GABA及受体参与视觉发育可塑性关键期终止机制研究
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摘要
目的:本研究采用视皮层脑片膜片钳全细胞记录技术,研究P3w-8w正常饲养、视皮层chABC处理以及双眼形觉剥夺(binocular form deprivation,BFD)大鼠视皮层Ⅳ层神经元突触后电流(postsynaptic currents,PSCs)、GABA_A受体介导的抑制性突触后电流(inhibitory postsynaptic currents,IPSCs)特性以及突触传递长时程增强(Long-term potentiation,LTP)现象,并对以上各组大鼠视皮层Ⅳ层中间神经元进行形态学研究,探讨成年动物视皮层可塑性终止的细胞内机制,以及参与这一过程的神经递质及受体,为儿童眼病治疗及成年后视功能损伤病人的康复提供理论基础。
方法:P3w-8w Long-Evans大鼠,分为正常饲养、视皮层chABC处理以及BFD组,分别制备新鲜视皮层脑片,利用脑片膜片钳全细胞记录技术,记录视皮层Ⅳ层神经元PSCs,利用CNQX(20μM)和AP-5(50μM),分离GABA_A-IPSCs,并采用低频刺激伴随突触后去极化法诱发LTP。电生理记录的同时,进行biocytin细胞内标记,之后进行免疫组化染色。结合HE染色及电镜技术,对以上各组大鼠视皮层Ⅳ层中间神经元进行形态学研究。
结果:1.P3w～P8w正常饲养大鼠视皮层Ⅳ层神经元被动膜学特性及PSCs保持稳定,各周龄无显著差异;GABA_A-IPSCs峰值、IPSCs/PSCs随周龄增加而逐渐增大,下降时间逐渐变长,在出生后5、6周龄时达到高峰,之后保持成年稳定的高水平;2.与正常饲养大鼠比较,chABC处理组大鼠视皮层Ⅳ层神经元被动膜学特性及PSCs各项指标无显著变化,但GABA_A-IPSCs的峰值、IPSCs/PSCs以及下降时间显著低于同周龄正常饲养组;chABC酶处理组大鼠视皮层Ⅳ层LTP诱发率明显降低,且LTP幅度明显低于正常大鼠。3.BFD大鼠视皮层Ⅳ层神经元PSCs峰值、10～90%下降时间(Decay time)明显下降,GABA_A-IPSCs的峰值、IPSCs/PSCs以及下降时间都显著低于同周龄正常饲养组,BFD组与chABC处理组同周龄间比较,以上各指标相差显著。4.ChABC处理组大鼠视皮层Ⅳ层每平方毫米细胞记数、神经元胞体大小、树突数目以及超微结构与正常饲养大鼠无显著差异,BFD大鼠视皮层神经元形态幼稚,树突分枝及树突棘数量较少,突触超微结构发生明显病理改变。
Objective: Patch-clamp whole cell recording and transmission electron microscope techniques were applied to investigate the electrophysiological and morphological property of layer IVneurons from visual cortex of the rats under different growth condition at different postnatal days and to explore the synaptic and cellular mechanism by which GABA transmitter and recepter involve in the ending of the critical period of rat visual development. Investigation of visual development will help interpret the principle of brain working, and provide theoretical basis for the visual reconstruction of adult visual damage and the treatment of infant-eye-disease.Methods: The visual cortex slices were prepared from normal, chondroitin sulphate proteoglycans (CSPGs) degradation and binocular form deprivation(BFD) rats of both genders. CSPGs degradation rats were obtained by injecting chABC into rat's visual cortex. Patch-clamp whole cell recording techniques were adopted. Pre-synaptic test stimulation was given at 0.07 Hz intervals through bipolar stimulating electrodes placed in white matter. PSCs of layer IV neurons were recorded by recording electrodes. Then GABA_A-IPSCs were isolated by adding AP-5 and CNQX into artificial cerebrospinal fluid (ACSF). Long-term potentiation (LTP) was induced by pairing 1Hz low frequency presynaptic stimulation with postsynaptic depolarization. During recording, 0.5% biocytin were injected into the cells. Cells were stained after recording for histological assessments. HE staining and transmission electron microscope techniques were also used for morphological study.Results: 1. Input resistance (IR), resting membrane potential (RMP) and the parameters of PSCs are not changed, while the peak value, 10-90% decaytime of GABA-IPSCs and ratio of IPSCs/PSCs increased with age from P3w to P8w in normal rat visual cortex, then reached the top at P5,6w and kept high level in adult. 2. Treatment with chABC had no effect on PSCs, IR and RMP, while peak value, 10-90% decaytime of GABA-IPSCs and ratio of IPSCs/PSCs were significantly decreased by chABC treatment.
