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Spike Timing Dependent Plasticity of Inhibitory Synapses
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3306 J S HAAS T NOWOTNY AND H D I ABARBANEL, ticity We construct a one dimensional chain and a two dimen. sional layer model and show that the inhibitory plasticity leads. to efficient and flexible control of network activity in both. cases We hypothesize that the observed plasticity of inhibitory. synapses is a mechanism to control inappropriate run away. seizure like activity Our results provide strong evidence for. the importance of inhibitory inputs in maintaining an appro. priate balance of synaptic signaling in the brain, All experiments were conducted as approved by the UCSD IACUC. Young 14 to 21 day old Long Evans rats were killed by overexpo. sure to CO2 and decapitated The brain was quickly removed and. immersed in cold 0 C oxygenated artificial cerebral spinal fluid. Downloaded from http jn physiology org by 10 220 33 3 on October 9 2016. ACSF which contained in mM 126 NaCl 3 KCl 1 25 NaH2PO4. 2 MgSO4 26 NaHCO3 10 glucose and 2 CaCl2 buffered to pH 7 4. with 95 O2 5 CO2 Horizontal slices were prepared using a. Vibratome cutter TPI Slices were allowed to recover for 1 h prior. to recording in a holding chamber at room temperature continuously. bathed in oxygenated ACSF Slices were transferred to an immersion. chamber RC 27L Warner Instruments and visualized with IR DIC. optics Zeiss Axioskop 2FS Plus Dage CCD100 maintained at 34 C. TC 344B Warner Electrodes of resistance 4 6 M were pulled on FIG 1 Response of a typical layer II SC to long hyperpolarizing and. a horizontal puller Sutter Instruments and filled with a recording depolarizing current steps A In response to presynaptic stimulation within. solution in mM 135 KGluconate 4 KCl 2 NaCl 10 HEPES 0 2 layer II SCs show a compound response B left plot which can be pharma. cologically separated into an excitatory component B middle plot and an. EGTA 4 MgATP 0 3 GTP tris and 10 phosphocreatine tris Intra. inhibitory component B right plot, cellular signals were amplified Axoclamp 2B Molecular Devices. low pass filtered 8 pole Butterworth at 5 kHz and digitized at 10 RESULTS. kHz with a DAQ card NI PCI 6035E controlled by lab made. software created in LabView National Instruments In most exper Bidirectional STDP of inhibitory synapses. iments excitatory synaptic transmission was blocked by 6 cyano 7. nitroquinoxalene 2 3 dione CNQX 10 M and D amino 5 phos In control solution synaptic responses of SCs to intra layer. phonovaleric acid D APV 50 M obtained from Sigma St stimulations are a mix of excitatory and inhibitory responses. Louis MO In some experiments 10 mM bis o aminophenoxy Fig 1 The excitatory effects can be blocked by addition of. N N N N tetraacetic acid BAPTA Sigma was added to the internal the antagonists CNQX 10 M blocking AMPA receptors. solution and 15 M nimodipine Tocris from stock solution of 10. mM in DMSO was added to the bath, and D APV 50 M blocking NMDA receptors The. We obtained whole cell recordings from superficial EC layer II inhibitory responses can be blocked by addition of bicuculline. neurons We selected SCs by their superficial most position in layer II 10 M blocking GABAA receptors to the bath solution To. and oblong cell bodies as well as particular characteristics of their focus on the inhibitory portion of the response all recordings. electrophysiological responses to long current steps a prominent reported here were made in the presence of CNQX and D. 30 sag in response to both depolarizing and hyperpolarizing APV In each experiment sets of 30 baseline responses. current injections Alonso and Klink 1993 Haas and White 2002 as recorded at 0 5 Hz were monitored every 5 min over a period. well as an early first spike in response to suprathreshold stimuli Fig of 10 15 min to ensure a stable synaptic response. 1A From a total of 78 neurons the average rest potential was We paired presynaptic stimulations with single induced. 61 2 4 8 mV without correction for junction potential Neurons postsynaptic spikes Fig 2 arrow and varied the interval. were recorded in current clamp mode with no extra holding current. Presynaptic extracellular stimulation was delivered as 1 ms 10 to. between those stimuli t tpost tpre between 25 and 25. 