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| However, spike trains extracellularly recorded from single cortical neurons exhibit high variability. | | However, spike trains extracellularly recorded from single cortical neurons exhibit high variability. |
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− | 然而,从单个皮质神经元记录的胞外'''<font color="#ff8000"> 尖峰序列spike trains </font>'''表现出高度的<font color="#32CD32">可变性</font>。 | + | 然而,从单个皮质神经元记录的胞外'''<font color="#ff8000"> 尖峰序列spike trains </font>'''表现出高度的可变性。 |
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− | For instance, the coefficient of variation of the inter-spike intervals (ISIs) of neurons firing in response to a sensory input for a period of several seconds, is approximately equal to 1, as expected from a Poisson process (Softky and Koch, 1993). | + | For instance, '''<font color="#32CD32">the coefficient of variation</font>''' of the inter-spike intervals (ISIs) of neurons firing in response to a sensory input for a period of several seconds, is approximately equal to 1, as expected from a Poisson process (Softky and Koch, 1993). |
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− | 例如,响应于数秒内的感觉输入而放电的神经元的放电间隔(ISIs)的<font color="#32CD32">变化系数</font>大约等于1,正如'''<font color="#ff8000"> 泊松过程Poisson process </font>'''所预期的那样(Softky和Koch,1993)。 | + | 例如,响应于数秒内的感觉输入而放电的神经元的放电间隔(ISIs)的'''<font color="#32CD32">变化系数</font>'''大约等于1,正如'''<font color="#ff8000"> 泊松过程Poisson process </font>'''所预期的那样(Softky和Koch,1993)。 |
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| This apparent paradox between simple probabilistic considerations and the observed statistics of cortical spike trains led to several proposed resolutions. | | This apparent paradox between simple probabilistic considerations and the observed statistics of cortical spike trains led to several proposed resolutions. |
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| The behavior of the membrane potential is approximated using a passive, single compartment, conductance-based model of the neuron, described by | | The behavior of the membrane potential is approximated using a passive, single compartment, conductance-based model of the neuron, described by |
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− | 膜电位的行为是用神经元的被动单隔室的基于电导的模型近似的,描述如下
| + | 膜电位的行为是用神经元的被动单'''<font color="#ff8000">隔室compartment</font>'''的<ref>基于电导的模型</ref>近似的,描述如下 |
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| <math>CdV/dt = -G_{leak}(V(t)-E_{leak}) - G_{ex}(t)(V(t)-E_{ex}) - G_{in}(t)(V(t)-E_{in})+I_{inj}</math> (1) | | <math>CdV/dt = -G_{leak}(V(t)-E_{leak}) - G_{ex}(t)(V(t)-E_{ex}) - G_{in}(t)(V(t)-E_{in})+I_{inj}</math> (1) |
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| where \(E_{leak}\) is the resting membrane potential of the neuron, \(C\) is its capacitance, \(G_{leak}\) is the mean conductance in absence of stimulation (the inverse of input resistance), \(E_{ex}\) and \(E_{in}\) are the reversal potentials of excitation and inhibition, and \(I_{inj}\) is the current injected through the recording pipette. | | where \(E_{leak}\) is the resting membrane potential of the neuron, \(C\) is its capacitance, \(G_{leak}\) is the mean conductance in absence of stimulation (the inverse of input resistance), \(E_{ex}\) and \(E_{in}\) are the reversal potentials of excitation and inhibition, and \(I_{inj}\) is the current injected through the recording pipette. |
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− | 其中<math>E_{leak}</math>是神经元的静息膜电位,C是其电容,<math>G_{leak}</math>是在没有刺激的情况下的平均电导(输入电阻的倒数),<math>E_{ex}</math>和<math>E_{in}</math>是兴奋和抑制的反转电位,<math>I_{in}</math>是通过记录移液管注入的电流。 | + | 其中<math>E_{leak}</math>是神经元的静息膜电位,<math>C</math>是其电容,<math>G_{leak}</math>是在没有刺激的情况下的平均电导(输入电阻的倒数),<math>E_{ex}</math>和<math>E_{in}</math>是兴奋和抑制的反转电位,<math>I_{inj}</math>是通过记录移液管注入的电流。 |