ChABC treatment also significantly decreased LTP incidence and amplitude of P35d rats. 3. BFD statistically decreased the peak value, 10-90% decaytime of PSCs and GABA-IPSCs and ratio of IPSCs/PSCs. There are also statistical differences of all the above parameters between BFD and chABC treatment group. 4. chABC had no effect on density of cells, cell body form, dentrite growth and ultrastructure, while BFD significantly changed those of layer IV interneurons in rat visual cortex.Summary: 1. Function of GABAa receptors of layer IV neurons in visual cortex is age dependent around the end of the critical period, reaching the top at P5,6w and keeping high level in adult. 2. The maturity of GABA circuits is involved in the end of critical period of visual development. The constant high level of GABAA receptors may result in the reduced plasticity in adult. 3. ChABC did not change PSCs, but changed GABAA-IPSCs through degradation of CSPGs, suggesting the maturity of GABA circuits is associated with the mechanism of inhibition of CSPGs for visual development plasticity. 4. CSPGs is suggested to be involved in LTP and long-term plasticity of layer IV in visual cortex. 5. Function of GABAa receptors of layer IV neuron in visual cortex is experience dependent. BFD affect inhibitory synaptic transmission through GABAa receptors. 6. Form information is crucial to the function development of visual cortical neurons. The kinetics basis of PSCs changes in BFD rats may lie in desensitization of postsynaptic receptors. 7. The changed morphological property and ultrastructure of layer IV intemeuron in visual cortex by BFD is the basis of the changed PSCs by BFD. 8. ChABC has no effect on morphological property and ultrastructure of layer IV intemeuron in visual cortex.
方法:P3w-8w Long-Evans大鼠,分为正常饲养、视皮层chABC处理以及BFD组,分别制备新鲜视皮层脑片,利用脑片膜片钳全细胞记录技术,记录视皮层Ⅳ层神经元PSCs,利用CNQX(20μM)和AP-5(50μM),分离GABA_A-IPSCs,并采用低频刺激伴随突触后去极化法诱发LTP。电生理记录的同时,进行biocytin细胞内标记,之后进行免疫组化染色。结合HE染色及电镜技术,对以上各组大鼠视皮层Ⅳ层中间神经元进行形态学研究。
结果:1.P3w～P8w正常饲养大鼠视皮层Ⅳ层神经元被动膜学特性及PSCs保持稳定,各周龄无显著差异;GABA_A-IPSCs峰值、IPSCs/PSCs随周龄增加而逐渐增大,下降时间逐渐变长,在出生后5、6周龄时达到高峰,之后保持成年稳定的高水平;2.与正常饲养大鼠比较,chABC处理组大鼠视皮层Ⅳ层神经元被动膜学特性及PSCs各项指标无显著变化,但GABA_A-IPSCs的峰值、IPSCs/PSCs以及下降时间显著低于同周龄正常饲养组;chABC酶处理组大鼠视皮层Ⅳ层LTP诱发率明显降低,且LTP幅度明显低于正常大鼠。3.BFD大鼠视皮层Ⅳ层神经元PSCs峰值、10～90%下降时间(Decay time)明显下降,GABA_A-IPSCs的峰值、IPSCs/PSCs以及下降时间都显著低于同周龄正常饲养组,BFD组与chABC处理组同周龄间比较,以上各指标相差显著。4.ChABC处理组大鼠视皮层Ⅳ层每平方毫米细胞记数、神经元胞体大小、树突数目以及超微结构与正常饲养大鼠无显著差异,BFD大鼠视皮层神经元形态幼稚,树突分枝及树突棘数量较少,突触超微结构发生明显病理改变。
Objective: Patch-clamp whole cell recording and transmission electron microscope techniques were applied to investigate the electrophysiological and morphological property of layer IVneurons from visual cortex of the rats under different growth condition at different postnatal days and to explore the synaptic and cellular mechanism by which GABA transmitter and recepter involve in the ending of the critical period of rat visual development. Investigation of visual development will help interpret the principle of brain working, and provide theoretical basis for the visual reconstruction of adult visual damage and the treatment of infant-eye-disease.Methods: The visual cortex slices were prepared from normal, chondroitin