50 A current pulses via 125 m concentric bipolar electrodes FHC ms Pairings were repeated at a rate of 2 Hz for 5 min After the. in layer II within 100 200 m of the recording electrode We paired pairings we monitored synaptic strength for up to an hour. synaptic responses to spikes by delivering extracellular stimulations recording sets of 30 postsynaptic responses at 0 5 Hz at 5 min. and forcing intracellular spikes using 1 ms 2 to 3 nA current injec intervals We quantified inhibitory postsynaptic potentials. tions through the recording electrode in current clamp mode at fixed IPSPs by their initial slopes the slope of a linear fit to the 1st. time delays We used 500 ms intervals between pairings for a total of 40 of the IPSP rise 11 6 3 5 ms and we normalized all. 3 5 min resulting in 320 600 pairings Baseline and postpairing responses to baseline We quantified the effective plasticity as. synaptic responses were collected as sets of 30 postsynaptic responses the mean IPSP slope between 20 and 30 min after pairings. to presynaptic stimulation collected at 0 5 Hz in 5 min intervals normalized to the mean of the slopes for 15 min preceding. Series and input resistances and resting potentials were monitored. throughout each experiment data from cells with variations 25 in. pairings We recorded IPSPs before and after pairings from a. those parameters were discarded Off line analysis was performed in total of 78 neurons IPSPs had initial sizes of 1 5 0 9 mV. Matlab Mathworks Numerical methods and modeling details are Examples of these results are shown in Fig 2 In the top. described in on line supplemental material Values are reported as panels we show the evolution of changes in IPSP initial slope. means SE statistical differences were measured with ANOVA after pairings with t 0 A and t 0 B Representative. unless indicated otherwise IPSPs for t 0 and t 0 are shown in the bottom panels. J Neurophysiol VOL 96 DECEMBER 2006 www jn org, SPIKE TIMING DEPENDENT PLASTICITY OF INHIBITORY SYNAPSES 3307.
experiments IPSPs were not significantly different after pair. ings in both of these cases n 4 P 0 2, In Fig 3A we show a summary plot of normalized change in. IPSP slope as a function of the pairing interval t For both. t 0 and t 0 significant changes in IPSPs were maximal. near t 10 ms and were restricted to relatively narrow. temporal windows The general trends potentiation depres. sion and temporal windows do not depend on the details of. evaluation using IPSP slope integral or amplitude to evaluate. the net change in synaptic strength yields the same overall. Pooled time course data are also shown in Fig 3B and show. on average that potentiation is expressed more slowly than. depression For responses that potentiated significantly P. 0 01 after pairings between 5 and 15 ms the first set of. IPSPs had a mean slope of 109 6 3 9 n 22 In, Downloaded from http jn physiology org by 10 220 33 3 on October 9 2016. comparison the first set of IPSPs that depressed to significant. levels P 0 01 had a mean slope of 82 2 7 7 n 15, Mechanisms of inhibitory STDP. To initiate the investigation of mechanisms for this bidirec. tional plasticity we repeated the pairings at delays of 10 and. FIG 2 After pairing presynaptic stimulations with evoked postsynaptic. spikes we monitored the evolution of changes in postsynaptic IPSPs We. observed long lasting potentiation of IPSPs for pairings with t tpost. tpre 0 A top individual and mean initial slopes for one cell in elapsed time. middle input resistance bottom each trace is the mean of 30 IPSPs traces are. shown over elapsed time We observed long lasting depression of IPSPs for. pairings with t tpost tpre 0 B top individual and mean initial slopes. for one cell in elapsed time middle input resistance bottom each trace is the. mean of 30 IPSPs traces are shown over elapsed time. We found that for pairings in which presynaptic stimulation. preceded postsynaptic stimulation t tpost tpre 0 IPSP. initial slopes potentiated This effect was maximal for delays. close to 10 ms and appears to be a very precise effect delays. of 5 or 20 ms were less effective in inducing potentiation. For all pairings with t between 5 and 15 ms IPSP slope. was enhanced on average to 134 3 5 9 n 26 P 0 02, of control values Potentiation tended to evolve slowly in time. after pairings, We found that for pairings in which t tpost tpre 0.
