Interneuronal mechanisms of hippocampal theta oscillations in a full-scale model of the rodent CA1 circuit

  1. Marianne J Bezaire  Is a corresponding author
  2. Ivan Raikov
  3. Kelly Burk
  4. Dhrumil Vyas
  5. Ivan Soltesz
  1. University of California, Irvine, United States
  2. Stanford University, United States
35 figures and 74 tables

Figures

Figure 1 with 2 supplements
CA1 network connectivity.

(A) The model network is arranged in a layered prism with the lengths of each dimension similar to the actual dimensions of the CA1 region and its layers. (B) The model cell somata within a small chunk of stratum pyramidale (as depicted in A) are plotted to show the regular distribution of model cells throughout the layer in which they are found. (C) Each pyramidal cell in the network has detailed morphology with realistic incoming synapse placement along the dendrites and soma. (D,E) Diagrams illustrate connectivity between types of cells. (D) The network includes one principal cell type (pyramidal cells) and eight interneuron types. Cell types that may connect are linked by a line colored according to the presynaptic cell type. Most cell types can connect to most other cell types. Total number of cells of each type are displayed, as are the number of local output synapses (boutons) from all cells of each type. (E) The number, position, and cell types of each connection are biologically constrained, as are the numbers and positions of the cells. See Figure 1—figure supplement 1) for details about the convergence onto each cell type. Also see Table 1 and Figure 1—figure supplement 2 for information about the cell-type combinations of the 5 billion connections and the axonal distributions followed by each cell type, as well as detailed connectivity results at http://doi.org/10.6080/K05H7D60.

https://doi.org/10.7554/eLife.18566.002
Figure 1—figure supplement 1
Quantitative network connectivity.

The average number of incoming synapses per postsynaptic cell of the given type are shown for (A) all inputs to the cells, (B) all excitatory inputs to the cells and (C) all inhibitory inputs to the cells.

https://doi.org/10.7554/eLife.18566.003
Figure 1—figure supplement 2
Anatomically constrained connectivity.

The axonal distributions are shown per presynaptic cell type. The distribution of boutons is plotted as a function of distance from the presynaptic cell’s soma. Boutons connecting to all possible types of postsynaptic cells are included in the plot. The colors correspond to each presynaptic cell type using the same color code as previous figures.

https://doi.org/10.7554/eLife.18566.004
Electrophysiology of the model network components.

(A) Ion channel densities vary as a function of location (top) in the morphologically detailed pyramidal cell model (bottom; adapted from Poolos et al., 2002). Scale bar: 100 μm and 0.01 μF/cm2. (B–C) The sodium channel found in the pyramidal cell soma is characterized in terms of (B) the activation/inactivation curves and (C) the current-voltage relation at peak (transient) current and steady state. (DG) Current sweeps are shown for four model cell types: (D) PV+ basket cell, (E) CCK+ basket cell, (F) O-LM cell, and (G) neurogliaform cell. Scale bar: 100 ms and 20 mV. (H–J) Electrophysiological properties for each cell type, including (H) input resistance, (I) membrane time constant, and (J) action potential threshold. (K–L) Pyramidal cell synaptic connections are characterized as post-synaptic currents with the postsynaptic cell voltage clamped at −50 mV; (K) synapses made onto the pyramidal cell from all other cell types and (L) synapses made by the pyramidal cell onto all network cell types. Cells represented by same colors as in Figure 1. Source Data available for electrophysiological characterizations shown here. Additional details available in the Methods, Table 3, and the Appendix.

https://doi.org/10.7554/eLife.18566.005
Figure 2—source data 1

Model sodium channel activation.

The ion channel characterized in this figure was an Nav channel, inserted into a single compartment cell of diameter and length 16.8 microns (a soma) with a density such that the maximum, macroscopic conductance was 0.001 μS/cm2. The reversal potential of the channel was + 55 mV and the settings during the characterization protocol were: temperature = 34 degrees Celsius, axial resistance = 210 ohm*cm, [Ca2+]internal=5.0000e-06 mM, specific membrane capacitance = 1 μF/cm2. For activation steps, the cell was held at −120 mV and then stepped to potential levels ranging from −90 mV to + 90 mV. For inactivation steps, the cell was held at various potential levels ranging from −90 mV to + 40 mV for 500 ms and then stepped to + 20 mV. Each current injection step is recorded in three columns, where t: time (ms), i: injection (nA), g: conductance (S/cm2). The column labels are followed by the voltage (hold or step, according to the file), with activation steps being recorded in the Na_Channel_Step.dat and inactivation steps being recorded in the Na_Channel_Hold.dat file.

https://doi.org/10.7554/eLife.18566.006
Figure 2—source data 2

Model sodium channel inactivation.

The ion channel characterized in this figure was an Nav channel, inserted into a single compartment cell of diameter and length 16.8 microns (a soma) with a density such that the maximum, macroscopic conductance was .001 μS/cm2. The reversal potential of the channel was + 55 mV and the settings during the characterization protocol were: temperature = 34 degrees Celsius, axial resistance = 210 ohm*cm, [Ca2+]internal=5.0000e-06 mM, specific membrane capacitance = 1 μF/cm2. For activation steps, the cell was held at −120 mV and then stepped to potential levels ranging from −90 mV to +90 mV. For inactivation steps, the cell was held at various potential levels ranging from −90 mV to +40 mV for 500 ms and then stepped to +20 mV. Each current injection step is recorded in three columns, where t: time (ms), i: injection (nA), g: conductance (S/cm2). The column labels are followed by the voltage (hold or step, according to the file), with activation steps being recorded in the Na_Channel_Step.dat and inactivation steps being recorded in the Na_Channel_Hold.dat file.

https://doi.org/10.7554/eLife.18566.007
Figure 2—source data 3

Model axo-axonic cell current injection sweep.

Simulated current injection sweep in AxoClamp ATF (tab-delimited) file format.

https://doi.org/10.7554/eLife.18566.008
Figure 2—source data 4

Model bistratified cell current injection sweep.

Simulated current injection sweep in AxoClamp ATF (tab-delimited) file format.

https://doi.org/10.7554/eLife.18566.009
Figure 2—source data 5

Model CCK+ basket cell current injection sweep.

Simulated current injection sweep in AxoClamp ATF (tab-delimited) file format.

https://doi.org/10.7554/eLife.18566.010
Figure 2—source data 6

Model ivy cell current injection sweep.

Simulated current injection sweep in AxoClamp ATF (tab-delimited) file format.

https://doi.org/10.7554/eLife.18566.011
Figure 2—source data 7

Model neurogliaform cell current injection sweep.

Simulated current injection sweep in AxoClamp ATF (tab-delimited) file format.

https://doi.org/10.7554/eLife.18566.012
Figure 2—source data 8

Model O-LM cell current injection sweep.

Simulated current injection sweep in AxoClamp ATF (tab-delimited) file format.

https://doi.org/10.7554/eLife.18566.013
Figure 2—source data 9

Model PV+ basket cell current injection sweep.

Simulated current injection sweep in AxoClamp ATF (tab-delimited) file format.

https://doi.org/10.7554/eLife.18566.014
Figure 2—source data 10

Model pyramidal cell current injection sweep.

Simulated current injection sweep in AxoClamp ATF (tab-delimited) file format.

https://doi.org/10.7554/eLife.18566.015
Figure 2—source data 11

Model Schaffer Collateral-Associated cell current injection sweep.

Simulated current injection sweep in AxoClamp ATF (tab-delimited) file format.

https://doi.org/10.7554/eLife.18566.016
Figure 2—source data 12

Model paired recording of an Axo-axonic cell to Pyramidal cell connection.

Simulated paired recordings where the postsynaptic cell was voltage-clamped at −50 mV and the reversal potential of the synapse was kept at its physiological potential, as defined in the network model code. Sodium channels in the postsynaptic cell were blocked to prevent a suprathreshold response. A spike was triggered in the presynaptic cell and the current response was measured in the postsynaptic cell at the soma. This recording was repeated 10 times, with a randomly chosen connection location (from anatomically likely locations) each time. Each of the 10 trials are included in this file.

https://doi.org/10.7554/eLife.18566.017
Figure 2—source data 13

Model paired recording of a Bistratified cell to Pyramidal cell connection.

Simulated paired recordings where the postsynaptic cell was voltage-clamped at −50 mV and the reversal potential of the synapse was kept at its physiological potential, as defined in the network model code. Sodium channels in the postsynaptic cell were blocked to prevent a suprathreshold response. A spike was triggered in the presynaptic cell and the current response was measured in the postsynaptic cell at the soma. This recording was repeated 10 times, with a randomly chosen connection location (from anatomically likely locations) each time. Each of the 10 trials are included in this file.

https://doi.org/10.7554/eLife.18566.018
Figure 2—source data 14

Model paired recording of a CA3 cell to Pyramidal cell connection.

Simulated paired recordings where the postsynaptic cell was voltage-clamped at −50 mV and the reversal potential of the synapse was kept at its physiological potential, as defined in the network model code. Sodium channels in the postsynaptic cell were blocked to prevent a suprathreshold response. A spike was triggered in the presynaptic cell and the current response was measured in the postsynaptic cell at the soma. This recording was repeated 10 times, with a randomly chosen connection location (from anatomically likely locations) each time. Each of the 10 trials are included in this file.

https://doi.org/10.7554/eLife.18566.019
Figure 2—source data 15

Model paired recording of a CCK+ basket cell to Pyramidal cell connection.

Simulated paired recordings where the postsynaptic cell was voltage-clamped at −50 mV and the reversal potential of the synapse was kept at its physiological potential, as defined in the network model code. Sodium channels in the postsynaptic cell were blocked to prevent a suprathreshold response. A spike was triggered in the presynaptic cell and the current response was measured in the postsynaptic cell at the soma. This recording was repeated 10 times, with a randomly chosen connection location (from anatomically likely locations) each time. Each of the 10 trials are included in this file.

https://doi.org/10.7554/eLife.18566.020
Figure 2—source data 16

Model paired recording of an ECIII cell to Pyramidal cell connection.

Simulated paired recordings where the postsynaptic cell was voltage-clamped at −50 mV and the reversal potential of the synapse was kept at its physiological potential, as defined in the network model code. Sodium channels in the postsynaptic cell were blocked to prevent a suprathreshold response. A spike was triggered in the presynaptic cell and the current response was measured in the postsynaptic cell at the soma. This recording was repeated 10 times, with a randomly chosen connection location (from anatomically likely locations) each time. Each of the 10 trials are included in this file.

https://doi.org/10.7554/eLife.18566.021
Figure 2—source data 17

Model paired recording of an Ivy cell to Pyramidal cell connection.

Simulated paired recordings where the postsynaptic cell was voltage-clamped at −50 mV and the reversal potential of the synapse was kept at its physiological potential, as defined in the network model code. Sodium channels in the postsynaptic cell were blocked to prevent a suprathreshold response. A spike was triggered in the presynaptic cell and the current response was measured in the postsynaptic cell at the soma. This recording was repeated 10 times, with a randomly chosen connection location (from anatomically likely locations) each time. Each of the 10 trials are included in this file.

https://doi.org/10.7554/eLife.18566.022
Figure 2—source data 18

Model paired recording of a Pyramidal cell to Pyramidal cell connection.

Simulated paired recordings where the postsynaptic cell was voltage-clamped at −50 mV and the reversal potential of the synapse was kept at its physiological potential, as defined in the network model code. Sodium channels in the postsynaptic cell were blocked to prevent a suprathreshold response. A spike was triggered in the presynaptic cell and the current response was measured in the postsynaptic cell at the soma. This recording was repeated 10 times, with a randomly chosen connection location (from anatomically likely locations) each time. Each of the 10 trials are included in this file.

https://doi.org/10.7554/eLife.18566.023
Figure 2—source data 19

Model paired recording of a Neurogliaform cell to Pyramidal cell connection.

Simulated paired recordings where the postsynaptic cell was voltage-clamped at −50 mV and the reversal potential of the synapse was kept at its physiological potential, as defined in the network model code. Sodium channels in the postsynaptic cell were blocked to prevent a suprathreshold response. A spike was triggered in the presynaptic cell and the current response was measured in the postsynaptic cell at the soma. This recording was repeated 10 times, with a randomly chosen connection location (from anatomically likely locations) each time. Each of the 10 trials are included in this file.

https://doi.org/10.7554/eLife.18566.024
Figure 2—source data 20

Model paired recording of an O-LM cell to Pyramidal cell connection.

