From 65a1b3372dde8e464d1c2ad962232ea09b013349 Mon Sep 17 00:00:00 2001
From: Michael Clerx <michael.clerx@nottingham.ac.uk>
Date: Thu, 19 Mar 2026 16:16:42 +0000
Subject: [PATCH 1/2] Cosmetic fix to level 0 schematic

---
 tutorial/img/vc-level-0.png | Bin 22325 -> 22679 bytes
 1 file changed, 0 insertions(+), 0 deletions(-)

diff --git a/tutorial/img/vc-level-0.png b/tutorial/img/vc-level-0.png
index b16aec722ba21c7bad1001a2c6b213bc640e799e..39904ef657329487a838855ecf8b287ef053a950 100644
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From 0f9131229be8fdfee22d8208d795665e7f82ec44 Mon Sep 17 00:00:00 2001
From: Michael Clerx <michael.clerx@nottingham.ac.uk>
Date: Thu, 19 Mar 2026 16:33:37 +0000
Subject: [PATCH 2/2] Finished model tour. Closes #3. Closes #4. Closes 16.

---
 README.md           |  13 ++---
 tutorial/README.md  |   8 +--
 tutorial/tour.ipynb | 122 +++++++++++++++++++-------------------------
 3 files changed, 64 insertions(+), 79 deletions(-)

diff --git a/README.md b/README.md
index 85762ae..f85ed1f 100644
--- a/README.md
+++ b/README.md
@@ -3,11 +3,11 @@
 This repository contains up-to-date reference versions of the voltage clamp model developed in Lei et al. 2020 and 2025.
 These models can be used to simulate manual or planar patch-clamp experiments in voltage-clamp mode.
 
-_Note: This URL originally hosted the data for the 2020 Lei et al. publication. This is now found at https://github.com/CardiacModelling/VoltageClampModel2020_
+_Note: This URL originally hosted data and code for the 2020 Lei et al. publication. This is now found at https://github.com/CardiacModelling/VoltageClampModel2020_
 
 ## The latest models
 
-We recently updated the voltage clamp model with improved filtering of input (stimulus filter) and output (Bessel filters), and an improved time-delay in the series resistance compensation pathway.
+We recently updated the voltage clamp model with improved filtering of input (stimulus filter) and output, and an improved time-delay in the series resistance compensation pathway.
 
 In addition to this full-featured model, we provide a set of increasingly simplified models:
 
@@ -26,14 +26,15 @@ All models are provided in Myokit ([models-mmt](./models-mmt)) and CellML ([mode
 
 In brief:
 - Levels 0, 1, and 2 can recreate the _fast_ artefacts seen in patch clamp experiments, and are very useful _to understand the patch clamp process_.
-- _To fit experimental data_, these fast artefacts are less important, and so level 3 is good to match data from the fastest currents. For slower currents, levels 4 or 5 can be used.
+- _To fit experimental data_, these fast artefacts are less important, and so level 3 is good to match data from the fastest currents.
+For slower currents, levels 4 or 5 can be used.
 
 ## Tutorials
 
-We provide four [Tutorial Notebooks](./tutorial/README.md) to understand these notebooks, and how they were derived.
+We provide four [Tutorial Notebooks](./tutorial/README.md) to understand these models, and how they were derived.
 
 The first notebook establishes a basic model of a patch clamp amplifier, and the second adds compensation and filtering, leading to the "Level 0" model.
-In the third notebook, this model is used to simulate the early stages of a (manual) patch-clamp experiment.
+In the third notebook, this model is used to simulate a manual patch-clamp experiment.
 
 The final notebook derives the Level 1 to 5 simplifications, and shows how they relate to our previous work (Lei et al., 2020 and 2025).
 