IPSP initial slopes were diminished As for potentiation this. effect is also very precise in its temporal requirements for. delays 5 or 15 ms no substantial effect was found For all. pairings with t between 15 and 5 ms IPSP slope was. diminished on average to 83 2 5 8 n 19 P 0 05 of, control values In contrast to potentiation depression was FIG 3 A summary results of change in postsynaptic IPSP initial slope. usually much faster and was usually expressed immediately IPSP expressed as a function of t tpost tpre No change corresponds to. following pairings normalized IPSP slope equal to unity equal to the baseline value before. pairing Each point represents data from one cell Change in slope is evaluated. In contrast to STDP of excitatory synpases for t near zero as the mean IPSP slope between 20 and 30 min following pairings normalized. we observed very little change in synaptic strength For all to the mean of the slopes for 15 min preceding pairings Data for which the. pairings with t between 5 and 5 ms IPSPs were on change in IPSP slope was significant P 0 01 ANOVA are plotted in open. average 103 6 3 3 n 11 P 0 3 of control values symbols The dashed line is a least square error fit used in modeling and is. Neither presynaptic stimulation alone nor postsynaptic spiking described in the text B pooled time series for potentiating circles and. depressing squares synapses evaluated for IPSP slopes showing significant. alone affected synaptic response at our stimulation rates As an P 0 01 potentiation following pairings between 5 to 15 ms and. experimental control we delivered isolated pre or postsynap significant P 0 01 depression following pairings between 15 to 5 ms. tic stimulation at the same interval and duration as in pairing Potentiation evolved more slowly than depression. J Neurophysiol VOL 96 DECEMBER 2006 www jn org, 3308 J S HAAS T NOWOTNY AND H D I ABARBANEL. 10 ms with 10 mM BAPTA added to the intracellular me. dium With intracellular calcium concentrations buffered no. significant potentiation was observed for pairings with t 0. 99 5 1 5 n 5 P 0 7 Potentiation but no significant. depression was observed for pairings with t 0 115 3. 3 3 n 5 P 0 01 Results of these experiments are, shown in Fig 4 along with representative IPSPs Our results. confirm previous reports Gaiarsa et al 2002 Woodin et al. 2003 that plasticity of inhibitory synapses is a process depen. dent on intracellular calcium dynamics, In our initial experiments we blocked both AMPA kainate. and NMDA receptors leaving voltage gated calcium channels. VGCCs as one possible site of calcium entry into the postsyn. aptic cell To investigate the role of VGCCs we repeated the. pairings at delays of 10 and 10 ms with 15 M nimodipine. to block L type calcium channels added to the bath solution. Downloaded from http jn physiology org by 10 220 33 3 on October 9 2016. Under this recording condition we saw no significant potenti. ation for pairings with t 0 101 89 1 9 n 5 P, 0 4 A small but insignificant depression was observed for.
pairings with t 0 95 1 3 9 n 4 P 0 2 Results, of these experiments are shown in Fig 5 along with represen. tative IPSPs As an experimental control we repeated pairings. in DMSO the solvent for nimodipine and as in control. conditions observed depression to 80 6 4 8 of control. n 5 P 001 for pairings with t 10 We observed, potentiation for t 0 to 136 0 8 3 of control n 3. P 0 01 FIG 5 With nimodipine in the bath solution we show that calcium enters. the postsynaptic cell through L type voltage gated calcium channels to partic. We also recorded paired pulse responses before and after ipate in the induction of inhibitory STDP We observed no significant change. IPSP spike pairings to investigate possible dependence of in IPSP slope after pairing for t 0 A top or for t 0 A bottom for. STDP on changes in presynaptic transmission In control this condition A representative set of IPSPs before and after pairings for each. temporal order of pairings is shown in B Summary data for n 9 cells some. data overlap completely is shown in C, conditions inhibitory synapses onto SCs exhibit paired pulse. depression measured as average of the second response di. vided by average of the first response Kim and Alger 2001. shown in Fig 6 Paired pulse ratios did not depend on IPSP. size for our dataset After induction of either LTP or LTD. paired pulse ratios were not significantly different from their. prepairing values n 10 P 0 4 providing one indication. that presynaptic release mechanisms remain unaffected by. IPSP spike pairings, Possible functions for inhibitory plasticity in the brain. To investigate functions of the observed plasticity rule we. first created a model of a unidirectional chain of 15 model SCs. each coupled reciprocally to a single model interneuron Fig. 7A A pool of background neurons firing with sinusoidally. Spike Timing Dependent Plasticity of Inhibitory Synapses in the Entorhinal Cortex Julie S Haas 1Thomas Nowotny and H D I Abarbanel1 2 1Institute for Nonlinear Science and 2Department of Physics and Marine Physical Laboratory Scripps Institution of Oceanography University of California San Diego La Jolla California

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