Simulated paired recordings where the postsynaptic cell was voltage-clamped at −50 mV and the reversal potential of the synapse was kept at its physiological potential, as defined in the network model code. Sodium channels in the postsynaptic cell were blocked to prevent a suprathreshold response. A spike was triggered in the presynaptic cell and the current response was measured in the postsynaptic cell at the soma. This recording was repeated 10 times, with a randomly chosen connection location (from anatomically likely locations) each time. Each of the 10 trials are included in this file.

https://doi.org/10.7554/eLife.18566.025
Figure 2—source data 21

Model paired recording of a PV+ basket cell to Pyramidal cell connection.

Simulated paired recordings where the postsynaptic cell was voltage-clamped at −50 mV and the reversal potential of the synapse was kept at its physiological potential, as defined in the network model code. Sodium channels in the postsynaptic cell were blocked to prevent a suprathreshold response. A spike was triggered in the presynaptic cell and the current response was measured in the postsynaptic cell at the soma. This recording was repeated 10 times, with a randomly chosen connection location (from anatomically likely locations) each time. Each of the 10 trials are included in this file.

https://doi.org/10.7554/eLife.18566.026
Figure 2—source data 22

Model paired recording of a Pyramidal cell to Axo-axonic cell connection.

Simulated paired recordings where the postsynaptic cell was voltage-clamped at −50 mV and the reversal potential of the synapse was kept at its physiological potential, as defined in the network model code. Sodium channels in the postsynaptic cell were blocked to prevent a suprathreshold response. A spike was triggered in the presynaptic cell and the current response was measured in the postsynaptic cell at the soma. This recording was repeated 10 times, with a randomly chosen connection location (from anatomically likely locations) each time. Each of the 10 trials are included in this file.

https://doi.org/10.7554/eLife.18566.027
Figure 2—source data 23

Model paired recording of a Pyramidal cell to Bistratified cell connection.

Simulated paired recordings where the postsynaptic cell was voltage-clamped at −50 mV and the reversal potential of the synapse was kept at its physiological potential, as defined in the network model code. Sodium channels in the postsynaptic cell were blocked to prevent a suprathreshold response. A spike was triggered in the presynaptic cell and the current response was measured in the postsynaptic cell at the soma. This recording was repeated 10 times, with a randomly chosen connection location (from anatomically likely locations) each time. Each of the 10 trials are included in this file.

https://doi.org/10.7554/eLife.18566.028
Figure 2—source data 24

Model paired recording of a Pyramidal cell to Ivy cell connection.

Simulated paired recordings where the postsynaptic cell was voltage-clamped at −50 mV and the reversal potential of the synapse was kept at its physiological potential, as defined in the network model code. Sodium channels in the postsynaptic cell were blocked to prevent a suprathreshold response. A spike was triggered in the presynaptic cell and the current response was measured in the postsynaptic cell at the soma. This recording was repeated 10 times, with a randomly chosen connection location (from anatomically likely locations) each time. Each of the 10 trials are included in this file.

https://doi.org/10.7554/eLife.18566.029
Figure 2—source data 25

Model paired recording of a Pyramidal cell to O-LM cell connection.

Simulated paired recordings where the postsynaptic cell was voltage-clamped at −50 mV and the reversal potential of the synapse was kept at its physiological potential, as defined in the network model code. Sodium channels in the postsynaptic cell were blocked to prevent a suprathreshold response. A spike was triggered in the presynaptic cell and the current response was measured in the postsynaptic cell at the soma. This recording was repeated 10 times, with a randomly chosen connection location (from anatomically likely locations) each time. Each of the 10 trials are included in this file.

https://doi.org/10.7554/eLife.18566.030
Figure 2—source data 26

Model paired recording of a Pyramidal cell to PV+ basket cell connection.

Simulated paired recordings where the postsynaptic cell was voltage-clamped at −50 mV and the reversal potential of the synapse was kept at its physiological potential, as defined in the network model code. Sodium channels in the postsynaptic cell were blocked to prevent a suprathreshold response. A spike was triggered in the presynaptic cell and the current response was measured in the postsynaptic cell at the soma. This recording was repeated 10 times, with a randomly chosen connection location (from anatomically likely locations) each time. Each of the 10 trials are included in this file.

https://doi.org/10.7554/eLife.18566.031
Figure 2—source data 27

Model paired recording of a Pyramidal cell to Schaffer Collateral-Associated cell connection.

Simulated paired recordings where the postsynaptic cell was voltage-clamped at −50 mV and the reversal potential of the synapse was kept at its physiological potential, as defined in the network model code. Sodium channels in the postsynaptic cell were blocked to prevent a suprathreshold response. A spike was triggered in the presynaptic cell and the current response was measured in the postsynaptic cell at the soma. This recording was repeated 10 times, with a randomly chosen connection location (from anatomically likely locations) each time. Each of the 10 trials are included in this file.

https://doi.org/10.7554/eLife.18566.032
Detailed network activity.

(A–D) One second of network activity is shown. (A–B) The LFP analog, filtered at (A) the theta range of 5–10 Hz and (B) the low gamma range of 25–40 Hz, shows consistent theta and gamma signals. Scale bar represents 100 ms and 0.2 mV (theta) or 0.27 mV (gamma) for filtered LFP traces. (C) Raster of all spikes from cells within 100 μm of the reference electrode point. (D) Representative intracellular somatic membrane potential traces from cells near the reference electrode point. Scale bar represents 100 ms and 50 mV for the intracellular traces.

https://doi.org/10.7554/eLife.18566.037
Figure 3—source data 1

Filtered analog local field potential of model network.

The theta-filtered and gamma-filtered local field potential (LFP) analog traces.

https://doi.org/10.7554/eLife.18566.038
Figure 3—source data 2

Spike Raster.

Spike times for the length of the entire simulation, from the specific cells displayed in raster shown in Figure 3 (spike times of every single cell in the network are available in the CRCNS repository entry for Bezaire et al. [2016b]).

https://doi.org/10.7554/eLife.18566.039
Figure 3—source data 3

Somatic membrane potential recordings.

Full duration, intracellular somatic membrane potential recordings from the specific cells shown in Figure 3.

https://doi.org/10.7554/eLife.18566.040
Figure 4 with 1 supplement
Spectral analysis of model activity.

(A) A spectrogram of the local pyramidal-layer LFP analog (including contributions from all pyramidal cells within 100 μm of the reference electrode and 10% of pyramidal cells outside that radius) shows the stability and strength of the theta oscillation over time. The oscillation also featured strong harmonics at multiples of the theta frequency of 7.8 Hz. (B,D) Welch’s periodogram of the spike density function for each cell type, normalized by cell type and by displayed frequency range, shows the dominant network frequencies of (B) theta (7.8 Hz) and (D) gamma (71 Hz). Power is normalized to the peak power displayed in the power spectrum for each cell type. (C) Cross-frequency coupling between theta and gamma components of the LFP analog shows that the gamma oscillation is theta modulated. The gamma envelope is a function of the theta phase with the largest amplitude gamma oscillations occurring at the trough of the theta oscillation. Following convention, the theta trough was designated 0°/360°; see e.g., Varga et al. (2012). A graphical explanation of the relation between a spike train and its spike density function is shown in Figure 4—figure supplement 1.

https://doi.org/10.7554/eLife.18566.041
Figure 4—source data 1

Raw analog local field potential of model network.

The raw local field potential (LFP) analog calculated from the network activity, as detailed in the Materials and methods section.

https://doi.org/10.7554/eLife.18566.042
Figure 4—source data 2

Spike Density Functions of each cell type in control network.

The power of the Spike Density Functions was calculated from a one-sided periodogram using Welch’s method where segments have a 50% overlap with a Hamming Window.

https://doi.org/10.7554/eLife.18566.043
Figure 4—figure supplement 1
Different views of cell activity.

Several ways of characterizing model cell activity per cell type are shown using the spikes from the ivy cells as an example. (A) The spike times of each ivy cell are plotted as a function of time and ivy cell number. A subset of ivy cells positioned within 100 μm of the reference electrode location (whose spikes are shown in black) are then carried forward in the remaining calculations. (B) The spikes of the local ivy cells are binned into 1 ms windows to give spike counts per window. (C) A continuous representation of the ivy cell spikes as a function of time is given in the spike density function (SDF) computed from the ivy cell spike times. (D) A Welch’s Periodogram is computed, which summarizes the power of each oscillation frequency in the ivy cell SDF Although only a part of the simulation is shown, the full simulation length (except the first 50 ms) was used in the spectral analysis.

https://doi.org/10.7554/eLife.18566.044
Figure 5 with 2 supplements
Model and experimental cell theta phases.

All model results are based on the spiking of the cells within 100 μm of the reference electrode. (A–B) Firing probability by cell type as a function of theta phase for (A) model and (B) experimental cells under anesthesia (histograms adapted with permission from Figure 2, Figure 5B left, and Figure 6F respectively from Klausberger and Somogyi, 2008; Fuentealba et al., 2008; Fuentealba et al., 2010). The model histograms are normalized; see Figure 5—figure supplement 1 for firing rates. (C) Theta phase preference and theta modulation level were correlated; better modulated cell types spiked closer to the LFP analog trough near the phase preference of pyramidal cells. (D) Theta phase preference plotted on an idealized LFP wave for model data (base of arrow signifies the model phase preference and head of the arrow shows the distance to anesthetized, experimental phase preference).

https://doi.org/10.7554/eLife.18566.045
Figure 5—source data 1

Spike times of axo-axonic cells.

All spike times from all axo-axonic cells, as well as the calculated theta phases (relative to the theta-filtered LFP analog) of each spike.

https://doi.org/10.7554/eLife.18566.046
Figure 5—source data 2

Spike times of bistratified cells.

All spike times from all bistratified cells, as well as the calculated theta phases (relative to the theta-filtered LFP analog) of each spike.

https://doi.org/10.7554/eLife.18566.047
Figure 5—source data 3

Spike times of proximal afferent cells.

All spike times from all proximal afferent cells, as well as the calculated theta phases (relative to the theta-filtered LFP analog) of each spike.

https://doi.org/10.7554/eLife.18566.048
Figure 5—source data 4

Spike times of CCK+ basket cells.

All spike times from all CCK+ basket cells, as well as the calculated theta phases (relative to the theta-filtered LFP analog) of each spike.

https://doi.org/10.7554/eLife.18566.049
Figure 5—source data 5

Spike times of distal afferent cells.

All spike times from all distal afferent cells, as well as the calculated theta phases (relative to the theta-filtered LFP analog) of each spike.

https://doi.org/10.7554/eLife.18566.050
Figure 5—source data 6

Spike times of ivy cells.

All spike times from all ivy cells, as well as the calculated theta phases (relative to the theta-filtered LFP analog) of each spike.

https://doi.org/10.7554/eLife.18566.051
Figure 5—source data 7

Spike times of neurogliaform cells.

All spike times from all neurogliaform cells, as well as the calculated theta phases (relative to the theta-filtered LFP analog) of each spike.

https://doi.org/10.7554/eLife.18566.052
Figure 5—source data 8

Spike times of O-LM cells.

All spike times from all O-LM cells, as well as the calculated theta phases (relative to the theta-filtered LFP analog) of each spike.

https://doi.org/10.7554/eLife.18566.053
Figure 5—source data 9

Spike times of PV+ basket cells.

All spike times from all PV+ basket cells, as well as the calculated theta phases (relative to the theta-filtered LFP analog) of each spike.

https://doi.org/10.7554/eLife.18566.054
Figure 5—source data 10

Spike times of pyramidal cells.

All spike times from all pyramidal cells, as well as the calculated theta phases (relative to the theta-filtered LFP analog) of each spike.

https://doi.org/10.7554/eLife.18566.055
Figure 5—source data 11

Spike times of Schaffer Collateral-associated cells.

All spike times from all Schaffer Collateral-associated cells, as well as the calculated theta phases (relative to the theta-filtered LFP analog) of each spike.

https://doi.org/10.7554/eLife.18566.056
Figure 5—figure supplement 1
Firing rates of model and experimental cells of each type.

For experimental cells, firing rates in both the anesthetized and awake states were included where available. See Table 6 for sources of experimental data.

https://doi.org/10.7554/eLife.18566.057
Figure 5—figure supplement 2
Theta phase-specific firing preferences of various biological hippocampal cell types as reported in the literature.

The trough of the pyramidal-layer LFP is designated as 0/360 and the peak as 180. There is variation in phase preference for given cell types as a function of the experimental preparation. Shown are (A) anesthetized and (B) awake experimental conditions. Reference subscripts correspond to: 1: Klausberger et al. (2003), 2: Klausberger et al. (2004), 3: Klausberger et al. (2005), 4: Lapray et al. (2012), 5: Varga et al. (2012), 6: Fuentealba et al. (2008), 7: Fuentealba et al. (2010), 8: Varga et al. (2014). See Table 6 for further details.

https://doi.org/10.7554/eLife.18566.058
Figure 6 with 2 supplements
Altered network configurations.