@@ -91,5 +92,5 @@ Code: https://github.com/CardiacModelling/VoltageClampModel2020
 
 ## Acknowledging this work
 
-If you publish any work based on the contents of this repository please cite the papers listed above.
+If you use the contents of this repository in your work, please cite the papers listed above.
 
diff --git a/tutorial/README.md b/tutorial/README.md
index 4d73918..1f360c6 100644
--- a/tutorial/README.md
+++ b/tutorial/README.md
@@ -1,9 +1,9 @@
 # Voltage clamp model tutorials
 
-In these notebooks we retrace the steps taken in the supplement to [Lei et al., 2020](https://doi.org/10.1098/rsta.2019.0348) and [2025](https://doi.org/10.1002/advs.202500691), and construct (1) a model of a patch-clamp experiment with various experimental artefacts, and (2) a model of the corrections applied by patch-clamp amplifiers to mitigate these effects.
+In these notebooks we construct (1) a model of a patch-clamp experiment with various experimental artefacts, and (2) a model of the corrections applied by patch-clamp amplifiers to mitigate these effects.
 The initial exposition draws on a book chapter by [Sigworth (1995a)](https://doi.org/10.1007/978-1-4419-1229-9_4), but replaces Laplace-domain analysis with an ODE formulation.
 
-To view the notebooks, use the GitHub or nbviewer links below, or clone the repository and run jupyter notebook locally.
+To view the notebooks, use the GitHub or nbviewer links below, or clone the repository and run Jupyter notebook locally.
 Running locally will require the dependencies from `requirements.txt`.
 
 A list of references and further reading is provided [here](./references.ipynb).
@@ -30,8 +30,8 @@ The model is updated to include the compensation circuitry commonly used in patc
 We walk through and simulate the early steps of a manual patch-clamp experiment.
 
 ## Simplified models 
-[![github](./img/github.svg)](./4-simplified.ipynb) 
-[![nbviewer](./img/nbviewer.svg)](https://nbviewer.jupyter.org/github/CardiacModelling/VoltageClampModel-new/tree/main/tutorial/4-simplified.ipynb)
+[![github](./img/github.svg)](./4-simplifications.ipynb) 
+[![nbviewer](./img/nbviewer.svg)](https://nbviewer.jupyter.org/github/CardiacModelling/VoltageClampModel-new/tree/main/tutorial/4-simplifications.ipynb)
 
 In the final notebook, we derive simplified models and compare with previous work.
 
