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"illumination": "11 mW/mm²", "intensityMWmm2": 11, "t75Sec": 64, "t50Sec": 155, "extrapolated": false, "source": { "label": "Chamberland 2017, Fig. 1—figure supplement 3A", "url": "https://doi.org/10.7554/eLife.25690" } }, { "modality": "1P", "illumination": "18 mW/mm²", "intensityMWmm2": 18, "t75Sec": 42, "t50Sec": null, "extrapolated": false, "source": { "label": "Platisa 2017, Fig. 3C", "url": "https://doi.org/10.1021/acschemneuro.6b00234" } }, { "modality": "1P", "illumination": "100 mW/mm²", "intensityMWmm2": 100, "t75Sec": 56, "t50Sec": null, "extrapolated": false, "source": { "label": "Abdelfattah 2016, Fig. 3I", "url": "https://doi.org/10.1523/JNEUROSCI.3484-15.2016" } } ] }, "t75ScaledEntriesSec": [ 1.77, 7.04, 7.56, 56 ], "fvCurve": { "voltageMV": [ -100, -70, -40, 30, 100 ], "dffPercent": [ 16, 0, -12, -30, -41 ], "source": { "label": "Milosevic 2020, Fig. 2C", "url": "https://doi.org/10.1523/ENEURO.0060-20.2020" } }, "researchPapers": [ { "title": "Single action potentials and subthreshold electrical events imaged in neurons with a fluorescent protein voltage probe", "authors": "Jin L, Han Z, Platisa J, Wooltorton JRA, Cohen LB, Pieribone VA", "journal": "Neuron", "year": 2012, "sample": "HEK293 cells, cultured mouse hippocampal neurons, in vitro", "url": "https://doi.org/10.1016/j.neuron.2012.06.040" }, { "title": "Genetically targeted optical electrophysiology in intact neural circuits", "authors": "Cao G, et al.", "journal": "Cell", "year": 2013, "sample": "Drosophila brain neurons, in vivo", "url": "https://doi.org/10.1016/j.cell.2013.07.027" }, { "title": "Fluorescent protein voltage probes derived from ArcLight that respond to membrane voltage changes with fast kinetics", "authors": "Han Z, et al.", "journal": "PLoS One", "year": 2013, "sample": "HEK293 cells and cultured rat hippocampal neurons, in vitro", "url": "https://doi.org/10.1371/journal.pone.0081295" }, { "title": "Rapid cellular phenotyping of human pluripotent stem cell-derived cardiomyocytes using a genetically encoded fluorescent voltage sensor", "authors": "Leyton-Mange JS, et al.", "journal": "Stem Cell Reports", "year": 2014, "sample": "Human iPSC-derived cardiomyocytes, in vitro", "url": "https://doi.org/10.1016/j.stemcr.2014.01.003" }, { "title": "Imaging neural spiking in brain tissue using FRET-opsin protein voltage sensors", "authors": "Gong Y, et al.", "journal": "Nature Communications", "year": 2014, "sample": "Cultured neurons, brain slices, mouse cerebellum in vivo", "url": "https://doi.org/10.1038/ncomms4674" }, { "title": "Mechanistic studies of the genetically encoded fluorescent protein voltage probe ArcLight", "authors": "Han Z, et al.", "journal": "PLoS One", "year": 2014, "sample": "HEK293 cells, in vitro", "url": "https://doi.org/10.1371/journal.pone.0113873" }, { "title": "Monitoring brain activity with protein voltage and calcium sensors", "authors": "Storace DA, et al.", "journal": "Scientific Reports", "year": 2015, "sample": "Mouse olfactory bulb, in vivo", "url": "https://doi.org/10.1038/srep10212" }, { "title": "Two-photon lifetime imaging of voltage indicating proteins as a probe of absolute membrane voltage", "authors": "Brinks D, et al.", "journal": "Biophysical Journal", "year": 2015, "sample": "HEK293 cells, in vitro", "url": "https://doi.org/10.1016/j.bpj.2015.07.038" }, { "title": "Monitoring human-induced pluripotent stem cell-derived cardiomyocytes with genetically encoded calcium and voltage fluorescent reporters", "authors": "Shinnawi R, et al.", "journal": "Stem Cell Reports", "year": 2015, "sample": "Human iPSC-derived cardiomyocytes, in vitro", "url": "https://doi.org/10.1016/j.stemcr.2015.08.009" }, { "title": "Dual optical recordings for action potentials and calcium handling in induced pluripotent stem cell models of cardiac arrhythmias", "authors": "Song L, et al.", "journal": "Stem Cells Translational Medicine", "year": 2015, "sample": "Human iPSC-derived cardiomyocytes (Timothy syndrome), in vitro", "url": "https://doi.org/10.5966/sctm.2014-0245" }, { "title": "Expression and testing in plants of ArcLight, a genetically-encoded voltage indicator used in neuroscience research", "authors": "Matzke AJ, et al.", "journal": "BMC Plant Biology", "year": 2015, "sample": "Nicotiana benthamiana and Arabidopsis thaliana, in vivo", "url": "https://doi.org/10.1186/s12870-015-0633-z" }, { "title": "Single-molecule fluorimetry and gating currents inspire an improved optical voltage indicator (ArcLightning)", "authors": "Treger JS, et al.", "journal": "eLife", "year": 2015, "sample": "HEK293 cells, Xenopus oocytes, in vitro", "url": "https://doi.org/10.7554/eLife.10482" }, { "title": "Ziram, a pesticide associated with increased risk for Parkinson's disease, differentially affects the presynaptic function of aminergic and glutamatergic nerve terminals", "authors": "Martin CA, et al.", "journal": "Experimental Neurology", "year": 2016, "sample": "Drosophila neuromuscular junction, ex vivo", "url": "https://doi.org/10.1016/j.expneurol.2015.09.017" }, { "title": "Optogenetic monitoring of synaptic activity with genetically encoded voltage indicators", "authors": "Nakajima R, et al.", "journal": "Frontiers in Synaptic Neuroscience", "year": 2016, "sample": "Mouse olfactory bulb brain slices and in vivo", "url": "https://doi.org/10.3389/fnsyn.2016.00022" }, { "title": "Directed evolution of key residues in fluorescent protein inverses the polarity of voltage sensitivity in ArcLight (Marina)", "authors": "Platisa J, et al.", "journal": "ACS Chemical Neuroscience", "year": 2017, "sample": "HEK293 cells, in vitro", "url": "https://doi.org/10.1021/acschemneuro.6b00234" }, { "title": "Genetically expressed voltage sensor ArcLight for imaging large scale cortical activity in the anesthetized and awake mouse", "authors": "Borden PY, et al.", "journal": "Neurophotonics", "year": 2017, "sample": "Mouse somatosensory cortex (barrel cortex), in vivo", "url": "https://doi.org/10.1117/1.NPh.4.3.031212" }, { "title": "Attenuation of synaptic potentials in dendritic spines", "authors": "Kwon T, et al.", "journal": "Cell Reports", "year": 2017, "sample": "Cultured rat hippocampal neurons, in vitro (two-photon)", "url": "https://doi.org/10.1016/j.celrep.2017.07.012" }, { "title": "Improving a genetically encoded voltage indicator by modifying the cytoplasmic charge composition (Bongwoori-R3)", "authors": "Lee S, et al.", "journal": "Scientific Reports", "year": 2017, "sample": "HEK293 cells and cultured rat hippocampal neurons, in vitro", "url": "https://doi.org/10.1038/s41598-017-08731-2" }, { "title": "MicroRNA-92a is a circadian modulator of neuronal excitability in Drosophila", "authors": "Chen X, et al.", "journal": "Nature Communications", "year": 2017, "sample": "Drosophila PDF pacemaker neurons, in vivo", "url": "https://doi.org/10.1038/ncomms14707" }, { "title": "Human induced pluripotent stem cell-derived cardiac cell sheets expressing genetically encoded voltage indicator for pharmacological and arrhythmia studies", "authors": "Shaheen N, et al.", "journal": "Stem Cell Reports", "year": 2018, "sample": "Human iPSC-derived cardiac cell sheets, in vitro", "url": "https://doi.org/10.1016/j.stemcr.2018.04.006" }, { "title": "A dimeric fluorescent protein yields a bright, red-shifted GEVI capable of population signals in brain slice (Ilmol)", "authors": "Yi B, et al.", "journal": "Scientific Reports", "year": 2018, "sample": "HEK293 cells, cultured neurons, mouse hippocampal brain slices", "url": "https://doi.org/10.1038/s41598-018-33297-y" }, { "title": "Mapping of excitatory and inhibitory postsynaptic potentials of neuronal populations in hippocampal slices using the GEVI, ArcLight", "authors": "Nakajima R, et al.", "journal": "Journal of Physics D: Applied Physics", "year": 2018, "sample": "Mouse hippocampal CA1 brain slices, ex vivo", "url": "https://doi.org/10.1088/1361-6463/aae2e3" }, { "title": "Optical consequences of a genetically-encoded voltage indicator with a pH sensitive fluorescent protein", "authors": "Kang BE, et al.", "journal": "Neuroscience Research", "year": 2019, "sample": "HEK293 cells and cultured hippocampal neurons, in vitro", "url": "https://doi.org/10.1016/j.neures.2018.10.006" }, { "title": "Comparative evaluation of genetically encoded voltage indicators", "authors": "Bando Y, et al.", "journal": "Cell Reports", "year": 2019, "sample": "HEK293 cells, cultured neurons, mouse visual cortex in vivo", "url": "https://doi.org/10.1016/j.celrep.2018.12.088" }, { "title": "Using genetically encoded voltage indicators (GEVIs) to study the input-output transformation of the mammalian olfactory bulb", "authors": "Storace DA, et al.", "journal": "Frontiers in Cellular Neuroscience", "year": 2019, "sample": "Mouse olfactory bulb, in vivo", "url": "https://doi.org/10.3389/fncel.2019.00342" }, { "title": "Single-cell RNA-sequencing and optical electrophysiology of human iPSC-derived cardiomyocytes reveal discordance between cardiac subtype-associated gene expression and electrophysiological phenotypes", "authors": "Biendarra-Tiegs SM, et al.", "journal": "Stem Cells and Development", "year": 2019, "sample": "Human iPSC-derived cardiomyocytes, in vitro", "url": "https://doi.org/10.1089/scd.2019.0030" }, { "title": "Characterization of the CACNA1C-R518C missense mutation in the pathobiology of long-QT syndrome using human iPSC cardiomyocytes", "authors": "Estes SI, et al.", "journal": "Circulation: Genomic and Precision Medicine", "year": 2019, "sample": "Human iPSC-derived cardiomyocytes (long-QT syndrome), in vitro", "url": "https://doi.org/10.1161/CIRCGEN.119.002534" }, { "title": "Human induced pluripotent stem cell line with genetically encoded fluorescent voltage indicator generated via CRISPR", "authors": "Sun YH, et al.", "journal": "Stem Cells", "year": 2020, "sample": "Human iPSC-derived cardiomyocytes (CRISPR