Oscillation power (in mV22/Hz) of the spike density function (SDF) for pyramidal cells within 100 μm of the reference electrode, at the peak frequency within theta range (5–10 Hz) in altered network configurations. For corresponding peak frequencies, see Figure 6—figure supplement 1. (A) Theta is present at some excitation levels. (B) Muting each cell type’s output caused a range of effects. (C) The stability and frequency of spontaneous theta in the network was sensitive to the presence and number of recurrent connections between CA1 pyramidal cells. (D) Partially muting the broad classes of PV+ or SOM+ cells by 50% showed that PV+ muting disrupted the network more than SOM+ muting. (E) Theta falls apart when all interneurons are given the same electrophysiological profile, whether it be of a PV+ basket, CCK+ basket, neurogliaform, or O-LM cell. (F) Gradually setting all interneuron properties to those of PV+ basket cells did not restore theta. From left to right: control network; PV+ basket cell electrophysiology; also weights of incoming synapses; also numbers of incoming synapses; then all interneurons being PV+ basket cells (with the addition of the output synapse numbers, weights, and kinetics); then variable RMP (normal distribution with standard deviation of 8 mV). (G) A wide range in excitation was unable to produce theta in the PV+ B. network. (H) Removing the GABAB component from the neurogliaform synapses onto other neurogliaform cells and pyramidal cells showed a significant drop in theta power. Massively increasing the weight of the GABAA component to produce a similar amount of charge transfer restored theta power (compare the IPSCs corresponding to each condition in Figure 6—figure supplement 2). Standard deviations (n = 3) shown; significance (p=1.8e-05).

https://doi.org/10.7554/eLife.18566.061
Figure 6—source data 1

Simulation name mapping.

Map the names of the simulations (used in the header of SDF_All_Conditions.txt) to the bar labels in the graphs of Figure 6.

https://doi.org/10.7554/eLife.18566.062
Figure 6—source data 2

SDF of each network condition.

The full length pyramidal cell Spike Density Function computed at a resolution of 1000 Hz from the spikes of all pyramidal cells within the local range of the electrode point in the model network, for each network condition studied in Figure 6.

https://doi.org/10.7554/eLife.18566.063
Figure 6—figure supplement 1
Peak frequencies of oscillations in altered networks.

Peak theta frequency (within 5–10 Hz) of the spike density function (SDF) for all pyramidal cells within 100 μm of the reference electrode in each altered network configuration. For networks where no pyramidal cells spiked, resulting in zero power within the spectral analysis of the pyramidal cell spike density function, their peak frequencies are listed as ‘not available’ or ‘n/a’. (A) Spontaneous theta oscillation accelerated out of theta range with more excitation. (B) Muting each cell type shifted the oscillation out of range (neurogliaform, CCK+ basket, and axo-axonic cells), disrupted theta but not gamma (not shown; pyramidal, PV+ basket, and bistratified cells), or had little effect (S.C.-A., O-LM, and ivy cells). (C) Doubling the connections between CA1 pyramidal cells increased the theta frequency, while networks with half the number or no recurrent collaterals lost the slow oscillation but kept gamma. (D) Removing 50% of PV+ cell inhibition (PV+ basket, bistratified, and axo-axonic cells) or 50% of SOM+ cell inhibition (bistratified or O-LM cells) shifted the oscillation out of theta range or lost the slow oscillation entirely but kept gamma. (E) Peak oscillation shifted out of theta range when all interneurons had the same electrophysiological profile, regardless of the profile used. (F) Converging all properties to PV+ basket cells, gamma was restored (not shown) but not theta (left to right: control; network with 1: diverse interneurons with same electrophysiology; 2: also with same weights of incoming synapses; 3: also with same numbers of incoming synapses; 4: complete conversion to PV+ basket cells; 5: added variability in resting membrane potential (normal distribution with st. dev.=8 mV)). (G) In the all-PV+ basket cell network, a wide range of excitation levels could not produce a spontaneous theta rhythm. (H) Removing GABAB increased the oscillation frequency.

https://doi.org/10.7554/eLife.18566.064
Figure 6—figure supplement 2
IPSCs from the neurogliaform to pyramidal cell synapse corresponding to the different conditions in Figure 6H.

These traces are from pyramidal cells clamped at −50 mV during a paired recording from a presynaptic neurogliaform cell with a GABAA reversal potential of −60 mV and a GABAB reversal potential of −90 mV. The currents shown are averages from 10 recordings. Scale bar = 100 ms and 5 pA.

https://doi.org/10.7554/eLife.18566.065
Appendix 1—figure 1
Firing Rates of Experimental Cells.

Rebound spiking, which occurs in some O-LM cells at hyperpolarized current injection levels, is not shown in this graph.

https://doi.org/10.7554/eLife.18566.067
Appendix 1—figure 2
Physiological properties of experimental and model cells.

Experimental data are shown with closed markers for the mean and error bars for cell types where n > 1. The model cell properties are plotted as open circles. Calculation of properties is explained in the text. (A) resting membrane potential, (B) threshold, and (C) spike amplitude.

https://doi.org/10.7554/eLife.18566.070
Appendix 1—figure 3
Physiological properties, continued.

(A) sag time constant, (B) sag amplitude, and (C) amplitude of afterhyperpolarization (AHP).

https://doi.org/10.7554/eLife.18566.071
Appendix 1—figure 4
Physiological properties, continued.

(A) rheobase, (B) membrane time constant, (C) interspike interval (ISI), and (D) input resistance.

https://doi.org/10.7554/eLife.18566.072
Appendix 1—figure 5
Pyramidal (A) model and (B) experimental current sweep.

(C) Firing rates of model and experimental cells.

https://doi.org/10.7554/eLife.18566.073
Appendix 1—figure 6
Connections onto (A) and (B) from model Pyramidal cells, under voltage clamp at −50 mV with physiological reversal potentials.
https://doi.org/10.7554/eLife.18566.081
Appendix 1—figure 7
Axo-axonic (A) model and (B) experimental current sweep.

(C) Firing rates of model and experimental cells.

https://doi.org/10.7554/eLife.18566.082
Appendix 1—figure 8
Connections onto (A) and (B) from model Axo-axonic cells, under voltage clamp at −50 mV with physiological reversal potentials.
https://doi.org/10.7554/eLife.18566.089
Appendix 1—figure 9
Bistratified (A) model and (B) experimental current sweep.

(C) Firing rates of model and experimental cells.

https://doi.org/10.7554/eLife.18566.090
Appendix 1—figure 10
Connections onto (A) and (B) from model Bistratified cells, under voltage clamp at −50 mV with physiological reversal potentials.
https://doi.org/10.7554/eLife.18566.098
Appendix 1—figure 11
CCK+ Basket (A) model and (B) experimental current sweep.

(C) Firing rates of model and experimental cells.

https://doi.org/10.7554/eLife.18566.099
Appendix 1—figure 12
Connections onto (A) and (B) from model CCK+ Basket cells, under voltage clamp at −50 mV with physiological reversal potentials.
https://doi.org/10.7554/eLife.18566.106
Appendix 1—figure 13
Ivy (A) model and (B) experimental current sweep.

(fig:ivypage:firing) Firing rates of model and experimental cells.

https://doi.org/10.7554/eLife.18566.107
Appendix 1—figure 14
Connections onto (A) and (B) from model Ivy cells, under voltage clamp at −50 mV with physiological reversal potentials.
https://doi.org/10.7554/eLife.18566.115
Appendix 1—figure 15
Neurogliaform (A) model and (B) experimental current sweep.

(C) Firing rates of model and experimental cells.

https://doi.org/10.7554/eLife.18566.116
Appendix 1—figure 16
Connections onto (A) and (B) from model Neurogliaform cells, under voltage clamp at −50 mV with physiological reversal potentials.
https://doi.org/10.7554/eLife.18566.124
Appendix 1—figure 17
O-LM (A) model and (B) experimental current sweep.

(C) Firing rates of model and experimental cells.

https://doi.org/10.7554/eLife.18566.125
Appendix 1—figure 18
Connections onto (A) and (B) from model O-LM cells, under voltage clamp at −50 mV with physiological reversal potentials.
https://doi.org/10.7554/eLife.18566.132
Appendix 1—figure 19
PV+ Basket (A) model and (B) experimental current sweep.

(C) Firing rates of model and experimental cells.

https://doi.org/10.7554/eLife.18566.133
Appendix 1—figure 20
Connections onto (A) and (B) from model PV+ Basket cells, under voltage clamp at −50 mV with physiological reversal potentials.
https://doi.org/10.7554/eLife.18566.141
Appendix 1—figure 21
Schaffer Collateral-Associated (A) model and (B) experimental current sweep.

(C) Firing rates of model and experimental cells.

https://doi.org/10.7554/eLife.18566.142
Appendix 1—figure 22
Connections onto (A) and (B) from model Schaffer Collateral-Associated cells, under voltage clamp at −50 mV with physiological reversal potentials.
https://doi.org/10.7554/eLife.18566.150
Appendix 1—figure 23
Calcium channel currents.
https://doi.org/10.7554/eLife.18566.156
Appendix 1—figure 24
HCN channel currents.
https://doi.org/10.7554/eLife.18566.157
Appendix 1—figure 25
Delayed rectifier potassium channel currents.
https://doi.org/10.7554/eLife.18566.158
Appendix 1—figure 26
A-type potassium channel currents.
https://doi.org/10.7554/eLife.18566.159
Appendix 1—figure 27
Other potassium channel currents.

Because they didn’t have a voltage-sensitive inactivation component, only the activation curve, which is equivalent to the IV Peak curve, need be shown here.

https://doi.org/10.7554/eLife.18566.160
Appendix 1—figure 28
Calcium-dependent potassium channel dependence on calcium concentration.

(a) The normalized conductance of the channels are plotted as a function of test voltage step and calcium concentration. (b) and (c) The current-voltage relation is shown at several calcium concentrations for (b) KvCaB channel and (c) KCaS channel. Note that the KCaS channel is only active at the highest calcium concentration and is not dependent on voltage (although the voltage continues to set the driving force) when it is active.

https://doi.org/10.7554/eLife.18566.161
Appendix 1—figure 29
Sodium channel voltage dependence.

The normalized conductance of the sodium channel is plotted as (a) a function of test voltage step to show activation and (b) as a function of holding voltage prior to the test step to show inactivation.

https://doi.org/10.7554/eLife.18566.162

Tables

Table 1

Number of synapses between each cell type. Connections between cells generally comprise 1–10 synapses each. Presynaptic cells are listed down the first column (corresponding to each row) and postsynaptic cells are listed along the first row (corresponding to each column).

https://doi.org/10.7554/eLife.18566.033
Pre/PostAxoBisCCK+BIvyNGFO-LMPyrPV+BSC-A
Axo0.00e + 000.00e + 000.00e + 000.00e + 000.00e + 000.00e + 001.12e + 070.00e + 000.00e + 00
Bis2.35e + 053.54e + 055.76e + 052.64e + 050.00e + 006.40e + 053.12e + 078.85e + 056.80e + 04
CCK+B1.41e + 052.12e + 059.79e + 055.64e + 050.00e + 002.62e + 053.24e + 075.31e + 058.32e + 04
Ivy3.53e + 055.30e + 053.42e + 062.11e + 061.00e + 062.23e + 061.28e + 081.33e + 064.08e + 05
NGF0.00e + 000.00e + 000.00e + 000.00e + 006.09e + 050.00e + 004.36e + 070.00e + 000.00e + 00
O-LM1.18e + 051.77e + 051.44e + 060.00e + 004.65e + 059.84e + 042.49e + 074.42e + 051.60e + 05
Pyr7.19e + 052.43e + 060.00e + 002.38e + 050.00e + 001.17e + 076.14e + 077.03e + 061.26e + 05
PV+B5.73e + 048.62e + 041.37e + 057.05e + 040.00e + 000.00e + 005.83e + 072.16e + 059.60e + 03
SC-A8.82e + 031.33e + 041.30e + 051.06e + 050.00e + 001.97e + 043.74e + 063.32e + 041.44e + 04
CA31.23e + 072.56e + 071.44e + 073.39e + 070.00e + 000.00e + 003.73e + 096.69e + 071.55e + 06
ECIII1.43e + 061.91e + 064.02e + 060.00e + 003.75e + 060.00e + 008.09e + 080.00e + 004.58e + 05
Table 2

Simulation time, exchange time, and load balance for simulations executed on various supercomputers and numbers of processors.