diff --git a/tutorial/tour.ipynb b/tutorial/tour.ipynb
index 6192c3e..ef4bc03 100644
--- a/tutorial/tour.ipynb
+++ b/tutorial/tour.ipynb
@@ -7,10 +7,10 @@
    "source": [
     "# A quick tour of voltage-clamp models\n",
     "\n",
-    "The voltage-clamp models are available in different levels of detail, ranging from the \"full\" model (level 0) to a greatly simplified one (level 5).\n",
+    "The model is available in different levels of detail, ranging from the \"full\" model (level 0) to a greatly simplified one (level 5).\n",
     "Here, we provide a quick tour of the most important models, and provide guidance for setting parameter values.\n",
     "\n",
-    "_This notebook presents, but does not derive or explain the models. For that, please see the tutorial notebooks_."
+    "_This notebook presents, but does not derive or explain the models. For that, please see the Tutorial Notebooks_."
    ]
   },
   {
@@ -26,7 +26,7 @@
     "\n",
     "It includes:\n",
     "\n",
-    "- A voltage offset $E^\\dagger_\\text{off}$, representing the remaining voltage error after zeroing\n",
+    "- A voltage offset $E^\\dagger_\\text{off}$ representing the remaining voltage error after zeroing\n",
     "- A leak current $I_\\text{leak}$\n",
     "- The effect of series resistance $R_s$ on the membrane potential\n",
     "- Series resistance compensation, $\\alpha$, and prediction, $\\beta$, both specified as fractions between 0 and 1."
@@ -53,26 +53,26 @@
    "id": "6d59f32b-7b30-4c3e-b4a2-0484d787cf74",
    "metadata": {},
    "source": [
-    "| Variable        | Units | Meaning                                                   |\n",
-    "|-----------------|-------|-----------------------------------------------------------|\n",
-    "| $V_m$           | mV    | Membrane potential                                        |\n",
-    "| $V_c$           | mV    | The command potential, set by the voltage clamp protocol  |\n",
-    "| $I$             | pA    | The ionic current                                         |\n",
-    "| $I_\\text{obs}$  | pA    | The observed (measured/output) current                    |\n",
-    "| $I_\\text{leak}$ | pA    | A leak current, e.g. $g_\\text{leak}(V_m - I_\\text{leak})$ |\n",
+    "| Variable        | Units | Meaning                                                 |\n",
+    "|-----------------|-------|---------------------------------------------------------|\n",
+    "| $V_m$           | mV    | Membrane potential                                      |\n",
+    "| $V_c$           | mV    | Command potential, set by the voltage clamp protocol    |\n",
+    "| $I$             | pA    | Ionic current                                           |\n",
+    "| $I_\\text{obs}$  | pA    | Observed (measured/output) current                      |\n",
+    "| $I_\\text{leak}$ | pA    | Leak current, e.g. $g_\\text{leak}(V_m - I_\\text{leak})$ |\n",
     "\n",
-    "The variables $I$ and $I_\\text{leak}$ should be supplied by an ionic current model and leak model, respectively.\n",
+    "The variables $I$ and $I_\\text{leak}$ should be supplied by an ionic current model and a leak model, respectively.\n",
     "\n",
     "| Parameter | Units     | Meaning                                             | Typical value   |\n",
     "|-----------|-----------|-----------------------------------------------------|-----------------|\n",
-    "| $C_m$     | pF        | The cell capacitance, from the amplifier estimate   | 25 pF           |\n",
-    "| $R_s$     | G$\\Omega$ | The series resistance, from the amplifier estimate  | 0.005 G$\\Omega$ |\n",
+    "| $C_m$     | pF        | Cell capacitance, from the amplifier estimate       | 25 pF           |\n",
+    "| $R_s$     | G$\\Omega$ | Series resistance, from the amplifier estimate      | 0.005 G$\\Omega$ |\n",
     "| $\\alpha$  | -         | Fraction of $R_s$ compensation, read from amplifier | 0.7             |\n",
-    "| $\\beta$   | -         | Fraction of $R_s$ compensation, read from amplifier | 0.7             |\n",
+    "| $\\beta$   | -         | Fraction of $R_s$ prediction, read from amplifier   | 0.7             |\n",
     "| $E^\\dagger_\\text{off}$ | mV | Remaining voltage offset after zeroing        | 0 mV            |\n",
     "\n",
     "In a simple simulation $E^\\dagger_\\text{off}$ is chosen by the user.\n",
-    "In inference settings, $E^\\dagger_\\text{off}$ may be a parameter inferred from the data."
+    "In inference settings it may be a parameter inferred from the data."
    ]
   },
   {
@@ -93,7 +93,7 @@
     "This level adds\n",
     "\n",
     "- A distinction between estimated and true membrane capacitance and series resistance\n",