knock-in), in vitro", "url": "https://doi.org/10.1002/stem.3085" }, { "title": "Human iPSC-derived non-cardiomyocytes modulate cardiac electrophysiological maturation through connexin 43-mediated cell-cell interactions", "authors": "Biendarra-Tiegs SM, et al.", "journal": "Stem Cells and Development", "year": 2020, "sample": "Human iPSC-derived cardiomyocytes and non-cardiomyocytes co-cultures, in vitro", "url": "https://doi.org/10.1089/scd.2019.0098" }, { "title": "Real-time detection of changes in yeast plasma membrane potential using genetically encoded voltage indicator proteins", "authors": "Limapichat W, et al.", "journal": "FEMS Yeast Research", "year": 2020, "sample": "Saccharomyces cerevisiae (yeast), in vitro", "url": "https://doi.org/10.1093/femsyr/foaa041" }, { "title": "In vitro testing of voltage indicators: Archon1, ArcLightD, ASAP1, ASAP2s, ASAP3b, Bongwoori-Pos6, BeRST1, FlicR1, and Chi-VSFP-Butterfly", "authors": "Milosevic MM, et al.", "journal": "eNeuro", "year": 2020, "sample": "HEK293 cells, cultured neurons, mouse brain slices", "url": "https://doi.org/10.1523/ENEURO.0060-20.2020" }, { "title": "Triclosan disrupts immune cell function by depressing Ca2+ influx following acidification of the cytoplasm", "authors": "Sangroula S, et al.", "journal": "Toxicology and Applied Pharmacology", "year": 2020, "sample": "RBL-2H3 mast cells and Jurkat T cells, in vitro", "url": "https://doi.org/10.1016/j.taap.2020.115205" }, { "title": "Analysing bioelectrical phenomena in the Drosophila ovary with genetic tools", "authors": "Schotthoefer SK, et al.", "journal": "BMC Developmental Biology", "year": 2020, "sample": "Drosophila melanogaster ovary follicular epithelium, ex vivo", "url": "https://doi.org/10.1186/s12861-020-00220-6" }, { "title": "Promise and potential peril with lumacaftor for the trafficking defective type 2 long-QT syndrome-causative variants", "authors": "O'Hare BJ, et al.", "journal": "Circulation: Genomic and Precision Medicine", "year": 2020, "sample": "Human iPSC-derived cardiomyocytes (LQT2), in vitro", "url": "https://doi.org/10.1161/CIRCGEN.120.002950" }, { "title": "GEVI cell-type specific labelling and a manifold learning approach provide evidence for lateral inhibition at the population level in mouse hippocampal CA1", "authors": "Nakajima R, et al.", "journal": "European Journal of Neuroscience", "year": 2021, "sample": "Mouse hippocampal CA1 brain slices, ex vivo", "url": "https://doi.org/10.1111/ejn.15177" }, { "title": "Improving the flexibility of genetically encoded voltage indicators via intermolecular FRET", "authors": "Leong LM, et al.", "journal": "Biophysical Journal", "year": 2021, "sample": "HEK293 cells, in vitro", "url": "https://doi.org/10.1016/j.bpj.2021.03.010" }, { "title": "Mechanism of ArcLight derived GEVIs involves electrostatic interactions that can affect proton wires", "authors": "Kang BE, et al.", "journal": "Biophysical Journal", "year": 2021, "sample": "HEK293 cells, in vitro", "url": "https://doi.org/10.1016/j.bpj.2021.03.009" }, { "title": "Enabling comprehensive optogenetic studies of mouse hearts by simultaneous opto-electrical panoramic mapping and stimulation", "authors": "Rieger M, et al.", "journal": "Nature Communications", "year": 2021, "sample": "Transgenic mouse hearts (ArcLight-Q239), ex vivo", "url": "https://doi.org/10.1038/s41467-021-26039-8" }, { "title": "Studying synaptically evoked cortical responses ex vivo with combination of single neuron recording and population voltage imaging", "authors": "Jang J, et al.", "journal": "Frontiers in Neuroscience", "year": 2021, "sample": "Mouse cortex brain slices, ex vivo", "url": "https://doi.org/10.3389/fnins.2021.773883" }, { "title": "Visualizing oscillations in brain slices with genetically encoded voltage indicators", "authors": "Rhee JK, et al.", "journal": "Frontiers in Neuroanatomy", "year": 2021, "sample": "Mouse motor cortex brain slices, ex vivo", "url": "https://doi.org/10.3389/fnana.2021.741711" }, { "title": "Simultaneous two-photon imaging of action potentials and subthreshold inputs in vivo", "authors": "Bando Y, et al.", "journal": "Nature Communications", "year": 2021, "sample": "Mouse visual cortex neurons (ArcLight-ST variant), in vivo", "url": "https://doi.org/10.1038/s41467-021-27444-9" }, { "title": "Voltage imaging in the olfactory bulb using transgenic mouse lines expressing ArcLight", "authors": "Platisa J, et al.", "journal": "Scientific Reports", "year": 2022, "sample": "Transgenic mouse olfactory bulb, in vivo", "url": "https://doi.org/10.1038/s41598-021-04482-3" }, { "title": "Conserved amino acids residing outside the voltage field can shift the voltage sensitivity and increase the signal speed and size of Ciona based GEVIs", "authors": "Sepehri Rad M, et al.", "journal": "Frontiers in Cell and Developmental Biology", "year": 2022, "sample": "HEK293 cells, in vitro", "url": "https://doi.org/10.3389/fcell.2022.868143" }, { "title": "Recombinant adeno-associated viral vector-mediated gene transfer of hTBX18 generates pacemaker cells from ventricular cardiomyocytes", "authors": "Farraha M, et al.", "journal": "International Journal of Molecular Sciences", "year": 2022, "sample": "Neonatal rat ventricular cardiomyocytes, in vitro", "url": "https://doi.org/10.3390/ijms23169230" }, { "title": "Generation of human iPSC line encoding for ArcLight A242", "authors": "Vivekanandan R, et al.", "journal": "Stem Cell Research", "year": 2023, "sample": "Human iPSC line with knock-in ArcLight A242, in vitro", "url": "https://doi.org/10.1016/j.scr.2022.102981" }, { "title": "Voltage to calcium transformation enhances direction selectivity in Drosophila T4 neurons", "authors": "Mishra A, et al.", "journal": "Journal of Neuroscience", "year": 2023, "sample": "Drosophila T4 direction-selective neurons, in vivo (two-photon)", "url": "https://doi.org/10.1523/JNEUROSCI.2297-22.2023" }, { "title": "Fast two-photon imaging of subcellular voltage dynamics in neuronal tissue with genetically encoded indicators", "authors": "Chamberland S, et al.", "journal": "eLife", "year": 2017, "sample": "Drosophila visual neurons (L2), in vivo; HEK293A cells, in vitro", "url": "https://doi.org/10.7554/eLife.25690" }, { "title": "Imaging different cell populations in the mouse olfactory bulb using ArcLight", "authors": "Leong LM, et al.", "journal": "Neurophotonics", "year": 2024, "sample": "Mouse olfactory bulb (multiple cell types), in vivo", "url": "https://doi.org/10.1117/1.NPh.11.3.033402" }, { "title": "Generation of a genetically encoded voltage indicator MARINA reporter human iPS cell line using Cas9", "authors": "Petruskeviciu-te A, et al.", "journal": "Stem Cell Research", "year": 2025, "sample": "Human iPSC line with knock-in MARINA, in vitro", "url": "https://doi.org/10.1016/j.scr.2024.103628" }, { "title": "Ultrafast two-photon imaging of a high-gain voltage indicator in awake behaving mice", "authors": "Villette V, et al.", "journal": "Cell", "year": 2019, "sample": "HEK293 cells, cultured neurons, mouse visual cortex in vivo", "url": "https://doi.org/10.1016/j.cell.2019.11.004" }, { "title": "All-optical electrophysiology in mammalian neurons using engineered microbial rhodopsins", "authors": "Hochbaum DR, et al.", "journal": "Nature Methods", "year": 2014, "sample": "HEK293T cells, cultured rat hippocampal neurons, human iPSC-derived neurons", "url": "https://doi.org/10.1038/nmeth.3000" }, { "title": "High-fidelity optical reporting of neuronal electrical activity with an ultrafast fluorescent voltage sensor", "authors": "St-Pierre F, et al.", "journal": "Nature Neuroscience", "year": 2014, "sample": "HEK293A cells, cultured rat hippocampal neurons, mouse brain slices, in vivo", "url": "https://doi.org/10.1038/nn.3709" }, { "title": "A bright and fast red fluorescent protein voltage indicator that reports neuronal activity in organotypic brain slices", "authors": "Abdelfattah AS, et al.", "journal": "Journal of Neuroscience", "year": 2016, "sample": "HEK293 cells, organotypic rat brain slices", "url": "https://doi.org/10.1523/JNEUROSCI.3484-15.2016" } ] }, { "id": "asap1", "name": "ASAP1", "url": "https://gevibench.org/gevi/asap1", "description": "First ASAP sensor, establishing the VSD/cpGFP architecture for fast, negative-going fluorescence responses to voltage.", "category": "VSD-cpGFP GEVI", "tags": [ "cpGFP" ], "year": 2014, "date": "2014-04-22", "paper": "Nature Neuroscience 2014", "paperUrl": "https://doi.org/10.1038/nn.3709", "parent": null, "crossBranchParent": null, "lineagePath": [ "GEVI", "VSD", "VSD-single", "asap1" ], "polarity": "negative", "spectrum": { "peakExcitationNm": 490, "peakEmissionNm": 510, "fluorophore": "ASAP1" }, "twoPhotonCompatible": true, "bioluminescent": false, "addgene": { "id": "52519", "url": "https://www.addgene.org/52519/", "note": null }, "lastUpdated": "2026-09-19", "display": { "brightness": "0.43×", "kinetics": "6.7 / 12", "dynamicRange": "-17.5%", "subthreshold": "0.21%", "sensitivity": "6.3%", "apWidth": "6.2 ms", "photostability": "11 s", "papers": "21" }, "values": { "brightnessRelEGFP": 0.4271, "tauOnMs": 6.7, "tauOffMs": 11.6, "dffPer100mV": -17.5, "dffPer100mVMedianAbs": 24.5, "dffPerMV": 0.21, "dffPerMVEstimatedFromFV": false, "dffPerAP": 6.3, "dffPerAPMedianAbs": 7.45, "apWidthMs": 6.15, "t75At100mWmm2Sec": 10.86, "papersUsingIt": 21 }, "measurements": { "kinetics": [ { "tauOnMs": 29.7, "tauOffMs": 29.5, "temperature": "22°C", "dye": null, "source": { "label": "St-Pierre 2014, Fig. 1e", "url": "https://doi.org/10.1038/nn.3709" } }, { "tauOnMs": 44, "tauOffMs": 66.9, "temperature": "22°C", "dye": null, "source": { "label": "Chamberland 2017, Table 1", "url": "https://doi.org/10.7554/eLife.25690" } }, { "tauOnMs": 6.7, "tauOffMs": 11.6, "temperature": "23°C", "dye": null, "source": { "label": "Zou 2014, Table 1", "url": "https://doi.org/10.1038/ncomms5625" } }, { "tauOnMs": 23.3, "tauOffMs": 