https://doi.org/10.7554/eLife.18566.034
Supercomputer# ProcessorsSim time (s)Exchange time (s)Load balance
Comet16802610.281.050.999
Comet17042566.760.650.999
Comet17282601.220.860.999
Comet via NSG17282060.880.830.999
Stampede via NSG20482471.641.711.000
Stampede20482578.320.291.000
Stampede25282189.561.780.999
Stampede30081844.220.910.999
Stampede34881641.910.860.999
Table 3

Electrophysiological characteristics of each model cell type. For more information about model electrophysiology, see the Appendix.

https://doi.org/10.7554/eLife.18566.035
ConditionPyrPV+BCCK+BSC-AAxoBisO-LMIvyNGF
Resting Membrane Potential (mV)−63.0−65.0−70.6−70.5−65.0−67.0−71.5−60.0−60.0
Input Resistance (MΩ)62.252.0211.0272.452.098.7343.8100.0100.0
Membrane Tau (ms)4.86.922.624.47.014.722.421.121.1
Rheobase (pA)250.0300.060.040.0200.0350.050.0160.0170.0
Threshold (mV)52.0−36.6−40.6−43.1−41.6−28.1100.2−27.6−27.7
Delay to 1st Spike (ms)12.474.6166.6127.743.528.48.9173.3119.0
Half-Width (ms)80.70.91.91.60.60.5112.90.60.6
Table 4

Current injection levels used to characterize interneuron current sweeps in Figure 2D–G.

https://doi.org/10.7554/eLife.18566.036
Cell typeHyper. (pA)Step size (pA)Depol. (pA)
PV+ B.−30050+500
CCK+ B.−10020+80
O-LM−13030+80
NGF−13020+190
Table 5

Preferred theta firing phases for each model cell type.

https://doi.org/10.7554/eLife.18566.059
Cell typeFiring rate (Hz)ModulationPhase (0o=trough)
Levelp
Axo.8.90.074.58e − 130163.4
Bis.18.00.760.00e + 00340.0
CCK+ B.54.40.100.00e + 00202.8
Ivy43.30.330.00e + 00142.1
NGF.55.10.071.46e − 32176.3
O-LM17.40.760.00e + 00334.7
Pyr.6.00.740.00e + 00339.7
PV+ B.0.90.460.00e + 00356.8
S.C.-A.5.20.031.13e − 07197.9
Table 6

Firing rates and theta phase preferences for various cell types in various conditions. Theta phase is relative to the LFP recorded in the pyramidal layer, where 0o and 360o are at the trough of the oscillation. non: non-theta/non-SWR state. SWR: sharp wave/ripple. u+k and x: urethane + supplemental doses of ketamine and xylazine.

https://doi.org/10.7554/eLife.18566.060
Cell typeFiring rate (Hz)Theta phase (o)State of animalAnimalRef.
ThetaNonSWR
ADI8.600.060.25156anesth: u+k and xrat(Klausberger et al., 2005)
Axo-axonic17.103.502.95185anesth: u+k and xrat(Klausberger et al., 2003)
Axo-axonic2727251awake, head restraintmouse(Varga et al., 2014)
Bistratified5.900.9042.801anesth: u+k and xrat(Klausberger et al., 2004)
Bistratified34360awake, head restraintmouse(Varga et al., 2014)
Bistratified30.4227.6535.822awakerat(Katona et al., 2014)
CCK+ Basket9.401.602.70174anesth: u+k and xrat(Klausberger et al., 2005)
Ivy0.701.700.8031anesth: u+k and xrat(Fuentealba et al., 2008)
Ivy2.802.105.2046awake, freerat(Lapray et al., 2012)
Ivy2.403.006.70awake, freerat(Fuentealba et al., 2008)
NGF6.002.652.30196anesth: u+k and xrat(Fuentealba et al., 2010)
O-LM4.902.300.2319anesth: u+k and xrat(Klausberger et al., 2003)
O-LM29.8010.4025.40346awake, head restraintmouse(Varga et al., 2012)
O-LM17.3011.8818.95342awakerat(Katona et al., 2014)
PPA5.751.951.50100anesth: u+k and xrat(Klausberger et al., 2005)
PV+ Basket7.302.7432.68271anesth: u+k and xrat(Klausberger et al., 2003)
PV+ Basket234anesth: u+k and xrat(Klausberger et al., 2005)
PV+ Basket21.006.50122.00289awake, freerat(Lapray et al., 2012)
PV+ Basket25.008.2075.00307awake, head restraintmouse(Varga et al., 2012)
PV+ Basket2877310awake, head restraintmouse(Varga et al., 2014)
Pyramidal20anesth: u+k and xrat(Klausberger et al., 2003)
Trilaminar0.200.1069.00troughanesth: u+k and xrat(Ferraguti et al., 2005)
Double Proj.0.900.5526.9377anesth: u+k and xrat(Jinno et al., 2007)
Oriens Retro.0.530.3753.3728anesth: u+k and xrat(Jinno et al., 2007)
Radiatum Retro.5.151.900.70298anesth: u+k and xrat(Jinno et al., 2007)
Table 7

Peak, theta and gamma frequencies and powers of the pyramidal cell spike density function using Welch’s Periodogram. As in Figure 6—figure supplement 1, networks where no pyramidal cells spiked - resulting in zero power within the spectral analysis of the pyramidal cell spike density function - have their peak frequencies listed as ‘n/a’ for ‘not available’.

https://doi.org/10.7554/eLife.18566.066
ThetaGammaOverall
ConditionFrequencyPowerFrequencyPowerFrequencyPower
Tonic excitation level (Hz)
0.20n/a0.0e + 00n/a0.0e + 00n/a0.0e + 00
0.405.95.6e + 0425.44.1e + 0413.76.5e + 04
0.509.88.1e + 0425.41.0e + 0519.55.6e + 05
0.65 (Ctrl.)7.85.0e + 0525.42.0e + 057.85.0e + 05
0.809.87.8e + 0529.32.6e + 059.87.8e + 05
1.009.86.8e + 0529.31.4e + 059.86.8e + 05
1.209.85.1e + 0533.21.8e + 0511.78.2e + 05
1.409.81.9e + 0525.43.4e + 0511.78.6e + 05
Single Interneuron E’phys. Profile
Ctrl7.85.0e + 0525.42.0e + 057.85.0e + 05
O-LMn/a0.0e + 00n/a0.0e + 00n/a0.0e + 00
CCK+B9.85.7e + 0362.56.9e + 0562.56.9e + 05
PV+Bn/a0.0e + 00n/a0.0e + 00n/a0.0e + 00
NGF5.92.6e + 0439.12.4e + 0639.12.4e + 06
Inh. Cells Converge to PV+ B. Cells
Ctrl7.85.0e + 0525.42.0e + 057.85.0e + 05
E’phys.n/a0.0e + 00n/a0.0e + 00n/a0.0e + 00
+input wgt7.86.8e + 0244.91.6e + 0621.53.4e + 06
+input #9.86.1e + 0331.31.1e + 0615.62.0e + 06
All PV+Bn/a0.0e + 00n/a0.0e + 00n/a0.0e + 00
Var. PV+Bn/a0.0e + 00n/a0.0e + 00n/a0.0e + 00
Outputs Muted
Ctrl7.85.0e + 0525.42.0e + 057.85.0e + 05
SOM7.84.7e + 0527.31.4e + 057.84.7e + 05
PV9.83.2e + 0427.38.1e + 0513.71.5e + 06
Pyr to Pyr
2.0x9.81.1e + 0525.47.3e + 0513.71.0e + 06
1.0x (Ctrl.)7.85.0e + 0525.42.0e + 057.85.0e + 05
0.5x7.88.0e + 0429.32.2e + 0529.32.2e + 05
None9.81.1e + 0070.33.7e + 0170.33.7e + 01
Outputs Muted From Each Cell Type
Ctrl7.85.0e + 0525.42.0e + 057.85.0e + 05
Pyr7.81.1e + 0070.33.8e + 0170.33.8e + 01
PV+B9.88.8e + 0329.31.9e + 0629.31.9e + 06
SC-A9.84.9e + 0527.31.8e + 059.84.9e + 05
O-LM7.85.1e + 0525.48.3e + 047.85.1e + 05
NGF9.85.2e + 0327.39.1e + 0513.71.6e + 06
Ivy7.85.3e + 0525.42.0e + 057.85.3e + 05
CCK+B5.95.5e + 0325.43.3e + 033.95.7e + 03
Bis5.91.3e + 0429.31.7e + 0629.31.7e + 06
Axo7.84.0e + 0333.21.2e + 0615.61.9e + 06
Pyr & PV+ B. Network: Tonic Excitation Level (Hz)
0.01n/a0.0e + 00n/a0.0e + 00n/a0.0e + 00
0.05n/a0.0e + 00n/a0.0e + 00n/a0.0e + 00
0.10n/a0.0e + 00n/a0.0e + 00n/a0.0e + 00
0.20n/a0.0e + 00n/a0.0e + 00n/a0.0e + 00
0.405.92.3e + 0225.41.2e + 023.92.4e + 02
0.65n/a0.0e + 00n/a0.0e + 00n/a0.0e + 00
0.80n/a0.0e + 00n/a0.0e + 00n/a0.0e + 00
1.20n/a0.0e + 00n/a0.0e + 00n/a0.0e + 00
Ctrl7.85.0e + 0525.42.0e + 057.85.0e + 05
Appendix 1—table 1

Intrinsic electrophysiological properties of experimental cells.

https://doi.org/10.7554/eLife.18566.068
Cell typenRMP (mV)Input resistance (MΩ)Sag amplitude (mV)Sag tau (ms)Membrane tau (ms)Rheobase (pA)ISI (ms)Threshold (mV)Spike amplitude (mV)AHP (mV)
From mouse
Pyr17−70.7 ± 1.2139.5 ± 38.87.0 ± 2.234.4 ± 11.021.5 ± 8.6182.4 ± 55.7134.0 ± 44.0−36.7 ± 2.678.2 ± 7.28.6 ± 2.1
Axo3−64.4 ± 4.5122.0 ± 57.51.7 ± 0.645.4 ± 6.911.9 ± 2.2283.3 ± 152.847.8 ± 28.5−31.8 ± 3.444.5 ± 6.716.6 ± 3.5
Bis3−63.6 ± 4.7109.1 ± 30.51.7 ± 0.662.3 ± 13.712.2 ± 0.6333.3 ± 57.724.5 ± 21.8−31.9 ± 4.247.3 ± 6.822.6 ± 0.7
O-LM3−64.8 ± 1.3592.3 ± 97.010.4 ± 3.878.5 ± 22.041.4 ± 11.720.0 ± 0.0101.9 ± 30.1−44.2 ± 2.376.3 ± 6.122.1 ± 4.7
PV+B7−61.4 ± 2.065.2 ± 16.21.8 ± 0.562.9 ± 16.313.3 ± 5.4307.1 ± 109.774.2 ± 36.4−35.3 ± 3.751.1 ± 9.018.0 ± 2.7
From rat
CCK+B1−61.2298.12.772.156.060.0261.0−37.763.715.5
Ivy2−62.3 ± 0.3267.2 ± 107.92.4 ± 2.591.1 ± 120.9171.9 ± 45.680.0 ± 28.374.9 ± 20.6−32.8 ± 0.748.2 ± 5.120.1 ± 2.6
NGF2−66.7 ± 13.4260.0 ± 73.61.8 ± 1.661.7 ± 77.077.2 ± 66.2110.0 ± 70.780.0 ± 28.4−34.0 ± 2.234.7 ± 4.916.2 ± 6.3
SC-A2−57.0 ± 4.3529.9 ± 2.97.9 ± 6.291.1 ± 21.774.2 ± 37.330.0 ± 14.1132.4 ± 29.4−34.3 ± 2.258.7 ± 4.512.6 ± 2.0
Appendix 1—table 2

AxoClamp raw data files. Sch. Coll.-Assoc.: Schaffer Collateral-Associated; Super: superficial. Current sweep injection levels are reported as minimum (most hyperpolarized) : step size : maximum (depolarized) level in units of pA.