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| By fitting equation (1) to the average responses at different holding potentials, the synaptic conductances evoked by the stimulus, \(G_{ex}(t)\) and \(G_{in}(t)\ ,\) can be computed (see Figure 1). | | By fitting equation (1) to the average responses at different holding potentials, the synaptic conductances evoked by the stimulus, \(G_{ex}(t)\) and \(G_{in}(t)\ ,\) can be computed (see Figure 1). |
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| An opposite scenario, where inhibitory inputs have narrower selectivity, was observed for frequency tuning in layer V '''intrinsically-bursting''' (but not regular-spiking) neurons of the primary auditory cortex (Sun et al., 2013). | | An opposite scenario, where inhibitory inputs have narrower selectivity, was observed for frequency tuning in layer V '''intrinsically-bursting''' (but not regular-spiking) neurons of the primary auditory cortex (Sun et al., 2013). |
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− | 在初级听觉皮层的第五层内在簇放电(但不是规则的尖峰)神经元中观察到了一种相反的情况,即抑制性输入的选择性较窄(Sun等人,2013年)。
| + | 在初级听觉皮层的第五层'''<font color="#ff8000">内在簇放电</font>'''(但不是规则的尖峰)神经元中观察到了一种相反的情况,即抑制性输入的选择性较窄(Sun等人,2013年)。 |
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| Also in the auditory cortex, some intensity-tuned neurons receive excitatory inputs which peak at the preferred intensity, whereas their inhibitory inputs increase monotonically with the stimulus strength (Wu et al., 2006), representing a case where the co-tuning of excitation and inhibition appears to break altogether. | | Also in the auditory cortex, some intensity-tuned neurons receive excitatory inputs which peak at the preferred intensity, whereas their inhibitory inputs increase monotonically with the stimulus strength (Wu et al., 2006), representing a case where the co-tuning of excitation and inhibition appears to break altogether. |
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| In the Up phase a barrage of synaptic inputs produces a reliable depolarization of 10-20 mV, which occasionally causes spiking (see Figure 1 in Up and down states). | | In the Up phase a barrage of synaptic inputs produces a reliable depolarization of 10-20 mV, which occasionally causes spiking (see Figure 1 in Up and down states). |
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− | 在上升阶段,大量突触输入会产生10-20 mV的可靠去极化,偶尔会导致尖峰([[上升和下降状态]]见图1)。 | + | 在上升阶段,大量突触输入会产生10-20 mV的可靠去极化,偶尔会导致尖峰(上升和下降状态见图1)。 |
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| The other end of the spectrum of brain states in awake mammals is the quiet wakefulness condition, which is somewhat similar to light anesthesia, and is characterized by rather short depolarizations ('bumps') and membrane potential distribution that is not bimodal, e.g., (DeWeese and Zador, 2006; Poulet and Petersen, 2008). | | The other end of the spectrum of brain states in awake mammals is the quiet wakefulness condition, which is somewhat similar to light anesthesia, and is characterized by rather short depolarizations ('bumps') and membrane potential distribution that is not bimodal, e.g., (DeWeese and Zador, 2006; Poulet and Petersen, 2008). |
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− | 清醒哺乳动物大脑状态谱的另一端是安静的清醒状态,这在某种程度上类似于轻度麻醉,其特征是相当短的去极化(“碰撞”)和非双峰的膜电位分布,例如(DeWeese和Zador,2006;Poulet和Petersen,2008)。
| + | 清醒哺乳动物大脑状态谱的另一端是安静的清醒状态,这在某种程度上类似于轻度麻醉,其特征是相当短的去极化('''<font color="#ff8000">“碰撞”</font>''')和非双峰的膜电位分布,例如(DeWeese和Zador,2006;Poulet和Petersen,2008)。 |
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− | In the quiet wakefulness condition and light state of anesthesia there are no stereotypic Up events nor does the activity resemble a single continuous Up phase, therefore the single-electrode conductance measurement method which requires averaging over multiple repeats of some stereotypic event, recorded at different holding potentials, cannot be applied. | + | In the quiet wakefulness condition and light state of anesthesia there are no '''<font color="#32CD32"> stereotypic</font>''' Up events nor does the activity resemble a single continuous Up phase, therefore the single-electrode conductance measurement method which requires averaging over multiple repeats of some stereotypic event, recorded at different holding potentials, cannot be applied. |