36.4, "temperature": "23°C", "dye": null, "source": { "label": "Villette 2019, Table S1", "url": "https://doi.org/10.1016/j.cell.2019.11.004" } } ], "dffPer100mV": [ { "dff": -17.5, "responseType": "steady-state", "modality": "1P", "dye": null, "source": { "label": "St-Pierre 2014, Fig. 1d", "url": "https://doi.org/10.1038/nn.3709" } }, { "dff": -23.3, "responseType": "steady-state", "modality": "1P", "dye": null, "source": { "label": "Chamberland 2017, Fig. 1C", "url": "https://doi.org/10.7554/eLife.25690" } }, { "dff": -25.7, "responseType": "steady-state", "modality": "1P", "dye": null, "source": { "label": "Milosevic 2020, Fig. 2D", "url": "https://doi.org/10.1523/ENEURO.0060-20.2020" } }, { "dff": -28.8, "responseType": "steady-state", "modality": "1P", "dye": null, "source": { "label": "Zou 2014, Table 1", "url": "https://doi.org/10.1038/ncomms5625" } } ], "dffPerMV": [ { "slopePercentPerMV": 0.21, "modality": "1P", "note": "Subthreshold voltage sensitivity ≈ 0.21 %/mV, derived from the Fig. 3b (top) quantification in voltage-clamped cultured rat hippocampal neurons: ASAP1 peak −ΔF/F was 1.20 / 2.05 / 2.85 / 4.52% for simulated subthreshold depolarization waveforms of 5 / 10 / 15 / 20 mV peak amplitude (17 ms peak FWHM, n = 6 neurons, 1P widefield, 22°C). Through-origin least squares over the four points gives 0.213 %/mV (free-intercept fit gives 0.215 %/mV with a ~0 intercept, i.e. the response is linear over this range). Read off the bar graph, so ±~5%. Comparable steady-state chord slope from the HEK293A F−V curve (Fig. 1d, −80 → −60 mV) is 0.31 %/mV; the neuronal waveform value is lower because these are brief transient responses, not steady-state steps. Cross-GEVI: ArcLight Q239 in the same panel gives ≈ 0.13 %/mV (0.80 / 1.63 / 2.12 / 2.28%).", "source": { "label": "St-Pierre 2014, Fig. 3b", "url": "https://doi.org/10.1038/nn.3709" } } ], "dffPerAP": [ { "dff": 6.3, "modality": "1P", "dye": null, "source": { "label": "St-Pierre 2014, Fig. 4a", "url": "https://doi.org/10.1038/nn.3709" } }, { "dff": 8.6, "modality": "1P", "dye": null, "source": { "label": "Chamberland 2017, Fig. 2I", "url": "https://doi.org/10.7554/eLife.25690" } }, { "dff": 2.8, "modality": "1P", "dye": null, "source": { "label": "Lee 2017, Table 3", "url": "https://doi.org/10.1038/s41598-017-08731-2" } }, { "dff": 17.8, "modality": "1P", "dye": null, "source": { "label": "Zou 2014, Table 1", "url": "https://doi.org/10.1038/ncomms5625" } } ], "apWidth": [ { "fwhmMs": 3.7, "samplingRateHz": 827, "sample": "Rat cultured hippocampal neurons, current-triggered single AP, in vitro", "modality": "1P", "temperature": "22°C", "source": { "label": "St-Pierre 2014, Fig. 1f", "url": "https://doi.org/10.1038/nn.3709" } }, { "fwhmMs": 3.8, "samplingRateHz": 3700, "sample": "Organotypic hippocampal slice cultures, current-triggered single AP, two-photon RAMP (FEVIR)", "modality": "2P", "temperature": "22°C", "source": { "label": "Chamberland 2017, Fig. 6E", "url": "https://doi.org/10.7554/eLife.25690" } }, { "fwhmMs": 9, "samplingRateHz": 1000, "sample": "Rat, cultured hippocampal neurons, in vitro", "modality": "1P", "temperature": null, "source": { "label": "Chamberland 2017, Fig. 2J", "url": "https://doi.org/10.7554/eLife.25690" } }, { "fwhmMs": 8.5, "samplingRateHz": 500, "sample": "Mouse, cultured primary cortical/hippocampal neurons (P0.5), in vitro", "modality": "1P", "temperature": null, "source": { "label": "Milosevic 2020, Fig. 1A", "url": "https://doi.org/10.1523/ENEURO.0060-20.2020" } } ], "brightness": [ { "ratio": 1, "reference": "ASAP2s", "source": { "label": "Chamberland 2017, Fig. S2", "url": "https://doi.org/10.7554/eLife.25690" } }, { "ratio": 0.61, "reference": "ArcLight", "source": { "label": "Milosevic 2020, Fig. 2F", "url": "https://doi.org/10.1523/ENEURO.0060-20.2020" } }, { "ratio": 0.94, "reference": "ASAP3", "source": { "label": "Nair 2025, Fig. 2C", "url": "https://doi.org/10.1021/acschemneuro.5c00670" } } ], "photobleaching": [ { "modality": "1P", "illumination": "0.12 mW/mm²", "intensityMWmm2": null, "t75Sec": 508, "t50Sec": null, "extrapolated": false, "source": { "label": "St-Pierre 2014, Fig. S9", "url": "https://doi.org/10.1038/nn.3709" } }, { "modality": "1P", "illumination": "11 mW/mm²", "intensityMWmm2": 11, "t75Sec": 52, "t50Sec": 150, "extrapolated": false, "source": { "label": "Chamberland 2017, Fig. 1—figure supplement 3A", "url": "https://doi.org/10.7554/eLife.25690" } }, { "modality": "1P", "illumination": "100 mW/mm²", "intensityMWmm2": 100, "t75Sec": 16, "t50Sec": null, "extrapolated": false, "source": { "label": "Abdelfattah 2016, Fig. 3I", "url": "https://doi.org/10.1523/JNEUROSCI.3484-15.2016" } }, { "modality": "2P", "illumination": "~1 mW (950 nm, 2P point scan)", "intensityMWmm2": null, "t75Sec": null, "t50Sec": 0.84, "extrapolated": false, "source": { "label": "Abdelfattah 2016, Fig. 4C", "url": "https://doi.org/10.1523/JNEUROSCI.3484-15.2016" } } ] }, "t75ScaledEntriesSec": [ 5.72, 16 ], "fvCurve": { "voltageMV": [ -120, -100, -80, -70, -60, -40, -20, 0, 30, 50 ], "dffPercent": [ 23.4, 11.7, 3.3, 0, -2.8, -6.7, -9.8, -12.8, -17.5, -21.1 ], "source": { "label": "St-Pierre 2014, Fig. 1d", "url": "https://doi.org/10.1038/nn.3709" } }, "researchPapers": [ { "title": "High-fidelity optical reporting of neuronal electrical activity with an ultrafast fluorescent voltage sensor", "authors": "St-Pierre F, Marshall JD, Yang Y, Gong Y, Schnitzer MJ, Lin MZ", "journal": "Nature Neuroscience", "year": 2014, "sample": "HEK293A cells; rat hippocampal neurons, in vitro", "url": "https://doi.org/10.1038/nn.3709" }, { "title": "A Bright and Fast Red Fluorescent Protein Voltage Indicator That Reports Neuronal Activity in Organotypic Brain Slices", "authors": "Abdelfattah AS, Farhi SL, Zhao Y, Brinks D, Zou P, Ruangkittisakul A, Platisa J, Pieribone VA, Ballanyi K, Cohen AE, Campbell RE", "journal": "Journal of Neuroscience", "year": 2016, "sample": "HEK293 cells; rat hippocampal neurons, in vitro", "url": "https://doi.org/10.1523/JNEUROSCI.3484-15.2016" }, { "title": "Biophysical Characterization of Genetically Encoded Voltage Sensor ASAP1: Dynamic Range Improvement", "authors": "Lee EEL, Bezanilla F", "journal": "Biophysical Journal", "year": 2017, "sample": "Xenopus laevis oocytes; rat dorsal root ganglion neurons, in vitro", "url": "https://doi.org/10.1016/j.bpj.2017.10.018" }, { "title": "Fast two-photon imaging of subcellular voltage dynamics in neuronal tissue with genetically encoded indicators", "authors": "Chamberland S, et al.", "journal": "eLife", "year": 2017, "sample": "HEK293A cells; mouse hippocampal neurons, in vitro", "url": "https://doi.org/10.7554/eLife.25690" }, { "title": "In Vitro Testing of Voltage Indicators: Archon1, ArcLightD, ASAP1, ASAP2s, ASAP3b, Bongwoori-Pos6, BeRST1, FlicR1, and Chi-VSFP-Butterfly", "authors": "Milosevic MM, et al.", "journal": "eNeuro", "year": 2020, "sample": "HEK293 cells; mouse brain slices, in vitro", "url": "https://doi.org/10.1523/ENEURO.0060-20.2020" }, { "title": "Comparative Evaluation of Genetically Encoded Voltage Indicators", "authors": "Bando Y, et al.", "journal": "Cell Reports", "year": 2019, "sample": "Mouse hippocampal neurons, in vitro", "url": "https://doi.org/10.1016/j.celrep.2018.12.088" }, { "title": "Optical interrogation of neuronal circuitry in zebrafish using genetically encoded voltage indicators", "authors": "Miyazawa H, et al.", "journal": "Scientific Reports", "year": 2018, "sample": "Zebrafish, cerebellum; spinal cord, in vivo", "url": "https://doi.org/10.1038/s41598-018-23906-1" }, { "title": "Visualization of Cellular Electrical Activity in Zebrafish Early Embryos and Tumors", "authors": "Silic MR, Zhang G", "journal": "Journal of Visualized Experiments", "year": 2018, "sample": "Zebrafish embryos; zebrafish tumors, in vivo", "url": "https://doi.org/10.3791/57330" }, { "title": "Real-time detection of yeast membrane potential using ASAP1 voltage indicator", "authors": "Limapichat W, et al.", "journal": "FEMS Yeast Research", "year": 2020, "sample": "Saccharomyces cerevisiae, in vitro", "url": "https://doi.org/10.1093/femsyr/foaa041" }, { "title": "In vivo wide-field voltage imaging in zebrafish with voltage-sensitive dye and genetically encoded voltage indicator", "authors": "Hiyoshi K, et al.", "journal": "Development, Growth & Differentiation", "year": 2021, "sample": "Zebrafish larvae, cerebellum; spinal cord, in vivo", "url": "https://doi.org/10.1111/dgd.12744" }, { "title": "Enabling comprehensive optogenetic studies of mouse hearts by simultaneous opto-electrical panoramic mapping and stimulation", "authors": "Rieger M, et al.", "journal": "Nature Communications", "year": 2021, "sample": "Mouse heart, ex vivo", "url": "https://doi.org/10.1038/s41467-021-26039-8" }, { "title": "In Vitro Matured Human Pluripotent Stem Cell-Derived Cardiomyocytes Form Grafts With Enhanced Structure and Function in Injured Hearts", "authors": "Dhahri W, et al.", "journal": "Circulation", "year": 2022, "sample": "Human PSC-derived cardiomyocytes; rat heart, ex vivo", "url": "https://doi.org/10.1161/CIRCULATIONAHA.121.053563" }, { "title": "Zebrafish Embryos Display Characteristic Bioelectric Signals during Early Development", "authors": "Silic MR, et al.", "journal": "Cells", "year": 2022, "sample": "Zebrafish embryos, in vivo", "url": "https://doi.org/10.3390/cells11223586" }, { "title": "Optogenetic Reporters Delivered as mRNA Facilitate Repeatable Action Potential and Calcium Handling Assessment in Human iPSC-Derived Cardiomyocytes", "authors": "Yiangou L, et al.", "journal": "Stem Cells", "year": 2022, "sample": "Human iPSC-derived cardiomyocytes, in vitro", "url": "https://doi.org/10.1093/stmcls/sxac029" }, { "title": "A fully-automated low-cost cardiac monolayer optical mapping robot", "authors": "Lee P, et al.", "journal": "Frontiers in Cardiovascular Medicine", "year": 2023, "sample": "Human PSC-derived cardiomyocyte monolayers, in