https://doi.org/10.7554/eLife.18566.069
Cell typeLabCell nameCurrent inj.Original use and methods reference
SpeciesLevels (pA)
Axo-axonicSolteszCA203LF57mouse−200:50:+500unpublished
Axo-axonicSolteszCA204LF59mouse−200:50:+300unpublished
Axo-axonicSolteszCA204RF59mouse−200:50:+400unpublished
BistratifiedSolteszPV16IMmouse−300:50:+400unpublished
BistratifiedSolteszPV74mouse−300:50:+350unpublished
BistratifiedSolteszPV27IMmouse−300:50:+450unpublished
PV+ BasketSolteszPV34mouse−300:50:+500Lee et al. (2014)
PV+ BasketSolteszPV36mouse−300:50:+800Lee et al. (2014)
PV+ BasketSolteszPV37mouse−300:50:+500Lee et al. (2014)
PV+ BasketSolteszPV38mouse−300:50:+300Lee et al. (2014)
PV+ BasketSolteszPV72mouse−300:50:+400Lee et al. (2014)
PV+ BasketSolteszPV80mouse−300:50:+450Lee et al. (2014)
Deep PyramidalSolteszD1_25abfmouse−400:50:+550Lee et al. (2014)
Deep PyramidalSolteszD1_45abfmouse−400:50:+550Lee et al. (2014)
Deep PyramidalSolteszD2_06abfmouse−400:50:+550Lee et al. (2014)
Deep PyramidalSolteszD2_49abfmouse−400:50:+550Lee et al. (2014)
Deep PyramidalSolteszD3_55abfmouse−400:50:+550Lee et al. (2014)
Deep PyramidalSolteszD4_11abfmouse−400:50:+550Lee et al. (2014)
Deep PyramidalSolteszD5_15abfmouse−400:50:+550Lee et al. (2014)
Deep PyramidalSolteszD6_19abfmouse−400:50:+550Lee et al. (2014)
Deep PyramidalSolteszD7mouse−400:50:+550Lee et al. (2014)
Super. PyramidalSolteszS1_04abfmouse−400:50:+550Lee et al. (2014)
Super. PyramidalSolteszS1_47abfmouse−400:50:+550Lee et al. (2014)
Super. PyramidalSolteszS2_08abfmouse−400:50:+550Lee et al. (2014)
Super. PyramidalSolteszS2_31abfmouse−400:50:+550Lee et al. (2014)
Super. PyramidalSolteszS2_51abfmouse−400:50:+550Lee et al. (2014)
Super. PyramidalSolteszS3_13abfmouse−400:50:+550Lee et al. (2014)
Super. PyramidalSolteszS4mouse−400:50:+550Lee et al. (2014)
Super. PyramidalSolteszS5_21abfmouse−400:50:+550Lee et al. (2014)
IvySoltesz0422–1 (File 5)rat−100:20:+890Krook-Magnuson et al. (2011)
IvySoltesz0428–1 (File 4)rat−100:20:+300Krook-Magnuson et al. (2011)
NeurogliaformSoltesz09o21 (File 4)rat−100:20:+120Krook-Magnuson et al. (2011)
NeurogliaformSoltesz09o27 (File 7)rat−100:20:+490Krook-Magnuson et al. (2011)
CCK+ BasketSolteszsh108_BCrat−100:20:+80Lee et al. (2010)
Sch. Coll.-Assoc.Solteszsh114_SCArat−100:20:+60Lee et al. (2010)
Sch. Coll.-Assoc.Solteszsh153_SCArat−100:20:+60Lee et al. (2010)
O-LMMaccaferri1May2012_P3mouse−100:30:+250Quattrocolo and Maccaferri (2013)
O-LMMaccaferri20Sept2011_P2mouse−100:30:+250Quattrocolo and Maccaferri (2013)
O-LMMaccaferri24October2012_C2mouse−100:30:+250Quattrocolo and Maccaferri (2013)
Appendix 1—table 3

Model Pyramidal cell electrophysiological properties.

https://doi.org/10.7554/eLife.18566.074
PropertyValue
RMP−63.0 mV
Input Resistance76.1 MΩ
Sag Amplitude6.5 mV
Sag Tau9.6 ms
Membrane Tau7.1 ms
Rheobase250.0 pA
ISI80.7 ms
Threshold−39.9 mV
Spike Amplitude80.3 mV
Slow AHP Amplitude14.3 mV
Appendix 1—table 4

Model Pyramidal cell ion channels and conductance at highest density location in cell.

https://doi.org/10.7554/eLife.18566.075
ChannelHighest conductance Gmax (S/cm2)
HCNp4.968e-03
Kdrp3.000e-03
KvAdistp4.682e-02
KvAproxp1.599e-02
Navaxonp6.400e-02
Navp3.200e-02
Appendix 1—table 5

Structural connection parameters for Pyramidal cells, based on Bezaire and Soltesz (2013).

https://doi.org/10.7554/eLife.18566.076
Other typeOther cell to pyrPyr to other cell
#Syn.s#Post#Syn.s#Post
Conn.s/Conn.#Loc.Conn.s/Conn.#Loc.
Axo6636axon132apical dendrite
Bis1010100any dendrite337apical dendrite
CCK+B138104any dendrite
Ivy4210420any dendrite030apical dendrite
NGF1410140apical dendrite
O-LM81080apical dendrite13337basal dendrite
Pyr1971197apical dendrite1971197apical dendrite
PV+B1711187soma8322apical dendrite
SC-A030apical dendrite
CA35985211970any dendrite
ECIII129922598any dendrite
Appendix 1—table 6

Experimental constraints for incoming connections onto Pyramidal cells (clamp: black=voltage; purple=current).

https://doi.org/10.7554/eLife.18566.077
Pre typeExp. ref.

Hold (mV)

Erev (mV)Amp. (pA,mV)Diff. %t10-90 (ms)Diff. %τdecay (ms)Diff. %
AxoMaccaferri et al., 2000−70.07.0323.78+5.10.83+3.111.20+0.0
BisMaccaferri et al., 2000−70.07.0143.21−4.52.22+11.215.40−4.3
CCK+BLee et al., 2010−70.0−26.0118.97+3.10.53−16.76.15−4.9
IvyFuentealba et al., 2008−50.0−88.08.17+2.13.50+25.015.43−3.9
NGFPrice et al., 2008−50.0−89.05.25+7.115.48−3.932.73−34.5
O-LMMaccaferri et al., 2000−70.07.024.35−6.34.68−24.618.88−9.3
PyrDeuchars and Thomson, 1996−67.00.00.60−14.56.00+122.220.55+22.3
PV+BSzabadics et al., 2007−70.0−26.091.94−13.90.50−5.76.70+4.7
SC-ALee et al., 2010−70.0−26.052.42−12.91.63+13.68.55+3.0
Appendix 1—table 7

Experimental constraints for outgoing connections from Pyramidal cells (clamp: black=voltage; purple=current).

https://doi.org/10.7554/eLife.18566.078
Post typeExp. ref.Hold (mV)Erev (mV)Amp. (pA,mV)Diff. %t10-90 (ms)Diff. %τdecay (ms)Diff. %
BisPawelzik et al., 2002−66.00.00.77−19.61.58+31.316.75+41.1
IvyFuentealba et al., 2008−65.8−70.00.06−97.91.38−8.321.35+41.1
PyrDeuchars and Thomson, 1996−67.00.00.60−14.56.00+122.219.05+22.3
PV+BLee et al., 2014−60.00.015.09−67.70.28−72.52.00+22.3
Appendix 1—table 8

Model synaptic parameters for Pyramidal cells in the control network.

https://doi.org/10.7554/eLife.18566.079
TypeOther cell to pyrPyr to other cell
Erev (mV)Gmax (nS)τrise (ms)τdecay (ms)Erev (mV)Gmax (nS)τrise (ms)τdecay (ms)
Axo−60.01.150e-030.288.400.04.000e-050.300.60
Bis−60.05.100e-040.119.700.01.900e-030.110.25
CCK+B−60.05.200e-040.204.20
Ivy−60.04.100e-051.1011.000.04.050e-040.300.60
NGF−60.06.500e-059.0039.00
O-LM−60.03.000e-040.1311.000.02.000e-040.300.60
Pyr0.07.000e-020.101.500.07.000e-020.101.50
PV+B−60.02.000e-040.306.200.07.000e-040.070.20
SC-A0.04.050e-040.300.60
CA30.02.000e-040.503.00
ECIII0.02.000e-040.503.00
Appendix 1—table 9

Model synaptic properties under voltage clamp at −50 mV with physiological reversal potentials

https://doi.org/10.7554/eLife.18566.080
TypeOther cell to pyrPyr to other cell
Hold (mV)Erev (mV)Amp. (pA)t10-90 (ms)τdecay (ms)Hold (mV)Erev (mV)Amp. (pA)t10-90 (ms)τdecay (ms)
Axo−50.0−60.036.450.8511.57−50.00.01.850.782.53
Bis−50.0−60.013.472.1715.20−50.00.064.480.281.42
CCK+B−50.0−60.024.860.526.03
Ivy−50.0−60.01.633.6315.35−50.00.040.700.581.28
NGF−50.0−60.01.1065.580.00
O-LM−50.0−60.00.543.7014.10−50.00.017.470.601.53
Pyr−50.00.022.132.229.65−50.00.022.132.229.65
PV+B−50.0−60.020.560.506.70−50.00.014.750.251.77
SC-A−50.00.017.420.683.05
CA3−50.00.07.151.837.08
ECIII−50.00.01.413.2513.63
Appendix 1—table 10

Model Axo-axonic cell electrophysiological properties.

https://doi.org/10.7554/eLife.18566.083
PropertyValue
RMP−65.0 mV
Input Resistance52.3 MΩ
Sag Amplitude
Sag Tau
Membrane Tau7.0 ms
Rheobase200.0 pA
ISI57.3 ms
Threshold−42.0 mV
Spike Amplitude94.3 mV
Slow AHP Amplitude33.4 mV
Appendix 1—table 11

Model Axo-axonic cell ion channels and conductance at highest density location in cell.

https://doi.org/10.7554/eLife.18566.084
ChannelGmax (S/cm2)
CavL5.000e-03
CavN8.000e-04
KCaS2.000e-06
Kdrfast1.300e-02
KvA1.500e-04
KvCaB2.000e-07
Nav1.500e-01
leak1.800e-04
Appendix 1—table 12

Structural connection parameters for Axo-axonic cells, based on Bezaire and Soltesz (2013).

https://doi.org/10.7554/eLife.18566.085
Other typeOther cell to axoAxo to other cell
#Syn.s#Post#Syn.s#Post
Conn.s/Conn.#Loc.Conn.s/Conn.#Loc.
Bis1610160any dendrite
CCK+B12896any dendrite
Ivy2410240any dendrite
O-LM81080apical dendrite
Pyr1623486apical dendrite127167628axon
PV+B39139soma
SC-A166any dendrite
CA3417028340any dendrite
ECIII4852970any dendrite
Appendix 1—table 13

Experimental constraints for outgoing connections from Axo-axonic cells (clamp: black=voltage; purple=current).

https://doi.org/10.7554/eLife.18566.086
Post typeExp. ref.Hold (mV)Erev (mV)Amp. (pA,mV)Diff. %t10-90 (ms)Diff. %τdecay (ms)Diff. %
PyrMaccaferri et al., 2000−70.07.0323.78+5.10.83+3.111.20+0.0
Appendix 1—table 14

Model synaptic parameters for Axo-axonic cells in the control network.

https://doi.org/10.7554/eLife.18566.087
TypeOther cell to axoAxo to other cell
Erev (mV)Gmax (nS)τrise (ms)τdecay (ms)Erev (mV)Gmax (nS)τrise (ms)τdecay (ms)
Bis−60.06.000e-040.292.67
CCK+B−60.07.000e-040.434.49
Ivy−60.05.700e-052.903.10
O-LM−60.01.200e-040.7310.00
Pyr0.04.000e-050.300.60−60.01.150e-030.288.40
PV+B−60.01.200e-040.292.67
SC-A−60.06.000e-040.424.99
CA30.01.200e-042.006.30
ECIII0.01.200e-042.006.30
Appendix 1—table 15

Model synaptic properties under voltage clamp at −50 mV with physiological reversal potentials

https://doi.org/10.7554/eLife.18566.088
TypeOther cell to axoAxo to other cell
Hold (mV)Erev (mV)Amp. (pA)t10-90 (ms)τdecay (ms)Hold (mV)Erev (mV)Amp. (pA)t10-90 (ms)τdecay (ms)
Bis−50.0−60.036.770.703.70
CCK+B−50.0−60.047.290.755.27
Ivy−50.0−60.04.342.136.57
O-LM−50.0−60.04.762.5512.03
Pyr−50.00.01.850.782.53−50.0−60.036.450.8511.57
PV+B−50.0−60.01.080.453.13
SC-A−50.0−60.024.001.006.13
CA3−50.00.010.852.308.80
ECIII−50.00.08.743.089.20
Appendix 1—table 16

Model Bistratified cell electrophysiological properties.

https://doi.org/10.7554/eLife.18566.091
PropertyValue
RMP−67.0 mV
Input Resistance98.8 MΩ
Sag Amplitude0.0 mV
Sag Tau
Membrane Tau14.7 ms
Rheobase350.0 pA
ISI39.0 ms
Threshold−28.1 mV
Spike Amplitude51.2 mV
Slow AHP Amplitude48.8 mV
Appendix 1—table 17

