大鼠海马CA1区神经活动对体感刺激响应的研究
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摘要
感觉信号的处理一直是神经科学领域研究的热点,体感感觉因其遍布全身的感受器以及在触摸技术领域的广泛应用越来越受到重视。海马作为学习和记忆的中枢结构,从内嗅皮质接收有关外界体感刺激的信号,并将其按某种模式编码,以便信息在大脑中的传播和储存。早先的研究发现体感刺激可以使海马CA1区的多单元放电活动(multiple unit activity, MUA)发生变化,但其对CA1区具体的作用效果和机制并不明确。由于对体感信息的处理与动物的生存和活动息息相关,因此研究体感信息在海马区中的信号响应和处理机制具有重要意义。
本文通过对麻醉大鼠施加夹尾刺激,利用记录位点高密度分布的微电极阵列,记录海马CA1区的各种神经活动信号。主要研究了场电位、锋电位以及正向诱发电位对体感刺激的响应,考察了刺激对海马CA1区的作用效果；并且建立了包含体感输入和局部抑制回路的CA1区神经网络模型,利用计算机仿真验证了动物实验结果的产生机制；最后对海马中体感信息的处理与编码进行了探讨。
本文的主要研究结果如下：
(1)体感刺激对大鼠海马CA1区的主神经元活动具有抑制作用
在夹尾的体感刺激作用下,CA1区的场电位变成以θ节律为主要功率成分,同时伴随有不一致的MUA响应(发放减少或者不变)；其中,锥体细胞的锋电位发放频率减小,中间神经元的发放频率增加；锥体细胞爆发式发放频率减小,每个爆发式发放所包含的锋电位数量无明显变化；Schaffer侧支上施加正向脉冲刺激所诱发的CA1区群峰电位的幅值显著减小；CA1区的MUA、锥体细胞的发放对重复体感刺激产生了适应性现象(adaptation)。这些结果表明体感刺激对海马CA1区,尤其是锥体细胞兴奋性有明显的抑制作用。
(2)数学模型的仿真结果表明体感刺激可以增强海马CA1区局部抑制性回路的作用
利用NEURON软件建立CA1区神经网络模型,以研究海马对体感刺激响应的机制。模型的仿真结果与动物实验的结果一致,证明了体感刺激后CA1区锥体细胞兴奋性降低是中间神经元通过局部抑制回路作用的结果,其中前馈抑制回路和反馈抑制回路共同发挥作用,但以前馈抑制回路的作用为主导。为了进一步考察这种抑制作用是否能够用于控制异常的神经活动,我们还仿真了体感刺激对于过度兴奋状态下(癫痫发作时)锥体细胞兴奋性的影响,结果表明体感刺激可以减弱单侧颞叶癫痫活动,但无法减弱双侧颞叶癫痫活动。
(3)锋电位与场电位之间的相位锁定可能是CA1区编码体感信息的方式
本文利用Rayleigh检验分析了神经元锋电位与场电位θ节律之间的时间相关性,揭示了一种可能的CA1区对体感信息编码的方式。研究结果表明：夹尾刺激前,少部分神经元(3/22)展现出与0节律周期相位锁定的特性；夹尾刺激作用下,呈现相位锁定特性的神经元数量显著增加(10/22),其锋电位发放集中在0节律周期的负相位附近。
总之,本文结合了大鼠在体实验记录和建模仿真两种研究方法,考察了海马CA1区神经活动对体感刺激的响应,证明了体感刺激的抑制性作用,并且表明这种作用的内在机制是中间神经元通过局部抑制回路降低了锥体细胞的兴奋性。本文为海马是如何处理体感信息的研究提供了实验和理论依据,对海马的学习功能和神经编码的研究具有重要意义。
Processing of somatosensory signal has always been a hot spot in neuroscience. It receives more attention due to the numerous receptors spreading all over the body and the ever-growing applications in haptic technology. However, the basic mechanism of this process is largely undiscovered. As a well-known structure for learning and memory, hippocampus receives somatosensory input from entorhinal cortex, and encodes them to certain pattern for communication and storage within the brain. Previous studies found multiple unit activity (MUA) of the hippocampal CA1region responded to somatosensory stimuli, yet the explicit effect of the stimulation as well as the mechanism is not clear. Since the somatosensory system is crucial for animal's survival and daily life, it is of great significance to study the hippocampal response to somatosensory stimulation and the underlying mechanism.
In this dissertation, we investigated the hippocampal response to somatosensory input by applying tail-clamping on anesthetized rats and recorded the neural signals in CA1region with a microelectrode array. We analyzed the local field potential (LFP), unit spike, as well as the orthodromic population spike (OPS) evoked by Schaffer Collateral stimulation. Besides, a computer model with somatosensory input and local inhibitory circuits was built to explain the result in in-vivo experiment and the possible mechanism behind the observation. Lastly, we studied the mechanism of processing and encoding somatosensory information in the hippocampus.
