哺乳动物嗅觉学习与记忆的结构基础研究
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摘要
嗅觉是一种最古老的感觉。在漫长的进化历史中,哺乳动物具备了非常惊人的气味感知以及学习和记忆能力。除了其基因组内拥有数量巨大的气味受体基因外,嗅觉系统本身具有复杂的神经连接回路,加上在鼻孔中的嗅上皮检测到气味信息后只需不超过三个突触即可连接到情感和认知中枢,以及各级嗅觉信息处理区域接受大量来自皮层神经反馈投射和脑干的调制神经输入有关。
神经回路的可塑性是学习和记忆的基础。突触可塑性是整个神经回路可塑性的核心。因而,揭示嗅觉学习与记忆的突触结构基础和相关的神经回路对于阐明学习记忆机制具有重要的基础意义。我们利用嗅觉缺失转基因小鼠模型,通过质粒电转标记结合兴奋性和抑制性突触抗体标记方法,发现嗅球中数量最大的中间神经元——颗粒细胞所接受的投射神经元的兴奋性输入和深层短轴突细胞的抑制性输入具有突触可塑性潜力,而接受的皮层离心兴奋性输入不具有突触可塑性潜力。因此,嗅觉的学习和记忆有可能只发生在颗粒细胞的抑制性突触和树树突触上。
此外,长期记忆的形成、巩固以及储存需要新的突触蛋白形成并参与构建新的突触连接。而CREB(cAMP反应元件结合蛋白)是调控突触可塑性和神经元自身可塑性的关键转录调控因子之一,通过检测CREB的磷酸化水平不仅可以判断与记忆相关的脑区,也能判断与记忆相关的特定脑区内的神经元类型,同时也能检测记忆在整个大脑水平上所形成的神经回路。因此,我们通过检测与嗅觉相关脑区的CREB磷酸化水平来研究参与小鼠气味与食物联系性学习和记忆的可能脑区的空间特征及其时间特性。结果发现CREB磷酸化水平在不同脑区不同时间内的动态具有显著的差别。因此,我们的结果为后期调控气味感知以及嗅觉的学习和记忆提供了可能的靶向基础。
The olfaction is a kind of most ancient sensory. In the long history of evolution, mammals have extraordinary and powerful ability of odor perception and olfactory learning and memory. Except the numerous odor receptor genes, there are many intricate neural circuits and no more than three synapses to the central areas related to the emotion and cognition, and each area processing olfactory information receives mass of centrifugal inputs from other regions and various modulate inputs from the brain stem.
Plasticity of neural circuits is the basis of learning and memory. Synaptic plasticity is the core of entire neural circuit plasticity. Thus, it is important to reveal the synaptic structural basis of olfactory learning and memory and related neural circuits for clarifying the learning and memory mechanisms. We use the methods of plasmid electroporation and excitatory and inhibitory synaptic protein labeling with adopting anosmia transgenic mice, and found that the excitatory synaptic inputs from projection neurons and inhibitory synaptic inputs from the deep short axon cells onto the granule cells, the most common interneurons in olfactory bulb, have the potential of structural plasticity, rather than the centrifugal inputs from the olfactory cortex. So, the inhibitory and dendrodendritic synapses of granule cells could be the sites of memory storage.
In addition, long-term memory formation, consolidation and storage need new protein synthesis to participate in the construction of new synaptic connections. CREB (cAMP response element-binding protein) is an important transcription factor to mediate synaptic plasticity and intrinsic plasticity. It is impossible to assess the regions and even the neuron types related to memory, but also the neural circuits in the overall brain through examining the changes of the pCREB. Therefore, we study the spacial and temporal characteristics of pCREB in major regions related to olfaction after odor-reward associative learning, and found that the dynamics of pCREB were very significantly different in different regions and at different time. The data we got supply the potential targeted basis to mediate odor perception and olfactory learning and memory.
神经回路的可塑性是学习和记忆的基础。突触可塑性是整个神经回路可塑性的核心。因而,揭示嗅觉学习与记忆的突触结构基础和相关的神经回路对于阐明学习记忆机制具有重要的基础意义。我们利用嗅觉缺失转基因小鼠模型,通过质粒电转标记结合兴奋性和抑制性突触抗体标记方法,发现嗅球中数量最大的中间神经元——颗粒细胞所接受的投射神经元的兴奋性输入和深层短轴突细胞的抑制性输入具有突触可塑性潜力,而接受的皮层离心兴奋性输入不具有突触可塑性潜力。因此,嗅觉的学习和记忆有可能只发生在颗粒细胞的抑制性突触和树树突触上。
此外,长期记忆的形成、巩固以及储存需要新的突触蛋白形成并参与构建新的突触连接。而CREB(cAMP反应元件结合蛋白)是调控突触可塑性和神经元自身可塑性的关键转录调控因子之一,通过检测CREB的磷酸化水平不仅可以判断与记忆相关的脑区,也能判断与记忆相关的特定脑区内的神经元类型,同时也能检测记忆在整个大脑水平上所形成的神经回路。因此,我们通过检测与嗅觉相关脑区的CREB磷酸化水平来研究参与小鼠气味与食物联系性学习和记忆的可能脑区的空间特征及其时间特性。结果发现CREB磷酸化水平在不同脑区不同时间内的动态具有显著的差别。因此,我们的结果为后期调控气味感知以及嗅觉的学习和记忆提供了可能的靶向基础。
The olfaction is a kind of most ancient sensory. In the long history of evolution, mammals have extraordinary and powerful ability of odor perception and olfactory learning and memory. Except the numerous odor receptor genes, there are many intricate neural circuits and no more than three synapses to the central areas related to the emotion and cognition, and each area processing olfactory information receives mass of centrifugal inputs from other regions and various modulate inputs from the brain stem.
Plasticity of neural circuits is the basis of learning and memory. Synaptic plasticity is the core of entire neural circuit plasticity. Thus, it is important to reveal the synaptic structural basis of olfactory learning and memory and related neural circuits for clarifying the learning and memory mechanisms. We use the methods of plasmid electroporation and excitatory and inhibitory synaptic protein labeling with adopting anosmia transgenic mice, and found that the excitatory synaptic inputs from projection neurons and inhibitory synaptic inputs from the deep short axon cells onto the granule cells, the most common interneurons in olfactory bulb, have the potential of structural plasticity, rather than the centrifugal inputs from the olfactory cortex. So, the inhibitory and dendrodendritic synapses of granule cells could be the sites of memory storage.
In addition, long-term memory formation, consolidation and storage need new protein synthesis to participate in the construction of new synaptic connections. CREB (cAMP response element-binding protein) is an important transcription factor to mediate synaptic plasticity and intrinsic plasticity. It is impossible to assess the regions and even the neuron types related to memory, but also the neural circuits in the overall brain through examining the changes of the pCREB. Therefore, we study the spacial and temporal characteristics of pCREB in major regions related to olfaction after odor-reward associative learning, and found that the dynamics of pCREB were very significantly different in different regions and at different time. The data we got supply the potential targeted basis to mediate odor perception and olfactory learning and memory.
引文
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