sulphate proteoglycans (CSPGs) degradation and binocular form deprivation(BFD) rats of both genders. CSPGs degradation rats were obtained by injecting chABC into rat's visual cortex. Patch-clamp whole cell recording techniques were adopted. Pre-synaptic test stimulation was given at 0.07 Hz intervals through bipolar stimulating electrodes placed in white matter. PSCs of layer IV neurons were recorded by recording electrodes. Then GABA_A-IPSCs were isolated by adding AP-5 and CNQX into artificial cerebrospinal fluid (ACSF). Long-term potentiation (LTP) was induced by pairing 1Hz low frequency presynaptic stimulation with postsynaptic depolarization. During recording, 0.5% biocytin were injected into the cells. Cells were stained after recording for histological assessments. HE staining and transmission electron microscope techniques were also used for morphological study.Results: 1. Input resistance (IR), resting membrane potential (RMP) and the parameters of PSCs are not changed, while the peak value, 10-90% decaytime of GABA-IPSCs and ratio of IPSCs/PSCs increased with age from P3w to P8w in normal rat visual cortex, then reached the top at P5,6w and kept high level in adult. 2. Treatment with chABC had no effect on PSCs, IR and RMP, while peak value, 10-90% decaytime of GABA-IPSCs and ratio of IPSCs/PSCs were significantly decreased by chABC treatment.
ChABC treatment also significantly decreased LTP incidence and amplitude of P35d rats. 3. BFD statistically decreased the peak value, 10-90% decaytime of PSCs and GABA-IPSCs and ratio of IPSCs/PSCs. There are also statistical differences of all the above parameters between BFD and chABC treatment group. 4. chABC had no effect on density of cells, cell body form, dentrite growth and ultrastructure, while BFD significantly changed those of layer IV interneurons in rat visual cortex.Summary: 1. Function of GABAa receptors of layer IV neurons in visual cortex is age dependent around the end of the critical period, reaching the top at P5,6w and keeping high level in adult. 2. The maturity of GABA circuits is involved in the end of critical period of visual development. The constant high level of GABAA receptors may result in the reduced plasticity in adult. 3. ChABC did not change PSCs, but changed GABAA-IPSCs through degradation of CSPGs, suggesting the maturity of GABA circuits is associated with the mechanism of inhibition of CSPGs for visual development plasticity. 4. CSPGs is suggested to be involved in LTP and long-term plasticity of layer IV in visual cortex. 5. Function of GABAa receptors of layer IV neuron in visual cortex is experience dependent. BFD affect inhibitory synaptic transmission through GABAa receptors. 6. Form information is crucial to the function development of visual cortical neurons. The kinetics basis of PSCs changes in BFD rats may lie in desensitization of postsynaptic receptors. 7. The changed morphological property and ultrastructure of layer IV intemeuron in visual cortex by BFD is the basis of the changed PSCs by BFD. 8. ChABC has no effect on morphological property and ultrastructure of layer IV intemeuron in visual cortex.
引文
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