The major findings are:
(1) Somatosensory stimulation could suppress the pricipal neurons' activities in hippocampal CA1region
We recorded hippocampal CA1neural activity before and during somatosensory stimulation, analyzing the response of LFP, unit spike and OPS. Results showed that during somatosensory stimulation:θ rhythm became the major power component of LFP, and mixed MUA responses (suppression or no change) appeared in CA1region; CA1pyramidal neurons showed firing decrease, while the interneurons showed firing increase; number of the burst of the pyramidal neurons decreased while the number of spikes within each burst did not show obvious change; the amplitude of OPS decreased significantly; the firing of MUA and pyramidal cells exhibited adaptation to the repetitive stimulation. These results demonstrated a strong suppressive effect on the hippocampal CA1neuronal activity, especially the excitability of pyramidal cells by somatosensory stimulation.
(2) Computer model showed the somatosensory stimulation could enhance the effect of local inhibitory circuits in hippocampal CA1region
We developed a NEURON model with somatosensory input and local inhibitory circuits, and the simulation result was in accordance to the experimental observation. Therefore it successfully demonstrated that the interneurons in the local circuits were responsible for the decreasing excitability of the pyramidal cells, and the suppressive effect by the somatosensory stimulation relied on both the feed-forward inhibitory circuit and the feedback inhibitory circuit, and the feed-forward inhibitory circuit plays a major role. Besides, we used an epileptic model to study the effect of somatosensory stimulation on over-excited neurons. Results showed that somatosensory stimulation could alleviate unilateral seizure-like activity, but not bilateral seizure-like activity.
(3) Somatosensory information could be encoded by the phase-locking relationship between spikes and LFP in the hippocampal CA1region
We analyzed the phasic relationship between unit spike and the θ cycles with Rayleigh test, showing that:a few neurons (3/22) exhibited phase-locking characteristic during spontaneous activity; tail-clamping stimulation increased the number of the phase-locking neurons significantly (10/22), and the spikes concentrated on the negative phase of the0cycle. These results suggested a possible mechanism of processing and encoding somatosensory information in the hippocampal CA1region.
In brief, this dissertation combined in-vivo experiment and computer modeling to study the neural response to the somatosensory stimulation in hippocampal CA1region. It demonstrated the suppressive effect of the somatosensory stimuli, and revealed the underlying mechanism. This study was significant for studying the processing of somatosensory information, it also help understand the neural encoding and learning function in the hippocampus.
本文通过对麻醉大鼠施加夹尾刺激,利用记录位点高密度分布的微电极阵列,记录海马CA1区的各种神经活动信号。主要研究了场电位、锋电位以及正向诱发电位对体感刺激的响应,考察了刺激对海马CA1区的作用效果；并且建立了包含体感输入和局部抑制回路的CA1区神经网络模型,利用计算机仿真验证了动物实验结果的产生机制；最后对海马中体感信息的处理与编码进行了探讨。
本文的主要研究结果如下：
(1)体感刺激对大鼠海马CA1区的主神经元活动具有抑制作用
在夹尾的体感刺激作用下,CA1区的场电位变成以θ节律为主要功率成分,同时伴随有不一致的MUA响应(发放减少或者不变)；其中,锥体细胞的锋电位发放频率减小,中间神经元的发放频率增加；锥体细胞爆发式发放频率减小,每个爆发式发放所包含的锋电位数量无明显变化；Schaffer侧支上施加正向脉冲刺激所诱发的CA1区群峰电位的幅值显著减小；CA1区的MUA、锥体细胞的发放对重复体感刺激产生了适应性现象(adaptation)。这些结果表明体感刺激对海马CA1区,尤其是锥体细胞兴奋性有明显的抑制作用。
(2)数学模型的仿真结果表明体感刺激可以增强海马CA1区局部抑制性回路的作用
利用NEURON软件建立CA1区神经网络模型,以研究海马对体感刺激响应的机制。模型的仿真结果与动物实验的结果一致,证明了体感刺激后CA1区锥体细胞兴奋性降低是中间神经元通过局部抑制回路作用的结果,其中前馈抑制回路和反馈抑制回路共同发挥作用,但以前馈抑制回路的作用为主导。为了进一步考察这种抑制作用是否能够用于控制异常的神经活动,我们还仿真了体感刺激对于过度兴奋状态下(癫痫发作时)锥体细胞兴奋性的影响,结果表明体感刺激可以减弱单侧颞叶癫痫活动,但无法减弱双侧颞叶癫痫活动。
(3)锋电位与场电位之间的相位锁定可能是CA1区编码体感信息的方式
本文利用Rayleigh检验分析了神经元锋电位与场电位θ节律之间的时间相关性,揭示了一种可能的CA1区对体感信息编码的方式。研究结果表明：夹尾刺激前,少部分神经元(3/22)展现出与0节律周期相位锁定的特性；夹尾刺激作用下,呈现相位锁定特性的神经元数量显著增加(10/22),其锋电位发放集中在0节律周期的负相位附近。
总之,本文结合了大鼠在体实验记录和建模仿真两种研究方法,考察了海马CA1区神经活动对体感刺激的响应,证明了体感刺激的抑制性作用,并且表明这种作用的内在机制是中间神经元通过局部抑制回路降低了锥体细胞的兴奋性。本文为海马是如何处理体感信息的研究提供了实验和理论依据,对海马的学习功能和神经编码的研究具有重要意义。