vitro", "url": "https://doi.org/10.3389/fcvm.2023.1096884" }, { "title": "Protocol for dual-optical mapping of voltage and calcium sensors in human pluripotent stem cell-derived cardiomyocytes", "authors": "Alibhai FJ, et al.", "journal": "STAR Protocols", "year": 2025, "sample": "Human PSC-derived cardiomyocytes, in vitro", "url": "https://doi.org/10.1016/j.xpro.2025.104044" }, { "title": "Absolute Membrane Potential Recording with ASAP-Type Genetically Encoded Voltage Indicators Using Fluorescence Lifetime Imaging", "authors": "Nair AG, et al.", "journal": "ACS Chemical Neuroscience", "year": 2025, "sample": "HEK293T cells, in vitro", "url": "https://doi.org/10.1021/acschemneuro.5c00670" }, { "title": "Bright and fast multicoloured voltage reporters via electrochromic FRET", "authors": "Zou P, Zhao Y, Douglass AD, Hochbaum DR, Brinks D, Werley CA, Harrison DJ, Campbell RE, Cohen AE", "journal": "Nature Communications", "year": 2014, "sample": "HEK293 cells; cultured rat hippocampal neurons, in vitro", "url": "https://doi.org/10.1038/ncomms5625" }, { "title": "High-speed recording of neural spikes in awake mice and flies with a fluorescent voltage sensor", "authors": "Gong Y, Huang C, Li JZ, Grewe BF, Zhang Y, Eismann S, Schnitzer MJ", "journal": "Science", "year": 2015, "sample": "HEK293 cells; mouse cortex, in vivo; Drosophila, in vivo", "url": "https://doi.org/10.1126/science.aab0810" }, { "title": "Ultrafast Two-Photon Imaging of a High-Gain Voltage Indicator in Awake Behaving Mice", "authors": "Villette V, Chavarha M, Dimov IK, Bradley J, Pradhan L, Mathieu B, Evans SW, Chamberland S, Shi D, Yang R, Kim BB, Ayon A, Jalil A, St-Pierre F, Schnitzer MJ, Bi G, Toth K, Ding J, Dieudonné S, Lin MZ", "journal": "Cell", "year": 2019, "sample": "HEK293A cells; mouse cortex and hippocampus, in vivo", "url": "https://doi.org/10.1016/j.cell.2019.11.004" }, { "title": "Improving a genetically encoded voltage indicator by modifying the cytoplasmic charge composition", "authors": "Lee S, Geiller T, Jung A, Nakajima R, Song YK, Baker BJ", "journal": "Scientific Reports", "year": 2017, "sample": "HEK293 cells, in vitro", "url": "https://doi.org/10.1038/s41598-017-08731-2" } ] }, { "id": "asap2s", "name": "ASAP2s", "url": "https://gevibench.org/gevi/asap2s", "description": "Higher-sensitivity ASAP1 variant that trades speed for larger fluorescence responses.", "category": "VSD-cpGFP GEVI", "tags": [ "cpGFP", "Two-photon" ], "year": 2017, "date": "2017-07-27", "paper": "eLife 2017", "paperUrl": "https://doi.org/10.7554/eLife.25690", "parent": { "id": "asap1", "name": "ASAP1" }, "crossBranchParent": null, "lineagePath": null, "polarity": "negative", "spectrum": { "peakExcitationNm": 490, "peakEmissionNm": 511, "fluorophore": "ASAP2s" }, "twoPhotonCompatible": true, "bioluminescent": false, "addgene": { "id": "101274", "url": "https://www.addgene.org/101274/", "note": null }, "lastUpdated": "2026-09-19", "display": { "brightness": "0.43×", "kinetics": "7.8 / 14", "dynamicRange": "-38.7%", "subthreshold": "0.15%", "sensitivity": "12.2%", "apWidth": "15 ms", "photostability": "5.5 s", "papers": "13" }, "values": { "brightnessRelEGFP": 0.4345, "tauOnMs": 7.785, "tauOffMs": 14.03, "dffPer100mV": -38.7, "dffPer100mVMedianAbs": 36, "dffPerMV": 0.15, "dffPerMVEstimatedFromFV": false, "dffPerAP": 12.2, "dffPerAPMedianAbs": 9.6, "apWidthMs": 14.8, "t75At100mWmm2Sec": 5.5, "papersUsingIt": 13 }, "measurements": { "kinetics": [ { "tauOnMs": 30.6, "tauOffMs": 65.8, "temperature": "22°C", "dye": null, "source": { "label": "Chamberland 2017, Table 1", "url": "https://doi.org/10.7554/eLife.25690" } }, { "tauOnMs": 3.58, "tauOffMs": 6.99, "temperature": "32-35°C", "dye": null, "source": { "label": "Villette 2019, Table 1", "url": "https://doi.org/10.1016/j.cell.2019.11.004" } }, { "tauOnMs": 11.99, "tauOffMs": 21.07, "temperature": "32-35°C", "dye": null, "source": { "label": "Liu 2022, Table S2", "url": "https://doi.org/10.1016/j.cell.2022.07.013" } }, { "tauOnMs": 16.71, "tauOffMs": 29.51, "temperature": "21-23°C", "dye": null, "source": { "label": "Liu 2022, Table S2", "url": "https://doi.org/10.1016/j.cell.2022.07.013" } } ], "dffPer100mV": [ { "dff": -38.7, "responseType": "steady-state", "modality": "1P", "dye": null, "source": { "label": "Chamberland 2017, Fig. 1C", "url": "https://doi.org/10.7554/eLife.25690" } }, { "dff": -36, "responseType": "steady-state", "modality": "1P", "dye": null, "source": { "label": "Villette 2019, Table 1", "url": "https://doi.org/10.1016/j.cell.2019.11.004" } }, { "dff": -28.5, "responseType": "steady-state", "modality": "1P", "dye": null, "source": { "label": "Lu 2023, Fig. 2c", "url": "https://doi.org/10.1038/s41467-023-41975-3" } }, { "dff": -33.3, "responseType": "steady-state", "modality": "2P", "dye": null, "source": { "label": "Liu 2022, Fig. 2C", "url": "https://doi.org/10.1016/j.cell.2022.07.013" } } ], "dffPerMV": [ { "slopePercentPerMV": 0.15, "modality": "2P", "note": "Subthreshold voltage sensitivity ≈ 0.15 %/mV, from the Fig. 5G peak-response-vs-depolarization plot in organotypic hippocampal slices under 2P (462 Hz). ASAP2s ΔF/F peaks were ~0.9%/1.35%/2.2%/3.15% for 5/10/15/20 mV subthreshold depolarization waveforms (least-squares slope 0.154 %/mV, read from the bar graph). This is a 2P slice measurement of responses to subthreshold-waveform commands (not a 1P near-rest voltage-clamp ramp), so it may not be directly comparable to 1P subthreshold slopes.", "source": { "label": "Chamberland 2017, Fig. 5G", "url": "https://doi.org/10.7554/eLife.25690" } } ], "dffPerAP": [ { "dff": 12.2, "modality": "1P", "dye": null, "source": { "label": "Chamberland 2017, Fig. 2I", "url": "https://doi.org/10.7554/eLife.25690" } }, { "dff": 15, "modality": "2P", "dye": null, "source": { "label": "Chamberland 2017, Fig. 4E", "url": "https://doi.org/10.7554/eLife.25690" } }, { "dff": 7, "modality": "1P", "dye": null, "source": { "label": "Villette 2019, Table 1", "url": "https://doi.org/10.1016/j.cell.2019.11.004" } } ], "apWidth": [ { "fwhmMs": 21.1, "samplingRateHz": 3700, "sample": "Organotypic hippocampal slice cultures, current-triggered single AP, two-photon RAMP (FEVIR)", "modality": "2P", "temperature": "22°C", "source": { "label": "Chamberland 2017, Fig. 6E", "url": "https://doi.org/10.7554/eLife.25690" } }, { "fwhmMs": 17.4, "samplingRateHz": 1000, "sample": "Rat, cultured hippocampal neurons, in vitro", "modality": "1P", "temperature": null, "source": { "label": "Chamberland 2017, Fig. 2J", "url": "https://doi.org/10.7554/eLife.25690" } }, { "fwhmMs": 17.3, "sample": "Rat, cultured hippocampal neurons, current-evoked single AP, room temperature, in vitro", "modality": "1P", "temperature": "22°C", "source": { "label": "Villette 2019, Fig. 2F", "url": "https://doi.org/10.1016/j.cell.2019.11.004" } }, { "fwhmMs": 12.3, "samplingRateHz": 500, "sample": "Mouse, cultured hippocampal neurons, current-triggered single AP, in vitro", "modality": "1P", "temperature": null, "source": { "label": "Milosevic 2020, Fig. 1B", "url": "https://doi.org/10.1523/ENEURO.0060-20.2020" } }, { "fwhmMs": 6.7, "samplingRateHz": 987, "sample": "HEK293A cells, optical response to a single AP-like command waveform (2-ms FWHM, +30 mV peak, −70 mV baseline), whole-cell voltage clamp, in vitro", "modality": "1P", "temperature": "25°C", "source": { "label": "Lu 2023, Fig. 2f", "url": "https://doi.org/10.1038/s41467-023-41975-3" } }, { "fwhmMs": 7.6, "samplingRateHz": 440, "sample": "HEK293A cells, optical response to a single AP-like command waveform (2-ms FWHM), whole-cell voltage clamp, in vitro", "modality": "2P", "temperature": "22°C", "source": { "label": "Liu 2022, Fig. 2F", "url": "https://doi.org/10.1016/j.cell.2022.07.013" } } ], "brightness": [ { "ratio": 1.11, "reference": "ASAP3", "source": { "label": "Liu 2022, Fig. 2I", "url": "https://doi.org/10.1016/j.cell.2022.07.013" } }, { "ratio": 1, "reference": "ASAP1", "source": { "label": "Chamberland 2017, Fig. 1—figure supplement 2A", "url": "https://doi.org/10.7554/eLife.25690" } }, { "ratio": 0.45454545454545453, "reference": "JEDI-1P", "source": { "label": "Lu 2023, Fig. 2h", "url": "https://doi.org/10.1038/s41467-023-41975-3" } }, { "ratio": 0.6666666666666666, "reference": "JEDI-2P", "source": { "label": "Liu 2022, Fig. 2I", "url": "https://doi.org/10.1016/j.cell.2022.07.013" } } ], "photobleaching": [ { "modality": "1P", "illumination": "46 mW/mm²", "intensityMWmm2": 46, "t75Sec": null, "t50Sec": null, "extrapolated": false, "source": { "label": "Lu 2023, Fig. S4", "url": "https://doi.org/10.1038/s41467-023-41975-3" } }, { "modality": "1P", "illumination": "15 mW/mm²", "intensityMWmm2": 15, "t75Sec": null, "t50Sec": 3, "extrapolated": false, "source": { "label": "Liu 2022, Fig. S4G", "url": "https://doi.org/10.1016/j.cell.2022.07.013" } }, { "modality": "1P", "illumination": "11 mW/mm²", "intensityMWmm2": 11, "t75Sec": 50, "t50Sec": null, "extrapolated": false, "source": { "label": "Chamberland 2017, Fig. 1—figure supplement 3A", "url": "https://doi.org/10.7554/eLife.25690" } }, { "modality": "1P", "illumination": "24 mW/mm²", "intensityMWmm2": 24, "t75Sec": 47.7, "t50Sec": null, "extrapolated": false, "source": { "label": "Chamberland 2017, Fig. 2—figure supplement 3B", "url": "https://doi.org/10.7554/eLife.25690" } }, { "modality": "2P", "illumination": "6.4 mW at objective back aperture (2P, 920 nm)", "intensityMWmm2": null, "t75Sec": 20.4, "t50Sec": null, "extrapolated": false, "source": { "label": "Chamberland 2017, Fig. 1—figure supplement 4A", "url": "https://doi.org/10.7554/eLife.25690" } } ] }, "t75ScaledEntriesSec": [ 0.45, 5.5, 11.45 ], "fvCurve": { "voltageMV": [ -100, -90, -80, -70, -60, -50, -40, -30, -20, -10, 0, 10, 20, 30, 40, 50 ], "dffPercent": [ 18, 13, 6, 0, -6, -11, -17, -21, -25, -28, -32, -34, -37, -39, -40, -42 ], "source": { "label": "Chamberland 2017, Fig. 1C", "url": "https://doi.org/10.7554/eLife.25690" } }, "researchPapers": [ { "title": "Fast two-photon imaging of subcellular voltage dynamics in neuronal tissue with genetically encoded indicators", "authors": "Chamberland S, Yang HH, et al.", "journal": "eLife", "year": 2017, "sample": "HEK293A cells; rat hippocampal neurons; Drosophila L2 neurons; mouse hippocampal slice cultures, in vitro/in vivo", "url": "https://doi.org/10.7554/eLife.25690" }, { "title": "In Vitro Testing of Voltage Indicators: Archon1, ArcLightD, ASAP1, ASAP2s, ASAP3b, Bongwoori-Pos6, BeRST1, FlicR1, and Chi-VSFP-Butterfly", "authors": "Milosevic MM, et al.", "journal": "eNeuro", "year": 2020, "sample": "Mouse, cortex; HEK293 cells, in vitro", "url": "https://doi.org/10.1523/ENEURO.0060-20.2020" }, { "title": "Ultrafast two-photon imaging of a high-gain voltage indicator in awake behaving mice", "authors": "Villette V, et al.", "journal": "Cell", "year": 2019, "sample": "Rat, hippocampal neurons; HEK293A cells; mouse, cerebellar slices, in vitro", "url": "https://doi.org/10.1016/j.cell.2019.11.004" }, { "title": "Imaging of Evoked Cortical Depolarizations Using Either ASAP2s, or chi-VSFP, or Di-4-Anepps, or Autofluorescence Optical Signals", "authors": "Milicevic KD, et al.", "journal": "Journal of Integrative Neuroscience", "year": 2023, "sample": "Mouse, cortex; ASAP2s-transgenic brain slices, ex vivo", "url": "https://doi.org/10.31083/j.jin2206160" }, { "title": "CaMKII Measures the Passage of Time to Coordinate Behavior and Motivational State", "authors": "Thornquist SC, et al.", "journal": "Neuron", "year": 2019, "sample": "Drosophila, abdominal ganglion neurons, ex vivo", "url": "https://doi.org/10.1016/j.neuron.2019.10.018" }, { "title": "Biochemical evidence accumulates across neurons to drive a network-level eruption", "authors": "Thornquist SC, et al.", "journal": "Molecular Cell", "year": 2021, "sample": "Drosophila, Corazonin neurons, ex vivo", "url": "https://doi.org/10.1016/j.molcel.2020.12.029" }, { "title": "Drosophila TRPγ is required in neuroendocrine cells for post-ingestive food selection", "authors": "Dhakal S, et al.", "journal": "eLife", "year": 2022, "sample": "Drosophila, DH44 neuroendocrine cells, in vivo", "url": "https://doi.org/10.7554/eLife.56726" }, { "title": "Triclosan disrupts immune cell function by depressing Ca2+ influx following acidification of the cytoplasm", "authors": "Sangroula S, et al.", "journal": "Toxicology and Applied Pharmacology", "year": 2020, "sample": "RBL mast cells, in vitro", "url": "https://doi.org/10.1016/j.taap.2020.115205" }, { "title": "Real-time detection of changes in yeast plasma membrane potential using genetically encoded voltage indicator proteins", "authors": "Limapichat W, et al.", "journal": "FEMS Yeast Research", "year": 2020, "sample": "Saccharomyces cerevisiae, in vitro", "url": "https://doi.org/10.1093/femsyr/foaa041" }, { "title": "A dark quencher genetically encodable voltage indicator (dqGEVI) exhibits high fidelity and speed", "authors": "Alich TC, et al.", "journal": "Proceedings of the National Academy of Sciences", "year": 2021, "sample": "HEK293T cells; rat hippocampal neurons, in vitro", "url": "https://doi.org/10.1073/pnas.2020235118" }, { "title": "Sustained deep-tissue voltage recording using a fast indicator evolved for two-photon microscopy", "authors": "Liu Z, Lu X, et al.", "journal": "Cell", "year": 2022, "sample": "HEK293A cells; mouse hippocampal neurons, in vitro", "url": "https://doi.org/10.1016/j.cell.2022.07.013" }, { "title": "Widefield imaging of rapid pan-cortical voltage dynamics with an indicator evolved for one-photon microscopy", "authors": "Lu X, Wang Y, Liu Z, et al.", "journal": "Nature Communications", "year": 2023, "sample": "HEK293A cells; HEK293-Kir2.1 cells; rat cortical and hippocampal neurons, in vitro", "url": "https://doi.org/10.1038/s41467-023-41975-3" }, { "title": "Physiological features of parvalbumin-expressing GABAergic interneurons contributing to high-frequency oscillations in the cerebral cortex", "authors": "Milicevic KD, et al.", "journal": "Current Research in Neurobiology", "year": 2023, "sample": "Mouse, cortex; 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acute slices, in vitro", "url": "https://doi.org/10.1016/j.isci.2020.101363" }, { "title": "In Vitro Testing of Voltage Indicators: Archon1, ArcLightD, ASAP1, ASAP2s, ASAP3b, Bongwoori-Pos6, BeRST1, FlicR1, and Chi-VSFP-Butterfly", "authors": "Milosevic MM, et al.", "journal": "eNeuro", "year": 2020, "sample": "Mouse, cortex; HEK293 cells, in vitro", "url": "https://doi.org/10.1523/ENEURO.0060-20.2020" }, { "title": "A red fluorescent protein with improved monomericity enables ratiometric voltage imaging with ASAP3", "authors": "Kim BB, et al.", "journal": "Scientific Reports", "year": 2022, "sample": "Rat, hippocampal neurons; human iPSC-derived cardiomyocytes; HEK293 cells, in vitro", "url": "https://doi.org/10.1038/s41598-022-07313-1" }, { "title": "Sustained deep-tissue voltage recording using a fast indicator evolved for two-photon microscopy", "authors": "Liu Z, et al.", "journal": "Cell", "year": 2022, "sample": "Drosophila, visual interneurons, in vivo; mouse, retina, ex vivo; mouse, visual cortex L2/3 and L5, in vivo", "url": "https://doi.org/10.1016/j.cell.2022.07.013" }, { "title": "Electrical signals in the ER are cell type and stimulus specific with extreme spatial compartmentalization in neurons", "authors": "Campbell EP, et al.", "journal": "Cell Reports", "year": 2023, "sample": "Rat, hippocampal neurons; HEK293T cells, in vitro", "url": "https://doi.org/10.1016/j.celrep.2022.111943" }, { "title": "High-speed low-light in vivo two-photon voltage imaging of large neuronal populations", "authors": "Platisa J, et al.", "journal": "Nature Methods", "year": 2023, "sample": "Mouse, somatosensory cortex L2/3, in vivo; acute slices, in vitro", "url": "https://doi.org/10.1038/s41592-023-01820-3" }, { "title": "A positively tuned voltage indicator for extended electrical recordings in the brain", "authors": "Evans SW, et al.", "journal": "Nature Methods", "year": 2023, "sample": "Mouse, hippocampus, cortex, in vivo; Drosophila, visual interneurons, in vivo; rat, hippocampal neurons, in vitro", "url": "https://doi.org/10.1038/s41592-023-01913-z" }, { "title": "Widefield imaging of rapid pan-cortical voltage dynamics with an indicator evolved for one-photon microscopy", "authors": "Lu X, Wang Y, Liu Z, Gou Y, Jaeger D, St-Pierre F", "journal": "Nature Communications", "year": 2023, "sample": "HEK293A cells, in vitro (ASAP3 benchmarked against JEDI-1P for response amplitude, spike-waveform width and photostability); rat hippocampal neurons, in vitro (ASAP3-Kv single-AP responses)", "url": "https://doi.org/10.1038/s41467-023-41975-3" }, { "title": "Dendritic mGluR2 and perisomatic Kv3 signaling regulate dendritic computation of mouse starburst amacrine cells", "authors": "Ledesma HA, et al.", "journal": "Nature Communications", "year": 2024, "sample": "Mouse, retina, starburst amacrine cells, ex vivo", "url": "https://doi.org/10.1038/s41467-024-46234-7" }, { "title": "Two-dimensional electro-optical multiphoton microscopy", "authors": "Farinella DM, et al.", "journal": "Neurophotonics", "year": 2024, "sample": "Mouse, visual cortex L2/3, in vivo", "url": "https://doi.org/10.1117/1.NPh.11.2.025005" }, { "title": "A novel rhodopsin-based voltage indicator for simultaneous two-photon optical recording with GCaMP in vivo", "authors": "Villette V, Yang S, Valenti R, Macklin JJ, Bradley J, Mathieu B, Lombardini A, Podgorski K, Dieudonné S, Schreiter ER, Abdelfattah AS", "journal": "bioRxiv", "year": 2024, "sample": "Mouse, cortex and cerebellum, in vivo (ASAP3 spike amplitude / spike FWHM / photon flux cross-plotted against 2Photron-ST552 and JEDI-2P-Kv)", "url": "https://doi.org/10.1101/2024.11.15.623698" }, { "title": "Cl⁻-dependent amplification of excitatory synaptic potentials at distal dendrites revealed by voltage imaging", "authors": "Higashi R, et al.", "journal": "Science Advances", "year": 2024, "sample": "Mouse, hippocampal neurons; 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melanoma cells; breast cancer cells, in vitro", "url": "https://doi.org/10.1021/acschemneuro.5c00670" }, { "title": "Voltage imaging reveals hippocampal inhibitory dynamics shaping pyramidal memory-encoding sequences", "authors": "Taxidis J, et al.", "journal": "Nature Neuroscience", "year": 2025, "sample": "Mouse, hippocampus CA1, PV and SST interneurons, in vivo", "url": "https://doi.org/10.1038/s41593-025-02016-y" }, { "title": "An axonal brake on striatal dopamine output by cholinergic interneurons", "authors": "Zhang YF, et al.", "journal": "Nature Neuroscience", "year": 2025, "sample": "Mouse, striatum, dopamine axons, in vivo; acute slices, ex vivo", "url": "https://doi.org/10.1038/s41593-025-01906-5" }, { "title": "Imaging high-frequency voltage dynamics in multiple neuron classes of behaving mammals", "authors": "Haziza S, et al.", "journal": "Cell", "year": 2025, "sample": "Mouse, hippocampus CA1/CA3, visual cortex, motor cortex, in vivo; freely moving and head-fixed", "url": "https://doi.org/10.1016/j.cell.2025.06.028" }, { "title": "Neuronal activity induces myelin voltage changes that reflect action potential dependent myelin potassium buffering", "authors": "Labarchède M, et al.", "journal": "bioRxiv", "year": 2025, "sample": "Mouse, oligodendrocytes, myelin, in vivo", "url": "https://doi.org/10.64898/2025.12.16.694668" }, { "title": "Sub-Millivolt Voltage Imaging Reveals Gap Junction-Mediated Bioelectric Contact Inhibition", "authors": "Rühl P, Hussein R, Reuter S, Frahnert K, Nair AG, Mrowka R, Schönherr R, Heinemann SH", "journal": "bioRxiv", "year": 2026, "sample": "HEK293T cells, in vitro (ASAP3 benchmarked alongside rEstus2s)", "url": "https://doi.org/10.64898/2026.02.10.701308" }, { "title": "Ultrasensitive voltage imaging reveals distinct electrical microdomains in neurons", "authors": "Hao