-    "- A state-estimator used to approximate $V_e$ in the \"prediction\" pathway"
+    "- A state-estimator used to approximate $V_m$ in the prediction pathway"
    ]
   },
   {
@@ -121,20 +121,20 @@
    "source": [
     "New or changed variables and parameters are\n",
     "\n",
-    "| Variable | Units | Meaning                                               |\n",
-    "|----------|-------|-------------------------------------------------------|\n",
-    "| $V_e$    | mV    | An estimate of $V_m$ used in the \"prediction\" pathway |\n",
+    "| Variable | Units | Meaning                                          |\n",
+    "|----------|-------|--------------------------------------------------|\n",
+    "| $V_e$    | mV    | Estimate of $V_m$ used in the prediction pathway |\n",
     "\n",
     "And\n",
     "\n",
-    "| Parameter | Units     | Meaning                                                 | Typical value   |\n",
-    "|-----------|-----------|---------------------------------------------------------|-----------------|\n",
-    "| $C_m$     | pF        | The true cell capacitance, unknown in real experiments  | 25 pF           |\n",
-    "| $C^*_m$   | pF        | The estimated cell capacitance, read from amplifier     | 25 pF           |\n",
-    "| $R_s$     | G$\\Omega$ | The true series resistance, unknown in real experiments | 0.005 G$\\Omega$ |\n",
-    "| $R^*_s$   | G$\\Omega$ | The series resistance, read from amplifier              | 0.005 G$\\Omega$ |\n",
+    "| Parameter | Units     | Meaning                                          | Typical value   |\n",
+    "|-----------|-----------|--------------------------------------------------|-----------------|\n",
+    "| $C_m$     | pF        | True cell capacitance, unknown                   | 25 pF           |\n",
+    "| $C^*_m$   | pF        | Estimated cell capacitance, read from amplifier  | 25 pF           |\n",
+    "| $R_s$     | G$\\Omega$ | True series resistance, unknown                  | 0.005 G$\\Omega$ |\n",
+    "| $R^*_s$   | G$\\Omega$ | Estimated series resistance, read from amplifier | 0.005 G$\\Omega$ |\n",
     "\n",
-    "For a straightforward simulation, $C_m$ and $R_s$ can be chosen by the user.\n",
+    "For a straightforward simulation, $C_m$ and $R_s$ are chosen by the user.\n",
     "In an inference setting, $C_m$ and $R_s$ may be inferred from the data long with $E^\\dagger_\\text{off}$."
    ]
   },
@@ -197,22 +197,21 @@
    "source": [
     "New variables and parameters are\n",
     "\n",
-    "| Variable | Units | Meaning                                                     |\n",
-    "|----------|-------|-------------------------------------------------------------|\n",
-    "| $V_r$    | mV    | The voltage on one side of the measurement electrode        |\n",
-    "| $V_s$    | mV    | The filtered command potential                              |\n",
-    "| $V_rc$   | mV    | The filtered voltage used in series resistance compensation |\n",
-    "| $V_s$    | mV    | The voltage on one side of the measurement electrode        |\n",
-    "| $I_1$    | pA    | The output current before filtering                         |\n",
+    "| Variable      | Units | Meaning                                     |\n",
+    "|---------------|-------|---------------------------------------------|\n",
+    "| $V_r$         | mV    | Reference voltage set by the voltage clamp  |\n",
+    "| $V_s$         | mV    | Filtered command potential                  |\n",
+    "| $V_\\text{rc}$ | mV    | Filtered voltage used in $R_s$ compensation |\n",
+    "| $I_1$         | pA    | Unfiltered output current                   |\n",
     "\n",
     "And\n",
     "\n",
-    "| Parameter        | Unit      | Meaning                                                    | Typical value |\n",
-    "|------------------|-----------|------------------------------------------------------------|---------------|\n",
-    "| $\\tau_\\text{RC}$ | ms        | Time constant of filter in series resistance compenstation, read from amplifier|0.01 ms|\n",
-    "| $\\tau_s$         | ms        | Time constant of stimlus filter, estimated from fits       | 0.025 ms      |\n",
-    "| $\\tau_o$         | ms        | Time constant of 1st order approximation of output filter  | 0.01 ms       |\n",
-    "| $R_f$            | G$\\Omega$ | The measurement resistor, found in headstage documentation | 0.5 G$\\Omega$ |\n",
+    "| Parameter        | Units     | Meaning                                            | Typical value |\n",
+    "|------------------|-----------|----------------------------------------------------|---------------|\n",