Processing of somatosensory signal has always been a hot spot in neuroscience. It receives more attention due to the numerous receptors spreading all over the body and the ever-growing applications in haptic technology. However, the basic mechanism of this process is largely undiscovered. As a well-known structure for learning and memory, hippocampus receives somatosensory input from entorhinal cortex, and encodes them to certain pattern for communication and storage within the brain. Previous studies found multiple unit activity (MUA) of the hippocampal CA1region responded to somatosensory stimuli, yet the explicit effect of the stimulation as well as the mechanism is not clear. Since the somatosensory system is crucial for animal's survival and daily life, it is of great significance to study the hippocampal response to somatosensory stimulation and the underlying mechanism.
In this dissertation, we investigated the hippocampal response to somatosensory input by applying tail-clamping on anesthetized rats and recorded the neural signals in CA1region with a microelectrode array. We analyzed the local field potential (LFP), unit spike, as well as the orthodromic population spike (OPS) evoked by Schaffer Collateral stimulation. Besides, a computer model with somatosensory input and local inhibitory circuits was built to explain the result in in-vivo experiment and the possible mechanism behind the observation. Lastly, we studied the mechanism of processing and encoding somatosensory information in the hippocampus.
The major findings are:
(1) Somatosensory stimulation could suppress the pricipal neurons' activities in hippocampal CA1region
We recorded hippocampal CA1neural activity before and during somatosensory stimulation, analyzing the response of LFP, unit spike and OPS. Results showed that during somatosensory stimulation:θ rhythm became the major power component of LFP, and mixed MUA responses (suppression or no change) appeared in CA1region; CA1pyramidal neurons showed firing decrease, while the interneurons showed firing increase; number of the burst of the pyramidal neurons decreased while the number of spikes within each burst did not show obvious change; the amplitude of OPS decreased significantly; the firing of MUA and pyramidal cells exhibited adaptation to the repetitive stimulation. These results demonstrated a strong suppressive effect on the hippocampal CA1neuronal activity, especially the excitability of pyramidal cells by somatosensory stimulation.
(2) Computer model showed the somatosensory stimulation could enhance the effect of local inhibitory circuits in hippocampal CA1region
We developed a NEURON model with somatosensory input and local inhibitory circuits, and the simulation result was in accordance to the experimental observation. Therefore it successfully demonstrated that the interneurons in the local circuits were responsible for the decreasing excitability of the pyramidal cells, and the suppressive effect by the somatosensory stimulation relied on both the feed-forward inhibitory circuit and the feedback inhibitory circuit, and the feed-forward inhibitory circuit plays a major role. Besides, we used an epileptic model to study the effect of somatosensory stimulation on over-excited neurons. Results showed that somatosensory stimulation could alleviate unilateral seizure-like activity, but not bilateral seizure-like activity.
(3) Somatosensory information could be encoded by the phase-locking relationship between spikes and LFP in the hippocampal CA1region
We analyzed the phasic relationship between unit spike and the θ cycles with Rayleigh test, showing that:a few neurons (3/22) exhibited phase-locking characteristic during spontaneous activity; tail-clamping stimulation increased the number of the phase-locking neurons significantly (10/22), and the spikes concentrated on the negative phase of the0cycle. These results suggested a possible mechanism of processing and encoding somatosensory information in the hippocampal CA1region.
In brief, this dissertation combined in-vivo experiment and computer modeling to study the neural response to the somatosensory stimulation in hippocampal CA1region. It demonstrated the suppressive effect of the somatosensory stimuli, and revealed the underlying mechanism. This study was significant for studying the processing of somatosensory information, it also help understand the neural encoding and learning function in the hippocampus.
引文
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