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神经科学

Optogenetic 在行为小鼠海马θ振荡中的作用

Published: June 29, 2018
doi:
10.3791/57349
, ,
1Systems Neurophysiology Research Group, Institute of Clinical Neuroscience and Medical Psychology, Medical Faculty,Heinrich Heine University Düsseldorf, 2Behavioural Neurodynamics Group,Leibniz Institute for Molecular Pharmacology (FMP)/ NeuroCure Cluster of Excellence, 3Neuronal Circuits and Behavior Research Group,Max Planck Institute for Metabolism Research

Summary

我们描述使用光遗传学和电生理记录的选择操作的海马θ振荡 (5-10 赫兹) 在行为小鼠。利用局部磁场电位监测节律夹带的效果。光和遗传药理学抑制的结合解决了海马同步的传出读数。

Abstract

广泛的数据, 神经网络振荡行为和组织神经元放电的大脑区域要求新的工具, 有选择地操纵大脑节律。在这里, 我们描述了一种方法, 结合投影特异光遗传学与细胞外电生理学的高保真度控制海马θ振荡 (5-10 Hz) 的行为小鼠。optogenetic 夹带的特异性是通过靶向 channelrhodopsin-2 (ChR2) 到内侧间隔细胞 GABAergic 的人群中实现的, 关键的是海马θ振荡的产生, 以及局部同步活化海马中抑制性间隔传入的子集。optogenetic 节律控制的功效通过同时监测 CA1 区和/或神经元放电的局部场电位 (LFP) 进行验证。利用这一易于实施的准备, 我们显示了各种 optogenetic 刺激协议的有效性, 以诱导θ振荡和操纵他们的频率和规律性。最后, 将θ节律控制与投影特异抑制相结合, 解决了传出区域海马同步的特定方面的读数。

Introduction

哺乳动物的神经元活动由网络振荡协调, 这有助于大脑区域内和脑区之间的信息传递1、2、3、4。大脑节奏包括从非常慢 ( 200 赫兹) 频率的振荡。大量的证据支持网络振荡参与不同的大脑功能, 包括认知5,6,7,8,9,10, 先天行为11,12以及精神疾病, 如帕金森病和癫痫13,14,15。因此, 对网络振荡进行实验性操作的选择性和世俗的精确方法, 对于发展生理上可行的同步模型和建立与行为的因果关系是至关重要的。

网络同步由不同的生物基质和过程介导, 从离子通道的分子识别和动力学到调节兴奋性和网络连通性。韵律发生器16的生物设计已经被揭示为许多大脑节律, 不同的方面 (例如, 频率, 振幅) 往往是由不同的细胞类型和网络的动态带来。例如, 抑制中间神经元靶向胞体的主要细胞是最重要的球员跨频段和脑区17,18, 包括θ19,20, 伽玛20,21, 波纹 (140-200 赫兹)22振荡。同时, 利用金字塔细胞的鲁棒前馈信号来保证远距离细胞的相位同步, 从而重置中间神经元的发射。振荡的一个关键参数, 即同步神经元种群的大小, 与测量的 LFP 振荡振幅密切相关, 至少对于快速振荡, 取决于兴奋性驱动到中间神经元2。相比之下, 较慢的振荡, 如三角洲和θ节律, 是由长距离折返循环产生的, 由皮质丘脑23,24和海马内侧间隔预测25组成, 分别为 26、27。在参与单元28、29、30中, 信号传播延迟、兴奋响应和频率偏好的相互作用带来了这种电路中的振荡,31,32. GABAergic parvalbumin (PV) 阳性细胞 (MS) 对海马25、33、海马旁区域和嗅皮质26的抑制性预测为中间神经元在颞叶中形成θ振荡的必要条件。因此, 利用光遗传学实时精度, 可以对网络振荡和神经元同步的生理机制进行操作。

细胞类型特异的 optogenetic 操作已被应用于研究海马和皮层振荡的体外34,35,36,37,38和在体内30,39,40,41,42,43,44,45, 包括功能调查伽玛5,12,36,46,47,48,49,50, 51,52和波纹振荡40,53,54和睡眠纺锤55,56。最近, 我们表达了一项依赖于 ChR2 病毒的 MS, 这是一个关键地区的海马θ节律的生成, 光伏的小鼠。利用该制剂, 海马θ振荡 (频率和时间稳定性) 的特征通过 optogenetic 刺激海马11中 MS 的抑制投影来控制。此外, θ频率 optogenetic 刺激抑制 septo-海马投射诱发θ节律在清醒静止。optogenetically 的θ节律显示了在 LFP 和神经元活动水平的小鼠自发θ振荡的性质。