YA, Jayne LL, Lee S, Dittrich MN, Zhang M, et al.", "journal": "bioRxiv", "year": 2026, "sample": "Mouse, cortex PV interneurons, in vivo (ASAP3 photostability vs ASAP7y); 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Fig. 5c", "url": "https://doi.org/10.1038/s41586-019-1641-1" } }, { "dff": 6.5, "modality": "1P", "dye": "JF525", "source": { "label": "Hasseman 2022, Fig. 2", "url": "https://doi.org/10.25378/janelia.21534411" } } ], "apWidth": [ { "fwhmMs": 3, "samplingRateHz": 985, "sample": "Cultured rat hippocampal neurons, uM Workstation whole-cell current clamp, single evoked AP (2 nA, 2 ms pulse)", "modality": "1P", "temperature": "23°C", "source": { "label": "Abdelfattah 2023, Fig. 2F", "url": "https://doi.org/10.1016/j.neuron.2023.03.009" } } ], "brightness": [ { "ratio": 4, "reference": "Ace2N-mNeon", "source": { "label": "Abdelfattah 2019, Fig. 1F", "url": "https://doi.org/10.1126/science.aav6416" } }, { "ratio": 5.2631578947368425, "reference": "HVI", "source": { "label": "Liu 2021, Table S4", "url": "https://doi.org/10.1038/s41557-021-00641-1" } }, { "ratio": 1.6949152542372883, "reference": "Positron", "source": { "label": "Abdelfattah 2020, Fig. 2b", "url": "https://doi.org/10.1038/s41467-020-17322-1" } }, { "ratio": 1.5384615384615383, "reference": "Voltron2", "source": { "label": "Abdelfattah 2023, Fig. 2H", "url": "https://doi.org/10.1016/j.neuron.2023.03.009" } } ], "photobleaching": [ { "modality": "1P", "illumination": "25 mW/mm² (510/25 nm)", "intensityMWmm2": null, "t75Sec": 30.5, "t50Sec": null, "extrapolated": false, "source": { "label": "Piatkevich 2019, Ext. Fig. 5e", "url": "https://doi.org/10.1038/s41586-019-1641-1" } }, { "modality": "1P", "illumination": "~23 mW/mm² (510/25 nm)", "intensityMWmm2": 23, "t75Sec": 61, "t50Sec": null, "extrapolated": false, "source": { "label": "Abdelfattah 2019, Fig. 1H", "url": "https://doi.org/10.1126/science.aav6416" } }, { "modality": "1P", "illumination": "14.7 mW/mm² (510/25 nm)", "intensityMWmm2": 14.7, "t75Sec": 245, "t50Sec": null, "extrapolated": false, "source": { "label": "Abdelfattah 2023, Fig. 2I", "url": "https://doi.org/10.1016/j.neuron.2023.03.009" } }, { "modality": "1P", "illumination": "70–140 mW/mm² (DMD patterned, per-cell)", "intensityMWmm2": null, "t75Sec": null, "t50Sec": null, "extrapolated": false, "source": { "label": "Abdelfattah 2023, Fig. S8d", "url": "https://doi.org/10.1016/j.neuron.2023.03.009" } } ] }, "t75ScaledEntriesSec": [ 14.03, 36.02 ], "fvCurve": { "voltageMV": [ -110, -90, -70, -50, -30, -10, 10, 30, 50 ], "dffPercent": [ 8, 4.5, 0, -4.5, -9, -13.5, -18, -22.5, -27 ], "source": { "label": "Abdelfattah 2019, Fig. 1E", "url": "https://doi.org/10.1126/science.aav6416" } }, "researchPapers": [ { "title": "Bright and photostable chemigenetic indicators for extended in vivo voltage imaging", "authors": "Abdelfattah AS, Kawashima T, Singh A, et al.", "journal": "Science", "year": 2019, "sample": "Mouse cortex L1, larval zebrafish, Drosophila, in vivo; HEK cells", "url": "https://doi.org/10.1126/science.aav6416" }, { "title": "High-fidelity estimates of spikes and subthreshold waveforms from 1-photon voltage imaging in vivo", "authors": "Xie ME, et al.", "journal": "Cell Reports", "year": 2021, "sample": "Mouse, somatosensory cortex layer 1, in vivo", "url": "https://doi.org/10.1016/j.celrep.2021.108954" }, { "title": "VolPy: Automated and scalable analysis pipelines for voltage imaging datasets", "authors": "Cai C, et al.", "journal": "PLOS Computational Biology", "year": 2021, "sample": "Mouse cortex L1 and larval zebrafish tegmentum, in vivo", "url": "https://doi.org/10.1371/journal.pcbi.1008806" }, { "title": "Single Photon Kilohertz Frame Rate Imaging of Neural Activity", "authors": "Tian T, Yuan Y, Mitra S, et al.", "journal": "Advanced Science", "year": 2022, "sample": "Rat, hippocampal CA1, acute brain slices, Voltron-JF525", "url": "https://doi.org/10.1002/advs.202203018" }, { "title": "Voltage-Seq: all-optical postsynaptic connectome-guided single-cell transcriptomics", "authors": "Csillag V, et al.", "journal": "Nature Methods", "year": 2023, "sample": "Mouse, periaqueductal gray (PAG), acute brain slices, Voltron-ST + JF585", "url": "https://doi.org/10.1038/s41592-023-01965-1" }, { "title": "All-optical voltage imaging-guided postsynaptic single-cell transcriptome profiling with Voltage-Seq", "authors": "Csillag V, Noble JC, Calvigioni D, et al.", "journal": "Nature Protocols", "year": 2024, "sample": "Mouse, acute brain slices, Voltron-ST", "url": "https://doi.org/10.1038/s41596-024-01005-y" }, { "title": "Dynamic assemblies of parvalbumin interneurons in brain oscillations", "authors": "Huang YC, et al.", "journal": "Neuron", "year": 2024, "sample": "Mouse, hippocampal CA1, PV interneurons, in vivo", "url": "https://doi.org/10.1016/j.neuron.2024.05.015" }, { "title": "Optical Recordings of Unitary Synaptic Connections Reveal High and Random Local Connectivity between CA3 Pyramidal Cells", "authors": "Layous R, et al.", "journal": "Journal of Neuroscience", "year": 2025, "sample": "Rat, hippocampal CA3 pyramidal cells, acute brain slices, Voltron-ST + JF549", "url": "https://doi.org/10.1523/JNEUROSCI.0102-25.2025" }, { "title": "Voltage imaging reveals circuit computations in the raphe underlying serotonin-mediated motor vigor learning", "authors": "Kawashima T, et al.", "journal": "Neuron", "year": 2025, "sample": "Larval zebrafish, serotonergic raphe (DRN), in vivo", "url": "https://doi.org/10.1016/j.neuron.2025.05.017" } ] }, { "id": "voltron2", "name": "Voltron2", "url": "https://gevibench.org/gevi/voltron2", "description": "Improved Voltron with enhanced sensitivity, retaining fast kinetics and photostability.", "category": "Chemigenetic GEVI", "tags": [ "Ace opsin", "HaloTag" ], "year": 2023, "date": "2023-04-03", "paper": "Neuron 2023", "paperUrl": "https://doi.org/10.1016/j.neuron.2023.03.009", "parent": { "id": "voltron", "name": "Voltron" }, "crossBranchParent": null, "lineagePath": null, "polarity": "negative", "spectrum": { "peakExcitationNm": 534, "peakEmissionNm": 554, "fluorophore": "Voltron2 (JF525)" }, "twoPhotonCompatible": true, "bioluminescent": false, "addgene": { "id": "172909", "url": "https://www.addgene.org/172909/", "note": null }, "lastUpdated": "2026-10-12", "display": { "brightness": "7.18×", "kinetics": "4.0 / 5.5", "dynamicRange": "-34%", "subthreshold": "0.51%", "sensitivity": "10.09%", "apWidth": "3.2 ms", "photostability": "42 s", "papers": "23" }, "values": { "brightnessRelEGFP": 7.176, "tauOnMs": 3.96, "tauOffMs": 5.53, "dffPer100mV": -34, "dffPer100mVMedianAbs": 27, "dffPerMV": 0.51, "dffPerMVEstimatedFromFV": false, "dffPerAP": 10.09, "dffPerAPMedianAbs": 12, "apWidthMs": 3.2, "t75At100mWmm2Sec": 41.5, "papersUsingIt": 23 }, "measurements": { "kinetics": [ { "tauOnMs": 1.5, "tauOffMs": 3.05, "temperature": "23°C", "dye": null, "source": { "label": "Abdelfattah 2023, Fig. 2E", "url": "https://doi.org/10.1016/j.neuron.2023.03.009" } }, { "tauOnMs": 3.96, "tauOffMs": 5.53, "temperature": "25°C", "dye": null, "source": { "label": "Liu 2025, Table S2", "url": "https://doi.org/10.1126/sciadv.ads1807" } } ], "dffPer100mV": [ { "dff": -34, "responseType": "steady-state", "modality": "1P", "dye": "JF525", "source": { "label": "Abdelfattah 2023, Fig. 2B", "url": "https://doi.org/10.1016/j.neuron.2023.03.009" } }, { "dff": -23, "responseType": "steady-state", "modality": "1P", "dye": "JF525", "source": { "label": "Cao 2025, Fig. 1C", "url": "https://doi.org/10.1101/2025.05.29.656768" } }, { "dff": -21.2, "responseType": "steady-state", "modality": "1P", "dye": "JF525", "source": { "label": "Liu 2025, Table S2", "url": "https://doi.org/10.1126/sciadv.ads1807" } }, { "dff": -22, "responseType": "steady-state", "modality": "1P", "dye": "JF585", "source": { "label": "Yang 2024, Fig. 1C", "url": "https://doi.org/10.1101/2024.02.02.578569" } }, { "dff": -31, "responseType": "steady-state", "modality": "1P", "dye": "JF525", "source": { "label": "Hasseman 2022, Fig. 1", "url": "https://doi.org/10.25378/janelia.21534411" } } ], "dffPerMV": [ { "slopePercentPerMV": 0.54, "modality": null, "note": "Voltron2525 subthreshold slope 0.54%/mV ± 0.01%/mV in the −90 to −50 mV range (cultured neurons, n = 8 cells), an almost 3-fold higher slope than Voltron525 (0.21%/mV). Reported in the running main text; not a labeled figure panel (Fig. 3D is the synPSP sensitivity index d'/mV of Voltron2585, a different dye/metric).", "source": { "label": "Abdelfattah 2023, Main text", "url": "https://doi.org/10.1016/j.neuron.2023.03.009" } }, { "slopePercentPerMV": 0.48, "modality": null, "note": "Voltron2525-Kv subthreshold voltage-sensitivity slope ≈ 0.48 %/mV, read from the Fig. 5C comparison in the Hao et al. 2024 ASAP5 paper (soma-targeted Voltron2525-Kv, cultured rat hippocampal neurons), where the paper's primary sensor ASAP5-Kv is ≈ 0.83 %/mV. A different prep/metric from the origin-paper 0.54 %/mV entry above (that one is the −90→−50 mV subthreshold slope of un-targeted Voltron2525).", "source": { "label": "Hao 2024, Fig. 5C", "url": "https://doi.org/10.1016/j.neuron.2024.08.019" } } ], "dffPerAP": [ { "dff": 10.09, "modality": "1P", "dye": "JF525", "source": { "label": "Abdelfattah 2023, Fig. 2G", "url": "https://doi.org/10.1016/j.neuron.2023.03.009" } }, { "dff": 12, "modality": "1P", "dye": "JF525", "source": { "label": "Liu 2025, Table S2", "url": "https://doi.org/10.1126/sciadv.ads1807" } }, { "dff": 15, "modality": "1P", "dye": "JF525", "source": { "label": "Hasseman 2022, Fig. 2", "url": "https://doi.org/10.25378/janelia.21534411" } }, { "dff": 12.3, "modality": "1P", "dye": "JF585", "source": { "label": "Yang 2024, Fig. 2B", "url": "https://doi.org/10.1101/2024.02.02.578569" } } ], "apWidth": [ { "fwhmMs": 3.2, "samplingRateHz": 985, "sample": "Cultured rat hippocampal neurons, uM Workstation whole-cell current clamp, single evoked AP (2 nA, 2 ms