Model Bistratified cell ion channels and conductance at highest density location in cell.

https://doi.org/10.7554/eLife.18566.092
ChannelGmax (S/cm2)
CavL4.000e-03
CavN4.000e-04
KCaS7.000e-07
Kdrfast1.600e-02
KvA5.000e-05
KvCaB7.000e-08
Navbis7.000e-02
leak9.001e-05
Appendix 1—table 18

Structural connection parameters for Bistratified cells, based on Bezaire and Soltesz (2013).

https://doi.org/10.7554/eLife.18566.093
Other typeOther cell to bisBis to other cell
#Syn.s#Post#Syn.s#Post
Conn.s/Conn.#Loc.Conn.s/Conn.#Loc.
Axo1110106any dendrite
Bis1610160any dendrite1610160any dendrite
CCK+B12896any dendrite2610260any dendrite
Ivy2410240any dendrite1210119any dendrite
O-LM81080apical dendrite2910289any dendrite
Pyr36631098apical dendrite14101014095any dendrite
PV+B39139soma4010400any dendrite
SC-A166any dendrite31030any dendrite
CA35782211564any dendrite
ECIII4322864any dendrite
Appendix 1—table 19

Experimental constraints for incoming connections onto Bistratified cells (clamp: black=voltage; purple=current).

https://doi.org/10.7554/eLife.18566.094
Pre typeExp. ref.Hold (mV)Erev (mV)Amp. (pA,mV)Diff. %t10-90 (ms)Diff. %τdecay (ms)Diff. %
PyrPawelzik et al., 2002−66.00.00.77−19.61.58+31.314.68+41.1
PV+BCobb et al., 1997−55.0−70.00.27−27.50.47−52.57.30+30.4
Appendix 1—table 20

Experimental constraints for outgoing connections from Bistratified cells (clamp: black=voltage; purple=current).

https://doi.org/10.7554/eLife.18566.095
Post typeExp. ref.Hold (mV)Erev (mV)Amp. (pA,mV)Diff. %t10-90 (ms)Diff. %τdecay (ms)Diff. %
PyrMaccaferri et al., 2000−70.07.0143.21−4.52.22+11.215.40−4.3
Appendix 1—table 21

Model synaptic parameters for Bistratified cells in the control network.

https://doi.org/10.7554/eLife.18566.096
TypeOther cell to bisBis to other cell
Erev (mV)Gmax (nS)τrise (ms)τdecay (ms)Erev (mV)Gmax (nS)τrise (ms)τdecay (ms)
Axo−60.06.000e-040.292.67
Bis−60.05.100e-040.292.67−60.05.100e-040.292.67
CCK+B−60.07.000e-040.434.49−60.08.000e-040.292.67
Ivy−60.07.700e-052.903.10−60.05.000e-040.292.67
O-LM−60.01.100e-040.6015.00−60.02.000e-051.008.00
Pyr0.01.900e-030.110.25−60.05.100e-040.119.70
PV+B−60.02.900e-030.180.45−60.09.000e-030.292.67
SC-A−60.06.000e-040.424.99−60.08.000e-040.292.67
CA30.01.500e-042.006.30
ECIII0.01.500e-042.006.30
Appendix 1—table 22

Model synaptic properties under voltage clamp at −50 mV with physiological reversal potentials

https://doi.org/10.7554/eLife.18566.097
TypeOther cell to bisBis to other cell
Hold (mV)Erev (mV)Amp. (pA)t10-90 (ms)τdecay (ms)Hold (mV)Erev (mV)Amp. (pA)t10-90 (ms)τdecay (ms)
Axo−50.0−60.036.770.703.70
Bis−50.0−60.034.340.703.72−50.0−60.034.340.703.72
CCK+B−50.0−60.048.130.785.35−50.0−60.048.550.734.15
Ivy−50.0−60.06.392.156.63−50.0−60.043.400.603.17
O-LM−50.0−60.06.312.7017.05−50.0−60.01.861.788.13
Pyr−50.00.064.480.281.42−50.0−60.013.472.1715.20
PV+B−50.0−60.024.450.170.73−50.0−60.0429.340.574.13
SC-A−50.0−60.026.431.026.20−50.0−60.050.350.704.10
CA3−50.00.013.812.388.82
ECIII−50.00.012.043.059.30
Appendix 1—table 23

Model CCK+ Basket cell electrophysiological properties.

https://doi.org/10.7554/eLife.18566.100
PropertyValue
RMP−70.6 mV
Input Resistance222.4 MΩ
Sag Amplitude9.2 mV
Sag Tau45.6 ms
Membrane Tau25.5 ms
Rheobase80.0 pA
ISI180.8 ms
Threshold−38.0 mV
Spike Amplitude65.9 mV
Slow AHP Amplitude32.1 mV
Appendix 1—table 24

Model CCK+ Basket cell ion channels and conductance at highest density location in cell.

https://doi.org/10.7554/eLife.18566.101
ChannelGmax (S/cm2)
CavL2.700e-03
CavN2.000e-05
HCN1.000e-04
KCaS4.000e-06
Kdrfast8.000e-05
KvA4.000e-04
KvCaB4.000e-05
KvGroup2.600e-03
Navcck1.800e-02
leak3.704e-05
Appendix 1—table 25

Structural connection parameters for CCK+ Basket cells, based on Bezaire and Soltesz (2013).

https://doi.org/10.7554/eLife.18566.102
Other typeOther cell to CCK+BCCK+B to other cell
#Syn.s#Post#Syn.s#Post
Conn.s/Conn.#Loc.Conn.s/Conn.#Loc.
Axo5839any dendrite
Bis1610160any dendrite7858any dendrite
CCK+B358280any dendrite358280any dendrite
Ivy9610960any dendrite208156any dendrite
O-LM4010400apical dendrite9872any dendrite
Pyr112588998any dendrite
PV+B38138soma188147any dendrite
SC-A6636any dendrite3824any dendrite
CA3200024000any dendrite
ECIII55921118any dendrite
Appendix 1—table 26

Experimental constraints for outgoing connections from CCK+ Basket cells (clamp: black=voltage; purple=current).

https://doi.org/10.7554/eLife.18566.103
Post typeExp. ref.Hold (mV)Erev (mV)Amp. (pA,mV)Diff. %t10-90 (ms)Diff. %τdecay (ms)Diff. %
PyrLee et al., 2010−70.0−26.0118.97+3.10.53−16.76.15−4.9
Appendix 1—table 27

Model synaptic parameters for CCK+ Basket cells in the control network.

https://doi.org/10.7554/eLife.18566.104
TypeOther cell to CCK+BCCK+B to other cell
Erev (mV)Gmax (nS)τrise (ms)τdecay (ms)Erev (mV)Gmax (nS)τrise (ms)τdecay (ms)
Axo−60.07.000e-040.434.49
Bis−60.08.000e-040.292.67−60.07.000e-040.434.49
CCK+B−60.04.500e-040.434.49−60.04.500e-040.434.49
Ivy−60.03.700e-052.903.10−60.03.000e-040.434.49
O-LM−60.01.200e-030.7320.20−60.07.000e-041.008.00
Pyr−60.05.200e-040.204.20
PV+B−60.01.200e-030.292.67−60.09.000e-030.434.49
SC-A−60.08.500e-040.424.99−60.07.000e-040.434.49
CA30.06.500e-042.006.30
ECIII0.06.500e-042.006.30
Appendix 1—table 28

Model synaptic properties under voltage clamp at −50 mV with physiological reversal potentials

https://doi.org/10.7554/eLife.18566.105
TypeOther cell to CCK+BCCK+B to other cell
Hold (mV)Erev (mV)Amp. (pA)t10-90 (ms)τdecay (ms)Hold (mV)Erev (mV)Amp. (pA)t10-90 (ms)τdecay (ms)
Axo−50.0−60.047.290.755.27
Bis−50.0−60.048.550.734.15−50.0−60.048.130.785.35
CCK+B−50.0−60.032.190.735.30−50.0−60.032.190.735.30
Ivy−50.0−60.03.002.256.95−50.0−60.022.340.805.05
O-LM−50.0−60.040.323.1028.42−50.0−60.054.981.359.05
Pyr−50.0−60.024.860.526.03
PV+B−50.0−60.011.310.423.08−50.0−60.0523.110.685.70
SC-A−50.0−60.033.811.056.90−50.0−60.049.550.705.38
CA3−50.00.055.242.539.35
ECIII−50.00.043.273.4010.87
Appendix 1—table 29

Model Ivy cell electrophysiological properties.

https://doi.org/10.7554/eLife.18566.108
PropertyValue
RMP−60.0 mV
Input Resistance100.7 MΩ
Sag Amplitude0.0 mV
Sag Tau
Membrane Tau21.3 ms
Rheobase160.0 pA
ISI305.5 ms
Threshold−27.7 mV
Spike Amplitude54.6 mV
Slow AHP Amplitude20.9 mV
Appendix 1—table 30

Model Ivy cell ion channels and conductance at highest density location in cell.

https://doi.org/10.7554/eLife.18566.109
ChannelGmax (S/cm2)
CavL5.611e-02
CavN5.817e-04
KCaS4.515e-07
Kdrfastngf1.551e-01
KvAngf5.220e-06
KvCaB1.024e-06
Navngf3.786e+00
leak8.471e-05
Appendix 1—table 31

Structural connection parameters for Ivy cells, based on Bezaire and Soltesz (2013).

https://doi.org/10.7554/eLife.18566.110
OtherTypeOther cell to ivyIvy to other cell
#Syn.s#Post#Syn.s#Post
Conn.s/Conn.#Loc.Conn.s/Conn.#Loc.
Axo41040any dendrite
Bis31030any dendrite61060any dendrite
CCK+B8864any dendrite3910392any dendrite
Ivy2410240any dendrite2410240any dendrite
NGF1110113any dendrite
O-LM2510253any dendrite
Pyr9327apical dendrite14851014850any dendrite
PV+B818soma1510150any dendrite
SC-A2612any dendrite51046any dendrite
CA3192323846any dendrite
Appendix 1—table 32

Experimental constraints for incoming connections onto Ivy cells (clamp: black=voltage; purple=current).

https://doi.org/10.7554/eLife.18566.111
Pre typeExp. ref.Hold (mV)Erev (mV)Amp. (pA,mV)Diff. %t10-90 (ms)Diff. %τdecay (ms)Diff. %
PyrFuentealba et al., 2008−65.8−70.00.06−97.91.38−8.3−4.9
Appendix 1—table 33

Experimental constraints for outgoing connections from Ivy cells (clamp: black=voltage; purple=current).

https://doi.org/10.7554/eLife.18566.112
Post typeExp. ref.Hold (mV)Erev (mV)Amp. (pA,mV)Diff. %t10-90 (ms)Diff. %τdecay (ms)Diff. %
PyrFuentealba et al., 2008−50.0−88.08.17+2.13.50+25.015.43−3.9
Appendix 1—table 34

Model synaptic parameters for Ivy cells in the control network.

https://doi.org/10.7554/eLife.18566.113
TypeOther cell to ivyIvy to other cell
Erev (mV)Gmax (nS)τrise (ms)τdecay (ms)Erev (mV)Gmax (nS)τrise (ms)τdecay (ms)
Axo−60.05.700e-052.903.10
Bis−60.05.000e-040.292.67−60.07.700e-052.903.10
CCK+B−60.03.000e-040.434.49−60.03.700e-052.903.10
Ivy−60.05.700e-052.903.10−60.05.700e-052.903.10
NGF−60.05.700e-052.903.10
O-LM−60.05.700e-052.903.10
Pyr0.04.050e-040.300.60−60.04.100e-051.1011.00
PV+B−60.01.600e-040.292.67−60.07.000e-042.903.10
SC-A−60.08.500e-040.424.99−60.03.700e-052.903.10
CA30.03.000e-042.006.30
Appendix 1—table 35

Model synaptic properties under voltage clamp at −50 mV with physiological reversal potentials

https://doi.org/10.7554/eLife.18566.114
TypeOther cell to ivyIvy to other cell
Hold (mV)Erev (mV)Amp. (pA)t10-90 (ms)τdecay (ms)Hold (mV)Erev (mV)Amp. (pA)t10-90 (ms)τdecay (ms)
Axo−50.0−60.04.342.136.57
Bis−50.0−60.043.400.603.17−50.0−60.06.392.156.63
CCK+B−50.0−60.022.340.805.05−50.0−60.03.002.256.95
Ivy−50.0−60.05.481.886.42−50.0−60.05.481.886.42
NGF−50.0−60.05.481.886.42
O-LM−50.0−60.05.322.106.33
Pyr−50.00.040.700.581.28−50.0−60.01.633.6315.35
PV+B−50.0−60.01.440.553.13−50.0−60.051.352.056.75
SC-A−50.0−60.046.620.855.58−50.0−60.03.092.226.88
CA3−50.00.029.422.058.60
Appendix 1—table 36