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− | 在安静的清醒状态和轻度的麻醉状态下,没有定型(??)的上升事件,也没有类似于单一连续上升阶段的活动,因此,无法应用单电极电导测量方法,该方法要求在不同的保持电位下记录的一些定型事件的多次重复中求平均值。
| + | 在安静的清醒状态和轻度的麻醉状态下,没有'''<font color="#32CD32">定型</font>'''的上升事件,也没有类似于单一连续上升阶段的活动,因此,无法应用单电极电导测量方法,该方法要求在不同的保持电位下记录的一些定型事件的多次重复中求平均值。 |
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| However, the substantial synchrony of synaptic inputs to closely located neurons (Lampl et al., 1999; Hasenstaub et al., 2005; Okun and Lampl, 2008; Poulet and Petersen, 2008) which exists in this case allows to continuously monitor both the excitatory and the inhibitory activity in the local network. | | However, the substantial synchrony of synaptic inputs to closely located neurons (Lampl et al., 1999; Hasenstaub et al., 2005; Okun and Lampl, 2008; Poulet and Petersen, 2008) which exists in this case allows to continuously monitor both the excitatory and the inhibitory activity in the local network. |
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| 为此,使用一对临近神经元的同步记录,其中一个细胞超极化,接近抑制的反转电位,另一个细胞去极化,足以接近兴奋的反转电位(Okun和Lampl,2008),图3。 | | 为此,使用一对临近神经元的同步记录,其中一个细胞超极化,接近抑制的反转电位,另一个细胞去极化,足以接近兴奋的反转电位(Okun和Lampl,2008),图3。 |
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− | This method reveals that in this type of spontaneous activity the excitatory and inhibitory inputs are interlocked in time, with inhibition lagging by several milliseconds behind excitation. | + | This method reveals that in this type of spontaneous activity the excitatory and inhibitory inputs are '''<font color="#32CD32">interlocked</font>''' in time, with inhibition lagging by several milliseconds behind excitation. |
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− | 该方法揭示了在这种自发活动中,兴奋性和抑制性输入在时间上是互锁(??)的,抑制滞后于兴奋数毫秒。
| + | 该方法揭示了在这种自发活动中,兴奋性和抑制性输入在时间上是'''<font color="#32CD32">互锁</font>'''的,抑制滞后于兴奋数毫秒。 |
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| Furthermore, the strength of excitatory and inhibitory inputs is (positively) correlated – large bumps typically contain both a strong excitatory and a strong inhibitory components, whereas small bumps are due to weak synaptic inputs, rather than strong inhibition that quenches the excitatory input. | | Furthermore, the strength of excitatory and inhibitory inputs is (positively) correlated – large bumps typically contain both a strong excitatory and a strong inhibitory components, whereas small bumps are due to weak synaptic inputs, rather than strong inhibition that quenches the excitatory input. |
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| 近年来,出现了一系列新的遗传工具,特别是对于小鼠(''Mus musculus'')物种。 | | 近年来,出现了一系列新的遗传工具,特别是对于小鼠(''Mus musculus'')物种。 |
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− | In addition, working with awake head-fixed mice is relatively straightforward. | + | In addition, working with awake head-fixed mice is relatively '''<font color="#32CD32">straightforward</font>'''. |
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− | 此外,使用清醒的头部固定的小鼠相对直截了当(??)。
| + | 此外,使用清醒的头部固定的小鼠相对'''<font color="#32CD32">直截了当</font>'''。 |
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| These and other recent developments are heavily relied upon in the current research which, in addition to the directions discussed in the previous sections, focuses on new aspects of E/I balance, as described in more detail below. | | These and other recent developments are heavily relied upon in the current research which, in addition to the directions discussed in the previous sections, focuses on new aspects of E/I balance, as described in more detail below. |