pulse)", "modality": "1P", "temperature": "23°C", "source": { "label": "Abdelfattah 2023, Fig. 2F", "url": "https://doi.org/10.1016/j.neuron.2023.03.009" } }, { "fwhmMs": 7, "samplingRateHz": 484, "sample": "Rat, hippocampal neurons, in vitro", "modality": "1P", "temperature": "25°C", "source": { "label": "Yang 2024, Fig. 2B", "url": "https://doi.org/10.1101/2024.02.02.578569" } }, { "fwhmMs": 0.85, "samplingRateHz": 3500, "sample": "Mouse, cerebellar granular-layer Golgi/glycinergic interneurons, in vivo, awake, Voltron2-ST552 (JF552)", "modality": "2P", "temperature": null, "source": { "label": "Villette 2024, Fig. S9d", "url": "https://doi.org/10.1101/2024.11.15.623698" } } ], "brightness": [ { "ratio": 0.65, "reference": "Voltron", "source": { "label": "Abdelfattah 2023, Fig. 2H", "url": "https://doi.org/10.1016/j.neuron.2023.03.009" } } ], "photobleaching": [ { "modality": "1P", "illumination": "14.7 mW/mm²", "intensityMWmm2": 14.7, "t75Sec": 307, "t50Sec": null, "extrapolated": false, "source": { "label": "Abdelfattah 2023, Fig. 2I", "url": "https://doi.org/10.1016/j.neuron.2023.03.009" } }, { "modality": "1P", "illumination": "50 mW/mm²", "intensityMWmm2": 50, "t75Sec": 83, "t50Sec": null, "extrapolated": false, "source": { "label": "Liu 2025, Fig. S7", "url": "https://doi.org/10.1126/sciadv.ads1807" } }, { "modality": "1P", "illumination": "46 mW/mm²", "intensityMWmm2": 46, "t75Sec": 77, "t50Sec": null, "extrapolated": true, "source": { "label": "Hao 2024, Fig. S5E", "url": "https://doi.org/10.1016/j.neuron.2024.08.019" } }, { "modality": "1P", "illumination": "70–140 mW/mm² (DMD patterned, per-cell)", "intensityMWmm2": null, "t75Sec": null, "t50Sec": null, "extrapolated": false, "source": { "label": "Abdelfattah 2023, Fig. S8d", "url": "https://doi.org/10.1016/j.neuron.2023.03.009" } } ] }, "t75ScaledEntriesSec": [ 45.13, 41.5, 35.42 ], "fvCurve": { "voltageMV": [ -110, -90, -70, -50, -30, -10, 10, 30, 50 ], "dffPercent": [ 23, 11, 0, -10, -18, -25, -30, -34, -37 ], "source": { "label": "Abdelfattah 2023, Fig. 2B", "url": "https://doi.org/10.1016/j.neuron.2023.03.009" } }, "researchPapers": [ { "title": "Sensitivity optimization of a rhodopsin-based fluorescent voltage indicator", "authors": "Abdelfattah AS, et al.", "journal": "Neuron", "year": 2023, "sample": "Cultured neurons; flies, zebrafish, and mice, in vivo", "url": "https://doi.org/10.1016/j.neuron.2023.03.009" }, { "title": "Photophysics-informed two-photon voltage imaging using FRET-opsin voltage indicators", "authors": "Brooks FP, et al.", "journal": "Science Advances", "year": 2025, "sample": "Mouse, barrel cortex, in vivo, two-photon", "url": "https://doi.org/10.1126/sciadv.adp5763" }, { "title": "Imaging neuronal voltage beyond the scattering limit", "authors": "Chen TW, et al.", "journal": "Nature Methods", "year": 2025, "sample": "Mouse, hippocampus CA1, in vivo", "url": "https://doi.org/10.1038/s41592-025-02692-5" }, { "title": "Large-scale deep tissue voltage imaging with targeted-illumination confocal microscopy", "authors": "Xiao S, et al.", "journal": "Nature Methods", "year": 2024, "sample": "Mouse, cortex, in vivo", "url": "https://doi.org/10.1038/s41592-024-02275-w" }, { "title": "Two-photon voltage imaging with rhodopsin-based sensors", "authors": "Grimm C, et al.", "journal": "Neuron", "year": 2026, "sample": "Mouse, hippocampal slices; zebrafish larvae; mouse cortex, in vivo, two-photon", "url": "https://doi.org/10.1016/j.neuron.2025.12.014" }, { "title": "Orange/far-red hybrid voltage indicators with reduced phototoxicity enable reliable long-term imaging in neurons and cardiomyocytes", "authors": "Liu Y, et al.", "journal": "PNAS", "year": 2023, "sample": "Rat, hippocampal neurons, in vitro; cardiomyocytes, in vitro", "url": "https://doi.org/10.1073/pnas.2306950120" }, { "title": "Miniaturized widefield microscope for high speed in vivo voltage imaging", "authors": "Saladrigas CA, et al.", "journal": "Biomedical Optics Express", "year": 2025, "sample": "Mouse, visual cortex interneurons, in vivo", "url": "https://doi.org/10.1364/BOE.576516" }, { "title": "Voltage imaging with periodic structured illumination", "authors": "Speed F, et al.", "journal": "Biomedical Optics Express", "year": 2025, "sample": "Mouse, somatosensory, visual, and motor cortex, in vivo, Voltron2-JF552", "url": "https://doi.org/10.1364/BOE.573621" }, { "title": "Relative phase of membrane potential theta oscillations between individual hippocampal neurons code space", "authors": "Athif M, et al.", "journal": "bioRxiv", "year": 2025, "sample": "Mouse, hippocampus CA1, in vivo", "url": "https://doi.org/10.1101/2025.11.14.688496" }, { "title": "Dendritic excitations govern back-propagation via a spike-rate accelerometer", "authors": "Park P, et al.", "journal": "Nature Communications", "year": 2025, "sample": "Mouse, hippocampus CA1 pyramidal neurons, acute brain slices, Optopatch-V (Voltron2-based)", "url": "https://doi.org/10.1038/s41467-025-55819-9" }, { "title": "A fast and responsive voltage indicator with enhanced sensitivity for unitary synaptic events", "authors": "Hao YA, et al.", "journal": "Neuron", "year": 2024, "sample": "Rat, hippocampal neurons, in vitro", "url": "https://doi.org/10.1016/j.neuron.2024.08.019" }, { "title": "Fast dendritic excitations primarily mediate back-propagation in CA1 pyramidal neurons during behavior", "authors": "Lee BH, et al.", "journal": "bioRxiv", "year": 2026, "sample": "Mouse, hippocampus CA1 pyramidal neurons, in vivo, Voltron2-JF608", "url": "https://doi.org/10.64898/2026.01.03.696606" }, { "title": "A dendrite-resolved, in vivo transfer function from spike patterns to dendritic Ca2+", "authors": "Wu X, et al.", "journal": "bioRxiv", "year": 2026, "sample": "Mouse, hippocampus CA2 pyramidal neurons, in vivo", "url": "https://doi.org/10.64898/2026.01.18.700189" }, { "title": "A scalable, all-optical method for mapping synaptic connectivity with cell-type specificity", "authors": "Moya MV, et al.", "journal": "bioRxiv", "year": 2025, "sample": "Mouse, motor cortex, acute brain slices, Voltron2-JF585", "url": "https://doi.org/10.1101/2025.06.25.661552" }, { "title": "Voltage dynamics of dendritic integration and back-propagation in vivo", "authors": "Wong-Campos JD, et al.", "journal": "bioRxiv", "year": 2023, "sample": "Mouse, barrel cortex L2/3, in vivo", "url": "https://doi.org/10.1101/2023.05.25.542363" }, { "title": "High-speed multiplane confocal microscopy for voltage imaging in densely labeled neuronal populations", "authors": "Weber TD, et al.", "journal": "Nature Neuroscience", "year": 2023, "sample": "Mouse, motor cortex, acute brain slices and in vivo, Voltron2-ST JF585", "url": "https://doi.org/10.1038/s41593-023-01408-2" }, { "title": "Fast and light-efficient remote focusing for volumetric voltage imaging", "authors": "Bohm UL, Judkewitz B", "journal": "Nature Communications", "year": 2024, "sample": "Zebrafish larvae, spinal cord, in vivo", "url": "https://doi.org/10.1038/s41467-024-53685-5" }, { "title": "All-optical reporting of inhibitory receptor driving force in the nervous system", "authors": "Selfe JS, et al.", "journal": "Nature Communications", "year": 2024, "sample": "Mouse, hippocampal organotypic slices and somatosensory cortex L1, in vitro and in vivo, Voltron2-ST", "url": "https://doi.org/10.1038/s41467-024-53074-y" }, { "title": "Synchronous ensembles of hippocampal CA1 pyramidal neurons during novel exploration", "authors": "Chen EL, et al.", "journal": "eLife", "year": 2025, "sample": "Mouse, hippocampus CA1 pyramidal neurons, in vivo, Voltron2-ST JF552", "url": "https://doi.org/10.7554/eLife.96718" }, { "title": "A novel rhodopsin-based voltage indicator for simultaneous two-photon optical recording with GCaMP in vivo", "authors": "Villette V, et al.", "journal": "bioRxiv", "year": 2024, "sample": "Rat, hippocampal neurons, in vitro; mouse, cerebellum Golgi cells, in vivo, two-photon, Voltron2-ST JF552", "url": "https://doi.org/10.1101/2024.11.15.623698" }, { "title": "Positive-going hybrid indicators for voltage imaging in excitable cells and tissues", "authors": "Liu S, et al.", "journal": "Science Advances", "year": 2025, "sample": "HEK293T cells and rat hippocampal neurons, in vitro; Voltron2525 (JF525) characterized head-to-head as a comparator to HVI+ (kinetics/dynamic range/sensitivity Table S2, photobleaching Fig. S7)", "url": "https://doi.org/10.1126/sciadv.ads1807" }, { "title": "Solaris: a panel of bright and sensitive hybrid voltage indicators for imaging membrane potential in cultured neurons", "authors": "Yang J, et al.", "journal": "bioRxiv", "year": 2024, "sample": "HEK293T cells and rat hippocampal neurons, in vitro; Voltron2585 (JF585) characterized as a comparator to the Solaris dye series (dynamic range, per-AP sensitivity, kinetics)", "url": "https://doi.org/10.1101/2024.02.02.578569" }, { "title": "A photostable genetically encoded voltage indicator for imaging neural activities in tissue and live animals", "authors": "Cao C, et al.", "journal": "bioRxiv", "year": 2025, "sample": "HEK293T cells; Voltron2525 (JF525) measured as a comparator (dynamic range −27.5%/100 mV, Fig. 1C) to the Vega indicator", "url": "https://doi.org/10.1101/2025.05.29.656768" } ] }, { "id": "vsfpbutterfly", "name": "VSFP-Butterfly", "url": "https://gevibench.org/gevi/vsfpbutterfly", "description": "FRET-based voltage indicator with a novel 'Butterfly' topology sandwiching the voltage-sensing domain between two fluorescent proteins.", "category": "VSD-FRET GEVI", "tags": [ "Ci-VSP", "mCitrine/mKate2", "FRET", "Two-photon" ], "year": 2012, "date": "2012-10-15", "paper": "J. Neurophysiol. 