+    "| $\\tau_\\text{RC}$ | ms        | Time constant of filter in $R_s$ compenstation, read from amplifier|0.01 ms|\n",
+    "| $\\tau_s$         | ms        | Time constant of stimulus filter, unknown          | 0.025 ms      |\n",
+    "| $\\tau_o$         | ms        | Time constant of output filter, see below          | 0.01 ms       |\n",
+    "| $R_f$            | G$\\Omega$ | Measurement resistor, from headstage documentation | 0.5 G$\\Omega$ |\n",
     "\n",
     "To estimate $\\tau_s$ in a real amplifier, you can record the filtered output voltage and fit to it.\n",
     "The value above corresponds to the \"slow\" (default) setting on a HEKA EPC-10.\n",
@@ -235,7 +234,7 @@
     "\n",
     "This level adds\n",
     "\n",
-    "- A pipette (conventional patch clamp) or parasitic (planar patch clamp) capacitance $C_p$, and its amplifier estimate $C^*p$\n",
+    "- A pipette (conventional patch clamp) or parasitic (planar patch clamp) capacitance $C_p$, and its amplifier estimate $C^*_p$\n",
     "- A stray capacitance on the measurement resistor $\\tilde{C}_f$, and a finite op-amp speed with time constant $\\tau_a$\n",
     "- A two-part output filter, where \"Filter 1\" affects series resistance compensation, while \"Filter 2\" does not.\n",
     "- Arbitrary equations for the output filters and the stimulus filter (allowing e.g. Bessel formulations)"
@@ -293,42 +292,27 @@
    "id": "7cc410ae-a332-4fdd-9f04-459790d43ef2",
    "metadata": {},
    "source": [
-    "# TODO\n",
-    "\n",
     "New variables and parameters are\n",
     "\n",
-    "| Variable | Units | Meaning                                                     |\n",
-    "|----------|-------|-------------------------------------------------------------|\n",
-    "| $V_r$    | mV    | The voltage on one side of the measurement electrode        |\n",
-    "| $V_s$    | mV    | The filtered command potential                              |\n",
-    "| $V_rc$   | mV    | The filtered voltage used in series resistance compensation |\n",
-    "| $V_s$    | mV    | The voltage on one side of the measurement electrode        |\n",
-    "| $I_1$    | pA    | The output current before filtering                         |\n",
+    "| Variable | Units | Meaning                |\n",
+    "|----------|-------|------------------------|\n",
+    "| $V_o$    | mV    | Op-amp output voltage  |\n",
+    "| $V_p$    | mV    | Pipette potential      |\n",
+    "| $V_1$    | mV    | Voltage after filter 1 |\n",
+    "| $V_2$    | mV    | Voltage after filter 2 |\n",
     "\n",
-    "And\n",
     "\n",
-    "| Parameter        | Unit      | Meaning                                                    | Typical value |\n",
-    "|------------------|-----------|------------------------------------------------------------|---------------|\n",
-    "| $\\tau_\\text{RC}$ | ms        | Time constant of filter in series resistance compenstation, read from amplifier|0.01 ms|\n",
-    "| $\\tau_s$         | ms        | Time constant of stimlus filter, estimated from fits       | 0.025 ms      |\n",
-    "| $\\tau_o$         | ms        | Time constant of 1st order approximation of output filter  | 0.01 ms       |\n",
-    "| $R_f$            | G$\\Omega$ | The measurement resistor, found in headstage documentation | 0.5 G$\\Omega$ |\n",
+    "And\n",
     "\n",
-    "To estimate $\\tau_s$ in a real amplifier, you can record the filtered output voltage and fit to it.\n",
-    "The value above corresponds to the \"slow\" (default) setting on a HEKA EPC-10.\n",
+    "| Parameter     | Units | Meaning                                            | Typical value |\n",
+    "|---------------|-------|----------------------------------------------------|---------------|\n",
+    "| $C_p$         | pF    | True pipette capacitance, unknown                  | 5 pF          |\n",
+    "| $C^*_p$       | pF    | Estimated pipette capacitance, read from amplifier | 5 pF          |\n",
+    "| $\\tilde{C}_f$ | pF    | Compensated stray capacitance on measuring resistor, unknown | 5e-3 pF |\n",
+    "| $\\tau_a$      | ms    | Time constant of measurement op-amp, unknown       | 13e-6 ms      |\n",
     "\n",
-    "To estimate $\\tau_o$, you can use $\\tau_o \\approx \\frac{1}{2 \\pi f}$, where $f$ is the filter's cut-off frequency in kHz.\n",
-    "\n",
-    "Note that exact values of $\\tau_s$ and $\\tau_o$ are seldom crucial."
+    "The given values of $\\tilde{C}_f$ and $\\tau_a$ were estimated by fitting to model cell experiments with a Heka EPC-10."
    ]
-  },
-  {
-   "cell_type": "code",
-   "execution_count": null,
-   "id": "2332084c-5e5d-4dc1-aed9-fdc3019ffe8b",
-   "metadata": {},
-   "outputs": [],
-   "source": []
   }
  ],
  "metadata": {