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Neuroscience

Optogenetic medrivning af hippocampus Theta svingninger i opfører sig mus

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

Vi beskriver brugen af optogenetics og elektrofysiologiske optagelser til selektiv manipulationer af hippocampus theta svingninger (5-10 Hz) i opfører mus. Effekten af rytme medrivning overvåges ved hjælp af lokale felt potentiale. En kombination af opto- og en hæmning adresser den efferente udlæsning af hippocampus synkronisering.

Abstract

Omfattende data om relationer af neurale netværk svingninger til adfærd og organisation af neuronal decharge på tværs af hjerneregioner kræver nye værktøjer til selektivt manipulere hjernen rytmer. Her beskriver vi en tilgang, der kombinerer projektion-specifikke optogenetics med ekstracellulære Elektrofysiologi til high-fidelity kontrol af hippocampus theta svingninger (5-10 Hz) i opfører mus. Optogenetic medrivning specificitet opnås ved at målrette channelrhodopsin-2 (ChR2) til GABAergic befolkningen af mediale septal celler, afgørende involveret i generation af hippocampus theta svingninger, og en lokal synkroniseret aktivering af en delmængde af hæmmende septal afferenter i hippocampus. Effekten af kontrolelementet optogenetic rytme er verificeret ved en samtidig overvågning af de lokale felt potentielle (LFP) på tværs af lamina af området CA1 og/eller neuronal decharge. Ved hjælp af denne let gennemførlige forberedelse viser vi effekten af forskellige optogenetic stimulation protokoller for induktion af theta svingninger og manipulation af deres hyppighed og regelmæssighed. Endelig, en kombination af kontrolelementet theta rytme med projektion-specifik hæmning adresser udlæsning af særlige aspekter af hippocampus synkroniseringen af efferente regioner.

Introduction

Neuronal aktivitet i pattedyr er koordineret af netværk svingninger, der bistår informationsoverførslen inden for og mellem hjernen regioner1,2,3,4. Hjernen rytmer omfatter svingninger spænder fra meget langsom ( 200 Hz) frekvenser. En stor mængde af beviser understøtter inddragelse af netværk svingninger i forskellige hjernefunktioner, herunder kognition5,6,7,8,9,10 , medfødte adfærd11,12 samt neuropsykiatriske lidelser som Parkinsons sygdom og epilepsi13,14,15. Selektiv og tidsligt præcise metoder til eksperimentelle manipulation af netværk svingninger er derfor afgørende for udviklingen af fysiologisk plausible modeller af synkronisering og for at etablere årsagssammenhænge med adfærd.

Netværkssynkronisering er medieret af forskellige biologiske substrater og processer, spænder fra molekylære identitet af ionkanaler og deres kinetik til Neuromodulationsbehandling af ophidselse og netværk connectivity. Den biologiske udformning af rytme generatorer16 er blevet afsløret for mange hjernen rytmer, forskellige aspekter afhvilke (, frequency, amplitude)er ofte forårsaget af dynamikken i forskellige celletyper og netværk. For eksempel, er hæmmende interneurons rettet mod somata af principal celler de vigtigste aktører på tværs af frekvensbånd og hjernen regioner17,18, herunder theta19,20, gamma20 , 21, og ripple (140-200 Hz)22 svingninger. Til gengæld er fase synkronisering af Fjern celler sikret af robust feed-forward signalering af pyramideformet celler, som nulstiller affyring af interneurons. En afgørende parameter for svingninger, den synkroniserede neuronal indbyggertal, er nært beslægtet med den målte LFP svingning amplitude og mindst for hurtige svingninger, afhænger det ophidsende drev på interneurons2. Derimod langsommere svingninger, som delta og theta rytmer, der genereres af langtrækkende tilbagevendende sløjfer, dannet af cortico-thalamic23,24 og hippocampus-mediale septal fremskrivninger25, 26,27, henholdsvis. Svingninger i sådanne kredsløb er tilvejebragt gennem interaktioner signal propagation forsinkelser, overgearet svar og deres hyppighed præference i deltagende celler28,29,30, 31 , 32. hæmmende fremskrivninger fra GABAergic parvalbumin (PV)-positive celler i de mediale septum (MS) til interneurons i hippocampus25,33, parahippocampal regioner og entorhinal cortex26 er afgørende for generation af theta svingninger i den mediale tindingelappen. Således kan fysiologiske mekanismer af netværk svingninger og neuronal synkronisering manipuleres ved hjælp af optogenetics med en real-time præcision.