该协议的主要特点包括: (1) 在生理上对自发θ振荡有重要影响的抑制通路的利用, 同时避免对海马兴奋性的不特异效应;(2) 轴突,即投射特异刺激, 以尽量减少对非海马 efferents MS 的直接影响;(3) 局部θ-韵律光刺激, 确保最小直接干扰的θ韵律 septo-海马动力学和全球双边夹带的θ振荡;(4) θ振荡频率和规律性的参数控制;(5) 利用 LFP 对具有高时间分辨率的夹带保真度进行量化, 以使行为动物的定量因果分析。由于这一准备基本上是利用 septo-海马抑制在θ25,30, 它的一个众所周知的作用, 它使得鲁棒控制的几个参数的θ振荡的行为小鼠。septo – 海马电路的其他研究较少的通路和细胞类型的研究纵了38,39,47,49,50,51,52,53,54,55,56,57,58揭示θ节奏的进一步机制。

Protocol

使用了59, 10-25 周大的光伏撞击雄性小鼠。老鼠被安置在动物设施的标准条件下, 并保持在12小时的光/暗循环。所有程序均按照国家和国际准则执行, 并经当地卫生当局批准 (Landesamt Natur、Umwelt 和 Verbraucherschutz、Nordrhein Westfalen)。 1. 病毒注射 在整个过程中, 遵循生物安全准则60。穿上实验室大衣、手术面罩、罩和两副手套。 用?…

Representative Results

如图 2A所示, ChR2 在 MS 中的靶向 GABAergic 细胞, 如第1节所述。Optogenetic GABAergic 细胞在背海马中的轴突通过一根被植入 CA1 区的光纤, 拽在同侧 (图 2B) 和对侧的刺激频率上的θ振荡。半球 (图 2C)。θ振荡可以或多或少有效的 optogenetic 刺激 (图 3A), 其效能, 计算为每个记录时代作为一个相对…

Discussion

在这里, 我们提出了一个广泛的方法, 以 entrain 和诱发海马θ振荡的行为动物。该方法对于研究θ节律在信息处理和行为中的作用具有一定的参考价值。该方法的关键方面包括: (1) 视蛋白的选择和靶向 ChR2 在海马中 MS 细胞的轴突, (2) 植入式光纤线阵列组件的鲁棒光学和电学特性, 以确保持续的刺激和 LFP记录在行为小鼠, (3) 应用最佳的光量在θ频率, (4) 后的夹带保真度的特殊量化, (5) 控制光电?…

Disclosures

The authors have nothing to disclose.

Acknowledgements

我们要感谢玛丽亚 Gorbati 的专家帮助与数据分析和珍妮弗 Kupferman 评论的手稿。这项工作得到德意志 Forschungsgemeinschaft (DFG) 的支持;激动 257 NeuroCure、TK 和 AP;优先方案1665、1799/1-1 (2)、海森堡方案、1799/2-1、AP)、德国-以色列科研开发基金会 (GIF;I-1326-421.13/2015, TK) 和人类前沿科学计划 (HFSP;RGY0076/2012, TK)。