Model Neurogliaform cell electrophysiological properties.

https://doi.org/10.7554/eLife.18566.117
PropertyValue
RMP−60.0 mV
Input Resistance100.8 MΩ
Sag Amplitude0.0 mV
Sag Tau
Membrane Tau21.3 ms
Rheobase170.0 pA
ISI170.3 ms
Threshold−27.8 mV
Spike Amplitude55.2 mV
Slow AHP Amplitude20.6 mV
Appendix 1—table 37

Model Neurogliaform cell ion channels and conductance at highest density location in cell.

https://doi.org/10.7554/eLife.18566.118
ChannelGmax (S/cm2)
CavL5.611e-02
CavN5.817e-04
KCaS4.515e-07
Kdrfastngf1.551e-01
KvAngf5.220e-06
KvCaB1.024e-06
Navngf3.786e+00
leak8.471e-05
Appendix 1—table 38

Structural connection parameters for Neurogliaform cells, based on Bezaire and Soltesz (2013).

https://doi.org/10.7554/eLife.18566.119
Other typeOther cell to NGFNGF to other cell
#Syn.s#Post#Syn.s#Post
Conn.s/Conn.#Loc.Conn.s/Conn.#Loc.
Ivy2810280any dendrite
NGF1710170apical dendrite1710170apical dendrite
O-LM1310130apical dendrite
Pyr12181012181apical dendrite
ECIII52321046any dendrite
Appendix 1—table 39

Experimental constraints for incoming connections onto Neurogliaform cells (clamp: black=voltage; purple=current).

https://doi.org/10.7554/eLife.18566.120
Pre typeExp. ref.Hold (mV)Erev (mV)Amp. (pA,mV)Diff. %t10-90 (ms)Diff. %τdecay (ms)Diff. %
NGFKarayannis et al., 2010−65.0−11.085.50+0.24.83−1.732.03−46.9
O-LMElfant et al., 2007−50.0−70.018.43−4.01.98−10.211.63+7.6
Appendix 1—table 40

Experimental constraints for outgoing connections from Neurogliaform cells (clamp: black=voltage; purple=current).

https://doi.org/10.7554/eLife.18566.121
Post typeExp. ref.Hold (mV)Erev (mV)Amp. (pA,mV)Diff. %t10-90 (ms)Diff. %τdecay (ms)Diff. %
NGFKarayannis et al., 2010−65.0−11.085.50+0.24.83−1.732.03−46.9
PyrPrice et al., 2008−50.0−89.05.25+7.115.48−3.932.73−34.5
Appendix 1—table 41

Model synaptic parameters for Neurogliaform cells in the control network.

https://doi.org/10.7554/eLife.18566.122
TypeOther cell to NGFNGF to other cell
Erev (mV)Gmax (nS)𝛕rise (ms)𝛕decay (ms)Erev (mV)Gmax (nS)𝛕rise (ms)𝛕decay (ms)
Ivy−60.05.700e-052.903.10
NGF−60.01.600e-043.1042.00−60.01.600e-043.1042.00
O-LM−60.09.800e-051.3010.20
Pyr−60.06.500e-059.0039.00
ECIII0.03.500e-032.006.30
Appendix 1—table 42

Model synaptic properties under voltage clamp at −50 mV with physiological reversal potentials

https://doi.org/10.7554/eLife.18566.123
TypeOther cell to NGFNGF to other cell
Hold (mV)Erev (mV)Amp. (pA)t10-90 (ms)𝛕decay (ms)Hold (mV)Erev (mV)Amp. (pA)t10-90 (ms)𝛕decay (ms)
Ivy−50.0−60.05.481.886.42
NGF−50.0−60.017.525.6714.32−50.0−60.017.525.6714.32
O-LM−50.0−60.09.141.9811.63
Pyr−50.0−60.01.1065.580.00
ECIII−50.00.0324.352.138.80
Appendix 1—table 43

Model O-LM cell electrophysiological properties.

https://doi.org/10.7554/eLife.18566.126
PropertyValue
RMP−68.0 mV
Input Resistance267.7 MΩ
Sag Amplitude26.5 mV
Sag Tau42.5 ms
Membrane Tau22.7 ms
Rheobase20.0 pA
ISI66.9 ms
Threshold−37.8 mV
Spike Amplitude42.6 mV
Slow AHP Amplitude34.6 mV
Appendix 1—table 44

Model O-LM cell ion channels and conductance at highest density location in cell.

https://doi.org/10.7554/eLife.18566.127
ChannelGmax (S/cm2)
HCNolm5.000e-04
Kdrfast1.174e-01
KvAolm4.950e-03
Nav2.340e-02
leak1.000e-05
Appendix 1—table 45

Structural connection parameters for O-LM cells, based on Bezaire and Soltesz (2013).

https://doi.org/10.7554/eLife.18566.128
Other typeOther cell to O-LMO-LM to other cell
#Syn.s#Post#Syn.s#Post
Conn.s/Conn.#Loc.Conn.s/Conn.#Loc.
Axo71071apical dendrite
Bis3910390any dendrite1110107apical dendrite
CCK+B208160any dendrite8810878apical dendrite
Ivy136101360any dendrite
NGF2810283apical dendrite
O-LM61060basal dendrite61060basal dendrite
Pyr237937137basal dendrite15201015195apical dendrite
PV+B2710269apical dendrite
SC-A101097apical dendrite
Appendix 1—table 46

Experimental constraints for outgoing connections from O-LM cells (clamp: black=voltage; purple=current).

https://doi.org/10.7554/eLife.18566.129
Post typeExp. ref.Hold (mV)Erev (mV)Amp. (pA,mV)Diff. %t10-90 (ms)Diff. %𝛕decay (ms)Diff. %
NGFElfant et al., 2007−50.0−70.018.43−4.01.98−10.211.63+7.6
PyrMaccaferri et al., 2000−70.07.024.35−6.34.68−24.618.88−9.3
SC-AElfant et al., 2007−50.0−70.017.06−12.54.07+114.530.08−3.6
Appendix 1—table 47

Model synaptic parameters for O-LM cells in the control network.

https://doi.org/10.7554/eLife.18566.130
TypeOther cell to O-LMO-LM to other cell
Erev (mV)Gmax (nS)𝛕rise (ms)𝛕decay (ms)Erev (mV)Gmax (nS)𝛕rise (ms)𝛕decay (ms)
Axo−60.01.200e-040.7310.00
Bis−60.02.000e-051.008.00−60.01.100e-040.6015.00
CCK+B−60.07.000e-041.008.00−60.01.200e-030.7320.20
Ivy−60.05.700e-052.903.10
NGF−60.09.800e-051.3010.20
O-LM−60.01.200e-030.257.50−60.01.200e-030.257.50
Pyr0.02.000e-040.300.60−60.03.000e-040.1311.00
PV+B−60.01.100e-030.257.50
SC-A−60.01.500e-040.0729.00
Appendix 1—table 48

Model synaptic properties under voltage clamp at −50 mV with physiological reversal potentials

https://doi.org/10.7554/eLife.18566.131
TypeOther cell to O-LMO-LM to other cell
Hold (mV)Erev (mV)Amp. (pA)t10-90 (ms)𝛕decay (ms)Hold (mV)Erev (mV)Amp. (pA)t10-90 (ms)𝛕decay(ms)
Axo−50.0−60.04.762.5512.03
Bis−50.0−60.01.861.788.13−50.0−60.06.312.7017.05
CCK+B−50.0−60.054.981.359.05−50.0−60.040.323.1028.42
Ivy−50.0−60.05.322.106.33
NGF−50.0−60.09.141.9811.63
O-LM−50.0−60.078.691.059.30−50.0−60.078.691.059.30
Pyr−50.00.017.470.601.53−50.0−60.00.543.7014.10
PV+B−50.0−60.035.531.6510.18
SC-A−50.0−60.07.913.9029.83
Appendix 1—table 49

Model PV+ Basket cell electrophysiological properties.

https://doi.org/10.7554/eLife.18566.134
PropertyValue
RMP−65.0 mV
Input Resistance52.1 MΩ
Sag Amplitude
Sag Tau
Membrane Tau7.0 ms
Rheobase300.0 pA
ISI151.4 ms
Threshold−36.7 mV
Spike Amplitude90.7 mV
Slow AHP Amplitude41.4 mV
Appendix 1—table 50

Model PV+ Basket cell ion channels and conductance at highest density location in cell.

https://doi.org/10.7554/eLife.18566.135
ChannelGmax (S/cm2)
CavL5.000e-03
CavN8.000e-04
KCaS2.000e-06
Kdrfast1.300e-02
KvA1.500e-04
KvCaB2.000e-07
Navaxonp1.500e-01
leak1.800e-04
Appendix 1—table 51

Structural connection parameters for PV+ Basket cells, based on Bezaire and Soltesz (2013).

https://doi.org/10.7554/eLife.18566.136
Other typeOther cell to PV+BPV+B to other cell
#Syn.s#Post#Syn.s#Post
Conn.s/Conn.#Loc.Conn.s/Conn.#Loc.
Axo10110soma
Bis1610160any dendrite16115soma
CCK+B12896any dendrite25124soma
Ivy2410240any dendrite13112soma
O-LM81080apical dendrite
Pyr42431272apical dendrite9581110533soma
PV+B39139soma39139soma
SC-A211soma
CA36047212094any dendrite
Appendix 1—table 52

Experimental constraints for incoming connections onto PV+ Basket cells (clamp: black=voltage; purple=current).

https://doi.org/10.7554/eLife.18566.137
Pre typeExp. ref.Hold (mV)Erev (mV)Amp. (pA,mV)Diff. %t10-90 (ms)Diff. %𝛕decay (ms)Diff. %
PyrLee et al., 2014−60.00.015.09−67.70.28−72.51.83−55.7
PV+BCobb et al., 1997−59.0−70.00.29+14.92.67+105.814.72−45.5
Appendix 1—table 53

Experimental constraints for outgoing connections from PV+ Basket cells (clamp: black=voltage; purple=current).

https://doi.org/10.7554/eLife.18566.138
Post typeExp. ref.Hold (mV)Erev (mV)Amp. (pA,mV)Diff. %t10-90 (ms)Diff. %𝛕decay (ms)Diff. %
BisCobb et al., 1997−55.0−70.00.27−27.50.47−52.511.85+30.4
PyrSzabadics et al., 2007−70.0−26.091.94−13.90.50−5.76.70+4.7
PV+BCobb et al., 1997−59.0−70.00.29+14.92.67+105.813.45−45.5
Appendix 1—table 54

Model synaptic parameters for PV+ Basket cells in the control network.

https://doi.org/10.7554/eLife.18566.139
TypeOther cell to PV+BPV+B to other cell
Erev (mV)Gmax(nS)𝛕rise (ms)𝛕decay (ms)Erev (mV)Gmax (nS)𝛕rise (ms)𝛕decay (ms)
Axo−60.01.200e-040.292.67
Bis−60.09.000e-030.292.67−60.02.900e-030.180.45
CCK+B−60.09.000e-030.434.49−60.01.200e-030.292.67
Ivy−60.07.000e-042.903.10−60.01.600e-040.292.67
O-LM−60.01.100e-030.257.50
Pyr0.07.000e-040.070.20−60.02.000e-040.306.20
PV+B−60.01.600e-030.084.80−60.01.600e-030.084.80
SC-A−60.06.000e-040.292.67
CA30.02.200e-042.006.30
Appendix 1—table 55

Model synaptic properties under voltage clamp at −50 mV with physiological reversal potentials

https://doi.org/10.7554/eLife.18566.140
TypeOther cell to PV+BPV+B to other cell
Hold (mV)Erev (mV)Amp. (pA)t10-90 (ms)𝛕decay (ms)Hold (mV)Erev (mV)Amp. (pA)t10-90 (ms)𝛕decay (ms)
Axo−50.0−60.01.080.453.13
Bis−50.0−60.0429.340.574.13−50.0−60.024.450.170.73
CCK+B−50.0−60.0523.110.685.70−50.0−60.011.310.423.08
Ivy−50.0−60.051.352.056.75−50.0−60.01.440.553.13
O-LM−50.0−60.035.531.6510.18
Pyr−50.00.014.750.251.77−50.0−60.020.560.506.70
PV+B−50.0−60.013.940.235.25−50.0−60.013.940.235.25
SC-A−50.0−60.05.710.423.08
CA3−50.00.019.712.388.78
Appendix 1—table 56

Model Schaffer Collateral-Associated cell electrophysiological properties.

