图1：由感觉刺激引起的突触电导的计算。
在注入4种不同的电流时（左图），以电流钳模式记录大鼠初级躯体感觉皮层的第IV层中，多刺星状神经元对晶须偏转（？）的平均响应。
此外，测量了神经元电容和漏电导（未显示）。
通过将响应拟合至方程（1），恢复了由刺激引起的平均兴奋性和抑制性突触电导（右图）（句意？？）。改编自（Heiss等人，2008）。
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In a pioneering study, Borg-Graham and colleagues used intracellular recordings to directly estimate the synaptic conductance changes evoked in cortical neurons by visual stimulation (Borg-Graham et al., 1996, 1998).
In a pioneering study, Borg-Graham and colleagues used intracellular recordings to directly estimate the synaptic conductance changes evoked in cortical neurons by visual stimulation (Borg-Graham et al., 1996, 1998).
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[[Image:Balance_of_inhibition_and_excitation_figure2vb.jpg|frame|right|F2|An example of a neuron in the auditory cortex with frequency and intensity co-tuned excitatory and inhibitory inputs. (a) Excitatory and inhibitory synaptic conductances evoked by stimuli of different frequencies and preferred intensity have a similar tuning. The measured conductances are shown at the bottom (green – excitatory conductance, red - inhibitory conductance, black – total conductance). (b) The excitatory and inhibitory inputs are also intensity co-tuned, notation as in (a). Adapted from (Wehr and Zador, 2003).
图2：听觉皮层中神经元的示例，其频率和强度共同调谐兴奋性和抑制性输入。
（a）由不同频率和偏好强度的刺激诱发的兴奋性和抑制性突触传导具有相似的调谐。
测量的电导显示在底部（绿色-兴奋性电导，红色-抑制性电导，黑色-总电导）。
（b） 兴奋性和抑制性输入也会由强度协同调谐，如（b）所示。改编自（Wehr和Zador，2003）
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The similar tuning of excitatory and inhibitory inputs to different features of the stimuli space appears to be a rather common organizational principle in the sensory areas, however there are several notable exceptions.
The similar tuning of excitatory and inhibitory inputs to different features of the stimuli space appears to be a rather common organizational principle in the sensory areas, however there are several notable exceptions.
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[[Image:Balance_of_inhibition_and_excitation_figure3.jpg|thumb|500px|right|F3|Excitatory and inhibitory inputs are synchronized during spontaneous activity. Two nearby neurons are simultaneously recorded when (a) both are at their resting potential, close to the reversal potential of inhibition (hyperpolarized-hyperpolarized mode); (b) both neurons are depolarized close to the reversal potential of excitation (depolarized-depolarized mode); (c-d) one of the neurons is in the hyperpolarized mode while the other is in the depolarized mode. In (a) the activity is dominated by excitatory inputs, which are seen to be highly synchronized between the neurons. Similarly, in (b) the activity is dominated by inhibitory inputs which are also highly synchronized. Finally, the mixed mode recordings (c-d) demonstrate that the excitatory and inhibitory inputs possess a high degree of synchrony. Adapted from (Okun and Lampl, 2008).
图3：在自发活动期间，兴奋性和抑制性输入是同步的。
在以下情景同时记录两个邻近的神经元
（a） 两者都处于静息电位，接近抑制反转电位（超极化-超极化模式）；
（b） 两个神经元都去极化，接近于兴奋的反转电位（去极化-去极化模式）；
（c-d）其中一个神经元处于超极化模式，而另一个处于去极化模式。
在（a）中，活动主要由兴奋性输入支配，在神经元之间将这种兴奋性输入视为高度同步的。
类似地，在（b）中，活动由抑制性输入支配，这些输入也是高度同步的。
最后，对混合模式的记录（c-d）表明兴奋性和抑制性输入具有高度的同步性。改编自（Okun and Lampl，2008）。
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