Materials

PV-Cre mice The Jackson Laboratory B6;129P2-Pvalbtm1(cre)Arbr/J
Name Company Catalog Number Comments
Surgery
Stereotaxis David Kopf Instruments, Tujunga, CA, USA Model 963 Ultra Precise Small Animal Stereotaxic Instrument
Drill bits, 0.8 mm Bijoutil, Allschwil, Switzerland 49080HM
0.01-1 ml syringe Braun, Melsungen, Germany 9161406V
Sterican cannulas Braun 26 G, 0.45×25 mm BL/LB
Fine and sharp scissors Fine Science Tools Inc., Vancouver, Canada 14060-09
Forceps Fine Science Tools Inc. 11210-10 Dumont AA – Epoxy Coated Forceps
Blunt stainless steel scissors Fine Science Tools Inc. 14018-14
Soldering station Weller Tools GmbH, Besigheim, Germany WSD 81
Erythromycin Rotexmedica GmbH, Trittau, Germany PZN: 10823932 1g Powder for Solution for Infusion
Name Company Catalog Number Comments
Optogenetics
Hamilton pump PHD Ultra, Harvard Apparatus, Holliston, MA, USA model 703008 PHD Ultra Syringe Pump with push/pull mechanism
Hamilton 5 µL Syringe, 26 gauge PHD Ultra, Harvard Apparatus Model 75 RN SYR
Hamilton 5 µL Plunger PHD Ultra, Harvard Apparatus Model 75 RN SYR
Tubing Fisher Scientific, Pittsburgh, USA PE 20 Inner diameter 0.38 mm (.015"), Outer diameter 1.09 mm (.043")
Sterican cannulas Braun, Melsungen, Germany 27 G, 25×0.40 mm, blunt
Precision drill/grinder Proxxon, Wecker, Luxemburg fbs 240/e
Cutting disks Proxxon NO 28812
Cre dependent channelrhodopsin Penn Vector Core, Philadelphia, PA, USA AV-1-18917P Contruct name: AAV2/1.CAGGS.flex.ChR2.tdTomato, titer: 1.42×1013 vg/ml
Cam kinase dependent halorhodopsin Penn Vector Core AV-1-26971P Construct name: eNpHR3.0, AAV2/1.CamKIIa.eNpHR3.0-EYFP.WPRE.hGH, titer: 2.08_1012 vg/ml
Multimode optic fiber ThorLabs, Dachau, Germany FG105LCA 0.22 NA, Low-OH, Ø105 µm Core, 400 – 2400 nm
Ceramic stick ferrule Precision Fiber Products, Milpitas, CA, USA CFLC126 Ceramic LC MM Ferrule, ID 126um
Polishing paper Thorlabs LF3D 6" x 6" Diamond Lapping (Polishing) Sheet
Power meter Thorlabs PM100D Compact Power and Energy Meter Console, Digital 4" LCD
Multimode fiber optic coupler Thorlabs FCMM50-50A-FC 1×2 MM Coupler, 50:50 Split Ratio, 50 µm GI Fibers, FC/PC
Fiberoptic patch cord Thorlabs FG105LCA CUSTOM-MUC custom made, 3 m long, with protective tubing, Tubing: FT030, Connector 1: FC/PC, Connector 2: 1.25mm (LC) Ceramic Ferrule
Sleeve Precision Fiber Products, Milpitas, CA, USA ADAL1 Ceramic Split Mating Sleeve for Ø1.25 mm (LC/PC) Ferrules
473 nm DPSS laser Laserglow Technologies, Toronto, ON, Canada R471005FX LRS-0473 Series
593 nm DPSS laser Laserglow Technologies R591005FX LRS-0594 Series
MC_Stimulus II Multichannel Systems, Reutlingen, Germany STG 4004
Impedance conditioning module Neural microTargeting worldwide, Bowdoin, USA ICM
Name Company Catalog Number Comments
Electrophysiology
Tungsten wires California Fine Wire Company, Grover Beach, CA, USA CFW0010954 40 µm, 99.95%
Capillary tubing Optronics 1068150020 ID: 100.4 µm
Omnetics nanoconnector Omnetics Connector Corporation, Minneapolis, USA A79038-001
Screws Bilaney, Düsseldorf, Germany 00-96×1/16 stainless-steel
Silicone probe NeuroNexus Technologies, Ann Arbor, MI, USA B32
Headstage Neuralynx, Bozeman, Montana USA HS-8 miniature headstage unity gain preamplifiers
Silver conductive paint Conrad electronics, Germany 530042
Liquid flux Felder GMBH Löttechnik, Oberhausen, Germany Lötöl ST DIN EN 29454.1, 3.2.2.A (F-SW 11)
LED Neuralynx HS-LED-Red-omni-10V
Name Company Catalog Number Comments
Software
MATLAB Mathworks, Natick, MA, USA
MC_Stimulus software Multichannel, Systems
Neurophysiological Data Manager NDManager, http://neurosuite.sourceforge.net
Klusters http://neurosuite.sourceforge.net, Hazan et al., 2006
Software of the recording system Neuralynx Cheetah https://neuralynx.com/software/cheetah
Multi-channel data analysis software Cambridge Electronic Design Limited, Cambridge, GB Spike2
DOWNLOAD MATERIALS LIST

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Optogenetic EntrainmentHippocampal Theta OscillationsBehaving MiceNeurophysiologySpatial NavigationVirus InjectionOptic FiberTungsten Wire Array

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Bender, F., Korotkova, T., Ponomarenko, A. Optogenetic Entrainment of Hippocampal Theta Oscillations in Behaving Mice. J. Vis. Exp. (136), e57349, doi:10.3791/57349 (2018).
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