https://doi.org/10.7554/eLife.18566.143
PropertyValue
RMP−70.5 mV
Input Resistance300.0 MΩ
Sag Amplitude12.9 mV
Sag Tau41.7 ms
Membrane Tau28.9 ms
Rheobase60.0 pA
ISI115.9 ms
Threshold−36.6 mV
Spike Amplitude80.3 mV
Slow AHP Amplitude35.2 mV
Appendix 1—table 57

Model Schaffer Collateral-Associated cell ion channels and conductance at highest density location in cell.

https://doi.org/10.7554/eLife.18566.144
ChannelGmax (S/cm2)
CavL1.000e-03
CavN2.000e-05
HCN7.000e-05
KCaS1.000e-06
Kdrfast6.000e-05
KvA1.000e-04
KvCaB7.000e-06
KvGroup2.200e-03
Navcck4.000e-02
leak2.857e-05
Appendix 1—table 58

Structural connection parameters for Schaffer Collateral-Associated cells, based on Bezaire and Soltesz (2013).

https://doi.org/10.7554/eLife.18566.145
Other typeOther cell to SC-ASC-A to other cell
#Syn.s#Post#Syn.s#Post
Conn.s/Conn.#Loc.Conn.s/Conn.#Loc.
Axo4622any dendrite
Bis1710170any dendrite6633any dendrite
CCK+B278216any dendrite546324any dendrite
Ivy102101020any dendrite446264any dendrite
O-LM4010400apical dendrite
Pyr1053315apical dendrite
PV+B24124soma
CA3194023880any dendrite
ECIII57321146any dendrite
Appendix 1—table 59

Experimental constraints for incoming connections onto Schaffer Collateral-Associated cells (clamp: black=voltage; purple=current).

https://doi.org/10.7554/eLife.18566.146
Pre typeExp. ref.Hold (mV)Erev (mV)Amp. (pA,mV)Diff. %t10-90 (ms)Diff. %𝛕decay (ms)Diff.%
O-LMElfant et al., 2007−50.0−70.017.06−12.54.07+114.530.08−3.6
SC-APawelzik et al., 2002−58.0−70.01.53−405.62.35−41.322.90−33.2
Appendix 1—table 60

Experimental constraints for outgoing connections from Schaffer Collateral-Associated cells (clamp: black=voltage; purple=current).

https://doi.org/10.7554/eLife.18566.147
Post typeExp. ref.Hold (mV)Erev (mV)Amp. (pA,mV)Diff. %t10-90 (ms)Diff. %𝛕decay (ms)Diff. %
PyrLee et al., 2010−70.0−26.052.42−12.91.63+13.68.55+3.0
SC-APawelzik et al., 2002−58.0−70.01.53−405.62.35−41.327.98−33.2
Appendix 1—table 61

Model synaptic parameters for Schaffer Collateral-Associated cells in the control network.

https://doi.org/10.7554/eLife.18566.148
Other cell to SC-ASC-A to other cell
TypeErev (mV)Gmax (nS)𝛕rise (ms)𝛕decay (ms)Erev (mV)Gmax (nS)𝛕rise (ms)𝛕decay (ms)
Axo−60.06.000e-040.424.99
Bis−60.08.000e-040.292.67−60.06.000e-040.424.99
CCK+B−60.07.000e-040.434.49−60.08.500e-040.424.99
Ivy−60.03.700e-052.903.10−60.08.500e-040.424.99
O-LM−60.01.500e-040.0729.00
Pyr0.04.050e-040.300.60
PV+B−60.06.000e-040.292.67
CA30.03.000e-042.006.30
ECIII0.04.500e-042.006.30
Appendix 1—table 62

Model synaptic properties under voltage clamp at −50 mV with physiological reversal potentials

https://doi.org/10.7554/eLife.18566.149
TypeOther cell to SC-ASC-A to other cell
Hold (mV)Erev (mV)Amp. (pA)t10-90 (ms)𝛕decay (ms)Hold (mV)Erev (mV)Amp. (pA)t10-90 (ms)𝛕decay (ms)
Axo−50.0−60.024.001.006.13
Bis−50.0−60.050.350.704.10−50.0−60.026.431.026.20
CCK+B−50.0−60.049.550.705.38−50.0−60.033.811.056.90
Ivy−50.0−60.03.092.226.88−50.0−60.046.620.855.58
O-LM−50.0−60.07.913.9029.83
Pyr−50.00.017.420.683.05
PV+B−50.0−60.05.710.423.08
CA3−50.00.027.102.359.13
ECIII−50.00.031.823.3810.47
Appendix 1—table 63

Measured dendritic lengths and somatic diameters for ivy and neurogliaform cells from the hippocampal CA1 area in Wistar rats, with calculation of somatic surface area included. Cells were characterized in our lab and their function has been reported in Krook-Magnuson et al. (2011). Source Data available in Appendix 1—table 1 - Source Data.

https://doi.org/10.7554/eLife.18566.151
Cell typeCell nameDendritic length (μM))Somatic dia-meter (μm)Calculated synap-tic Area (μm2)
# SectionsSOSPSRSLM
Ivy0217–1 DAB 3_2_10 left slice1129.264.51200.6038.91188.5
Ivy9 n23-7 DAB 12_16_09 left+middle slice2002703.2300.345.21604.6
Ivy9 n23-6 DAB 06_10 left slice175.4133.82115.4036.61052.1
Ivy9 n16-3 DAB 12_29_09 left slice1001015.2052.52164.8
IvyAverage51.1549.5751758.675.0751502.5
Neurogliaform9n 12–5 DAB 1_06_091002097.752534.4929.4
Neurogliaform91021 DAB 3_18_10 second,third,fourth from left slice3001230.7780.128615.8
Neurogliaform9d 8–3 DAB 1_15_10 left and right slice2002328.21382.432.2814.3
NeurogliaformAverage001885.5895.8786.5
Appendix 1—table 64

Estimated or observed somatic area and dendritic length. Experimental observations of the dendritic length of broad interneuron classes were used as the basis for these estimations. The relative lengths for PV+ basket cells and axo-axonic cells were further differentiated based on experimental observations in region CA3 Papp et al. (2013). The observations published in Mátyás et al. (2004) for CCK+ basket cells were also applied to the CCK+ Schaffer Collateral-Associated cells, based on the discussion in Mátyás et al. (2004). The data for ivy and neurogliaform cells were based on measurements from filled cells from slices. Due to the compact nature of their morphology, especially the neurogliaform cells, the dendritic lengths within the slices were assumed to comprise most or all of the dendritic extents of those cells. See section below for raw data. The O-LM cell morphological measurements were taken from Blasco-Ibáñez and Freund (1995).

https://doi.org/10.7554/eLife.18566.152
InterneuronSoma area (100 μm2)Dendritic length (μm2)Reference
TotalSOSPSRSLM
Ivy15021934.451.1549.5751758.675.075See below
Neurogliaform7862781.4001885.5895.8See below
PV+ basket3428435914936971877292(Papp et al., 2013)
Bistratified10064347.751074.57248.282369.24655.66(Gulyás et al., 1999)
Axo-axonic232928255706591259337(Papp et al., 2013)
CCK+ basket9666338.311213.92310.613522.61291.18(Mátyás et al., 2004)
SCA9666338.311213.92310.613522.61291.18(Mátyás et al., 2004)
O-LM3007.784165.684165.68000(Blasco-Ibáñez and Freund, 1995)
Appendix 1—table 65

The synaptic densities (# boutons per 100 μm of dendritic length, or # boutons per 100 μm2 of somatic area) on the soma and dendrites of PV cells, given in Gulyás et al. (1999), were applied to the axo-axonic, PV+ basket, and bistratified cells. The synaptic densities of CCK+ cells Mátyás et al. (2004) were applied to the CCK+ basket and the Schaffer Collateral-Associated cells. For the ivy, neurogliaform, and O-LM cells, there were not sufficient experimental data published to constrain the synaptic density, and so an average of all synaptic densities for all cell classes was computed and applied to these cell types.

https://doi.org/10.7554/eLife.18566.153
Dendritic
SomaticSOSPSRSLM
ReferenceExcInhExcInhExcInhExcInhExcInhRef
Ivy21.816.1172.223.7163.338.5193.825.397.431.7Calc. from average
Neurogliaform21.816.1172.223.7163.338.5193.825.397.431.7Calc. from average
PV+ basket40.718.1342.519.234516.1371.218.3132.228.6(Gulyás et al., 1999)
Bistratified40.718.1342.519.234516.1371.218.3132.228.6(Gulyás et al., 1999)
Axo-axonic40.718.1342.519.234516.1371.218.3132.228.6(Gulyás et al., 1999)
CCK+ basket3.416.184.332.552.787.48237.886.558.8(Mátyás et al., 2004)
SCA3.416.184.332.552.787.48237.886.558.8(Mátyás et al., 2004)
O-LM21.816.1172.223.7163.338.5193.825.397.431.7Calc. from average
Appendix 1—table 66

Estimated numbers of excitatory and inhibitory synapses on each cell type, calculated by multiplying the somatic area or dendritic length by the respective synaptic density. About 20% of synapses onto O-LM cells are GABAergic, while at least 60% are from local excitatory collaterals Kispersky et al. (2012). Therefore, we conserved the total (inhibitory + excitatory) synaptic density of O-LM cells as calculated previously, but set 20% of that total to be inhibitory and the rest to be excitatory synapses.

https://doi.org/10.7554/eLife.18566.154
Dendritic
SomaticSOSPSRSLMTotal
RefExcInhExcInhExcInhExcInhExcInhExcInhRef
Ivy326.9242.388128219340844573243651500Calculated
Neurogliaform171.1126.8000036544778732844527761Calculated
PV+ basket1395.2620.1423023786640944946684018215385925Calculated
Bistratified409.4182368120685640879643486718814200868Calculated
Axo-axonic947.9421.31615908193863383139692109741651Calculated
CCK+ basket32.8155.7102439416427128871332111775951922756Calculated
SCA32.8155.7102439416427128871332111775951922756Calculated
O-LM654.6485.26527163200000065271632Calculated
Appendix 1—table 67

Ion channels included in the model. GHK: based on Goldman-Hodgkin-Katz equation; Q-O: quasi-ohmic; Hyperpol.-act: Hyperpolarization-activated; Nucleo.-gated: Nucleotide-gated; voltage-act.: voltage activated; voltage-dep.: voltage dependent; Calcium-act.: Calcium-activated; Pyr.: pyramidal; NGF: neurogliaform; dist.: distal; prox.: proximal.

https://doi.org/10.7554/eLife.18566.155
IonModel
ChannelDescriptionTypePyramidalAxo-axonicBistratifiedCCK+ BasketIvyNeurogliaformO-LMPV+ BasketS.C.-Assoc.
Cav,LL-type CalciumGHK
Cav,NN-type CalciumQ-O
HCNHyperpol.-act, Cyclic Nucleo.-gatedQ-O
HCNOLMHyperpol.-act, Cyclic Nucleo.-gated for O-LM cellsQ-O
HCNpHyperpol.-act, Cyclic Nucleo.-gated for Pyr. cellsQ-O
KCa,SSmall (SK) Calcium-activated potassiumQ-O
Kdr,fastFast delayed rectifier potassiumQ-O
Kdr,fast,ngfFast delayed rectifier potassium for NGF-family cellsQ-O
Kdr,pDelayed rectifier potassium for Pyr. cellsQ-O
Kv,AA-type voltage-act. potassiumQ-O
Kv,A,dist,pA-type voltage-act. potassium for dist. Pyr. dendritesQ-O
Kv,A,ngfA-type voltage-act. potassium for NGF-family cellsQ-O
Kv,A,olmA-type voltage-act. potassium for O-LM cellsQ-O
Kv,A,prox,pA-type voltage-act. potassium for prox. Pyr. dendritesQ-O
Kv,Ca,BBig (BK) Calcium-act., voltage-dep. potassiumQ-O
Kv,GroupMultiple slower voltage-dep. potassiumQ-O
leakLeakQ-O
NavVoltage-dep. sodiumQ-O
Nav,bisVoltage-dep. sodium for bistratified cellsQ-O
Nav,cckVoltage-dep. sodium for CCK+ cellsQ-O
Nav,ngfVoltage-dep. sodium for NGF-family cellsQ-O
Nav,pVoltage-dep. sodium for Pyr. cellsQ-O
pasLeakQ-O

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  1. Marianne J Bezaire
  2. Ivan Raikov
  3. Kelly Burk
  4. Dhrumil Vyas
  5. Ivan Soltesz
(2016)
Interneuronal mechanisms of hippocampal theta oscillations in a full-scale model of the rodent CA1 circuit
eLife 5:e18566.
https://doi.org/10.7554/eLife.18566