Celle typespecifikke optogenetic manipulationer er blevet anvendt til undersøgelser af hippocampus og kortikale svingninger i vitro34,35,36,37,38 og in vivo30,39,40,41,42,43,44,45, herunder funktionelle undersøgelser af gamma5,12,36,46,47,48,49,50, 51,52 og ripple svingninger40,53,54 og søvn spindler55,56. For nylig udtrykte vi et Cre-afhængige ChR2 virus i MS, en vigtig region for generation af hippocampus theta rytme, af PV-Cre mus. Bruger dette præparat, blev funktioner af hippocampus theta svingninger (hyppighed og tidsmæssige stabilitet) kontrolleret af optogenetic stimulation af hæmmende fremskrivninger af MS i hippocampus11. Desuden vakte theta-frekvens optogenetic stimulation af hæmmende septo-hippocampus fremskrivninger theta rytme under vågen immobilitet. Optogenetically medrives theta rytme vises egenskaberne for spontan theta svingninger i musen på LFP og neuronal aktivitetsniveauer.

Nøglefunktioner af denne protokol omfatter: (1) udnyttelse af en hæmmende pathway, som er fysiologisk kritisk for spontan theta svingninger samtidig undgå uspecifik effekter på Hippocampus ophidselse; (2) cytoskeletale, dvs, projektion-specifik stimulation til at minimere en direkte indflydelse på ikke-hippocampus MS efferents; (3) lokale theta-rytmisk lys stimulation, at sikre en minimal direkte indblanding med theta-rytmisk septo-hippocampus dynamik og en global bilaterale medrivning af theta svingninger; (4) parametrisk kontrol af theta svingninger hyppighed og regelmæssighed; og (5) kvantificering af medrivning troskab med høj tidsmæssige opløsning ved hjælp af LFP aktivere kvantitative kausalitet analyse i opfører dyr. Da dette præparat udnytter hovedsagelig en kendt rollen som den septo-hippocampus disinhibition i theta generation25,30, det giver mulighed for robust kontrol over flere parametre af theta svingninger i opfører mus. Studier hvor andre mindre undersøgte veje og celletyper af septo-hippocampus kredsløb blev manipuleret38,39,47,49,50,51 , 52 , 53 , 54 , 55 , 56 , 57 , 58 afsløre yderligere mekanismer af theta rytme.

Protocol

PV-Cre knock-i mandlige mus59, 10-25 uger gammel, blev anvendt. Mus var opstaldet under standardbetingelser i det animalske facilitet og holdt på en 12-timers lys/mørke cyklus. Alle procedurer er udført i overensstemmelse med nationale og internationale retningslinjer, og blev godkendt af de lokale sundhedsmyndigheder (Landesamt für Natur, Umwelt und Verbraucherschutz, Nordrhein-Westfalen). 1. viral injektion Følg biologiske sikkerhed retningslinjer<sup c…

Representative Results

Målretning af ChR2 GABAergic celler i MS som beskrevet i afsnit 1 er illustreret i figur 2A. Optogenetic stimulation af axoner af MS GABAergic celler i den dorsale hippocampus via en optisk fiber, der er implanteret over området CA1 medriver theta svingninger på hyppigheden af stimulus i ipsilaterale (figur 2B) samt kontralaterale halvkugle (figur 2 c). Theta svingninger kan være mere eller mindr…

Discussion

Her fremlagde vi et almindeligt tilgængelige metode til entrain og fremkalde hippocampus theta svingninger i de opfører dyr. Denne tilgang kan være nyttig for studier af theta rytme funktioner i informationsbehandling og adfærd. Kritiske aspekter af denne metode omfatter: (1) valg af opsin og målretning af ChR2 til axoner af MS-celler i hippocampus, (2) robust optisk og elektriske funktioner af implanterede optisk fiber-wire array forsamlinger til at sikre kontinuerlig stimulering og LFP optagelse i opfører sig mus…

Disclosures

The authors have nothing to disclose.

Acknowledgements

Vi vil gerne takke Maria Gorbati ekspert hjælp til dataanalyse og Jennifer Kupferman for kommentarer på manuskriptet. Dette arbejde blev støttet af Deutsche Forschungsgemeinschaft (DFG; Exc 257 NeuroCure, TK og AP; Prioriterede Program 1665, 1799/1-1(2), Heisenberg program, 1799/2-1, AP), tysk-israelske grundlaget for videnskabelig forskning og udvikling (GIF; Jeg-1326-421.13/2015, TK) og Human Frontier Science Program (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
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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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