2012", "paperUrl": "https://doi.org/10.1152/jn.00452.2012", "parent": { "id": "vsfp2", "name": "VSFP2.1" }, "crossBranchParent": null, "lineagePath": null, "polarity": "positive", "spectrum": { "peakExcitationNm": 516, "peakEmissionNm": 633, "fluorophore": "VSFP-Butterfly" }, "twoPhotonCompatible": true, "bioluminescent": false, "addgene": { "id": "47978", "url": "https://www.addgene.org/47978/", "note": null }, "lastUpdated": "2026-09-19", "display": { "brightness": "1.74×", "kinetics": "7.7 / 90", "dynamicRange": "+6%", "subthreshold": "0.042%", "sensitivity": "0.3%", "apWidth": null, "photostability": null, "papers": "28" }, "values": { "brightnessRelEGFP": 1.74, "tauOnMs": 7.7, "tauOffMs": 89.9, "dffPer100mV": 6, "dffPer100mVMedianAbs": 3.95, "dffPerMV": 0.042, "dffPerMVEstimatedFromFV": false, "dffPerAP": 0.3, "dffPerAPMedianAbs": 0.3, "apWidthMs": null, "t75At100mWmm2Sec": null, "papersUsingIt": 28 }, "measurements": { "kinetics": [ { "tauOnMs": 7.7, "tauOffMs": 89.9, "temperature": "32-34°C", "dye": null, "source": { "label": "Akemann 2012, Fig. 2E", "url": "https://doi.org/10.1152/jn.00452.2012" } } ], "dffPer100mV": [ { "dff": 6, "responseType": "steady-state", "modality": "1P", "dye": null, "source": { "label": "Mishina 2014, Table 1", "url": "https://doi.org/10.3389/fnmol.2014.00078" } }, { "dff": 1.9, "responseType": "steady-state", "modality": "1P", "dye": null, "source": { "label": "Akemann 2012, Fig. 3C", "url": "https://doi.org/10.1152/jn.00452.2012" } } ], "dffPerMV": [ { "slopePercentPerMV": 0.042, "modality": null, "note": null, "source": { "label": "Akemann 2012, Fig. 3C", "url": "https://doi.org/10.1152/jn.00452.2012" } } ], "dffPerAP": [ { "dff": 0.3, "modality": "1P", "dye": null, "source": { "label": "Akemann 2012, Fig. 3D", "url": "https://doi.org/10.1152/jn.00452.2012" } } ], "apWidth": [], "brightness": [ { "ratio": 1.74, "reference": "EGFP", "source": { "label": "doi:EC×QY: mCitrine EC=77000, QY=0.76 vs EGFP EC=56000, QY=0.60", "url": "EC×QY: mCitrine EC=77000, QY=0.76 vs EGFP EC=56000, QY=0.60" } } ], "photobleaching": [] }, "t75ScaledEntriesSec": [], "fvCurve": { "voltageMV": [ -140, -120, -100, -80, -70, -60, -40, -20, 0, 20, 40, 60 ], "dffPercent": [ -9, -8, -6, -2, 0, 2, 5, 6, 6, 6, 6, 6 ], "source": { "label": "Akemann 2012, Fig. 2C", "url": "https://doi.org/10.1152/jn.00452.2012" } }, "researchPapers": [ { "title": "Imaging neural circuit dynamics with a voltage-sensitive fluorescent protein", "authors": "Akemann W, Mutoh H, Perron A, Park YK, Iwamoto Y, Knöpfel T", "journal": "Journal of Neurophysiology", "year": 2012, "sample": "Mouse, somatosensory cortex layer 2/3, in vivo; hippocampus CA3, brain slices; PC12 cells, in vitro", "url": "https://doi.org/10.1152/jn.00452.2012" }, { "title": "Exploration of genetically encoded voltage indicators based on a chimeric voltage sensing domain", "authors": "Mishina Y, Mutoh H, Song C, Knöpfel T", "journal": "Frontiers in Molecular Neuroscience", "year": 2014, "sample": "Mouse, somatosensory cortex, in vivo; PC12 and HEK293T cells, in vitro", "url": "https://doi.org/10.3389/fnmol.2014.00078" }, { "title": "Two-photon voltage imaging using a genetically encoded voltage indicator", "authors": "Akemann W, et al.", "journal": "Scientific Reports", "year": 2013, "sample": "Mouse, cortex layer 2/3, brain slices; mouse, somatosensory cortex, in vivo", "url": "https://doi.org/10.1038/srep02231" }, { "title": "Voltage imaging of waking mouse cortex reveals emergence of critical neuronal dynamics", "authors": "Scott G, et al.", "journal": "Journal of Neuroscience", "year": 2014, "sample": "Mouse, cortex, in vivo", "url": "https://doi.org/10.1523/JNEUROSCI.3474-14.2014" }, { "title": "Imaging the awake visual cortex with a genetically encoded voltage indicator", "authors": "Carandini M, et al.", "journal": "Journal of Neuroscience", "year": 2015, "sample": "Mouse, visual cortex, in vivo, awake", "url": "https://doi.org/10.1523/JNEUROSCI.0594-14.2015" }, { "title": "Validation of optical voltage reporting by the genetically encoded voltage indicator VSFP-Butterfly from cortical layer 2/3 pyramidal neurons in mouse brain slices", "authors": "Empson RM, et al.", "journal": "Physiological Reports", "year": 2015, "sample": "Mouse, cortex layer 2/3, brain slices", "url": "https://doi.org/10.14814/phy2.12468" }, { "title": "Cortical entropy, mutual information and scale-free dynamics in waking mice", "authors": "Fagerholm ED, et al.", "journal": "Cerebral Cortex", "year": 2016, "sample": "Mouse, cortex, in vivo", "url": "https://doi.org/10.1093/cercor/bhw200" }, { "title": "Transgenic mice for intersectional targeting of neural sensors and effectors with high specificity and performance", "authors": "Madisen L, et al.", "journal": "Neuron", "year": 2015, "sample": "Mouse, cortex, transgenic Ai78(TITL-VSFPB) line, in vivo", "url": "https://doi.org/10.1016/j.neuron.2015.02.022" }, { "title": "Transgenic strategies for sparse but strong expression of genetically encoded voltage and calcium indicators", "authors": "Song C, et al.", "journal": "International Journal of Molecular Sciences", "year": 2017, "sample": "Mouse, cortex layer 2/3, transgenic", "url": "https://doi.org/10.3390/ijms18071461" }, { "title": "Imaging of brain slices with a genetically encoded voltage indicator", "authors": "Quicke P, et al.", "journal": "Methods in Molecular Biology", "year": 2017, "sample": "Mouse, cortex layer 2/3, brain slices", "url": "https://doi.org/10.1007/978-1-4939-6810-7_5" }, { "title": "Cortical signatures of wakeful somatosensory processing", "authors": "Song C, et al.", "journal": "Scientific Reports", "year": 2018, "sample": "Mouse, somatosensory cortex, in vivo, awake", "url": "https://doi.org/10.1038/s41598-018-30422-9" }, { "title": "Effects of arousal on mouse sensory cortex depend on modality", "authors": "Shimaoka D, et al.", "journal": "Cell Reports", "year": 2018, "sample": "Mouse, visual cortex; barrel cortex; auditory cortex, in vivo, awake", "url": "https://doi.org/10.1016/j.celrep.2018.02.092" }, { "title": "Optogenetic sensors in the zebrafish heart: a novel in vivo electrophysiological tool to study cardiac arrhythmogenesis", "authors": "van Opbergen CJM, et al.", "journal": "Theranostics", "year": 2018, "sample": "Zebrafish, heart, in vivo", "url": "https://doi.org/10.7150/thno.26108" }, { "title": "Slow moving neural source in the epileptic hippocampus can mimic progression of human seizures", "authors": "Chiang CC, et al.", "journal": "Scientific Reports", "year": 2018, "sample": "Mouse, hippocampal slices, in vitro", "url": "https://doi.org/10.1038/s41598-018-19925-7" }, { "title": "Single-neuron level one-photon voltage imaging with sparsely targeted genetically encoded voltage indicators", "authors": "Quicke P, et al.", "journal": "Frontiers in Cellular Neuroscience", "year": 2019, "sample": "Mouse, cortex layer 2/3, brain slices", "url": "https://doi.org/10.3389/fncel.2019.00039" }, { "title": "In vitro testing of voltage indicators: Archon1, ArcLightD, ASAP1, ASAP2s, ASAP3b, Bongwoori-Pos6, BeRST1, FlicR1, and Chi-VSFP-Butterfly", "authors": "Milosevic MM, et al.", "journal": "eNeuro", "year": 2020, "sample": "Mouse, cortex, brain slices; HEK293 cells, in vitro", "url": "https://doi.org/10.1523/ENEURO.0060-20.2020" }, { "title": "Subcellular resolution three-dimensional light-field imaging with genetically encoded voltage indicators", "authors": "Quicke P, et al.", "journal": "Neurophotonics", "year": 2020, "sample": "Mouse, brain slices, in vitro", "url": "https://doi.org/10.1117/1.NPh.7.3.035006" }, { "title": "Population imaging discrepancies between a genetically-encoded calcium indicator (GECI) versus a genetically-encoded voltage indicator (GEVI)", "authors": "Zhu MH, et al.", "journal": "Scientific Reports", "year": 2021, "sample": "Mouse, somatosensory cortex, brain slices", "url": "https://doi.org/10.1038/s41598-021-84651-6" }, { "title": "Studying synaptically evoked cortical responses ex vivo with combination of a single neuron recording (whole-cell) and population voltage imaging (genetically encoded voltage indicator)", "authors": "Jang J, et al.", "journal": "Frontiers in Neuroscience", "year": 2021, "sample": "Mouse, cortex, brain slices", "url": "https://doi.org/10.3389/fnins.2021.773883" }, { "title": "Cortex-wide dynamics of intrinsic electrical activities: propagating waves and their interactions", "authors": "Liang Y, Song C, Liu M, Gong P, Zhou C, Knöpfel T", "journal": "Journal of Neuroscience", "year": 2021, "sample": "Mouse, cortex, in vivo, anesthetized, CaMK2A-tTA;chi-VSFP transgenic", "url": "https://doi.org/10.1523/JNEUROSCI.0623-20.2021" }, { "title": "Combining cortical voltage imaging and hippocampal electrophysiology for investigating global, multi-timescale activity interactions in the brain", "authors": "Pedrosa R, et al.", "journal": "International Journal of Molecular Sciences", "year": 2022, "sample": "Mouse, cortex; hippocampus, in vivo, awake", "url": "https://doi.org/10.3390/ijms23126814" }, { "title": "Evoked cortical depolarizations before and after the amyloid plaque accumulation: voltage imaging study", "authors": "Zhu MH, Jogdand AH, Jang J, Nagella SC, Das B, Milosevic MM, Yan R, Antic SD", "journal": "Journal of Alzheimer's Disease", "year": 2022, "sample": "Mouse, cortex, brain slices, Alzheimer's disease model (APPswe/PS1dE9), chi-VSFP transgenic", "url": "https://doi.org/10.3233/JAD-220249" }, { "title": "Cortical correlates of psychedelic-induced shaking behavior revealed by voltage imaging", "authors": "Buchborn T, et al.", "journal": "International Journal of Molecular Sciences", "year": 2023, "sample": "Mouse, cortex layer 2/3, in vivo, awake", "url": "https://doi.org/10.3390/ijms24119463" }, { "title": "Complexity of cortical wave patterns of the wake mouse cortex", "authors": "Liang Y, Liang J, Song C, Liu M, Knöpfel T, Gong P, Zhou C", "journal": "Nature Communications", "year": 2023, "sample": "Mouse, cortex, in vivo, awake, CaMK2A-tTA;chi-VSFP transgenic", "url": "https://doi.org/10.1038/s41467-023-37088-6" }, { "title": "Imaging of evoked cortical depolarizations using either ASAP2s, or chi-VSFP, or Di-4-Anepps, or autofluorescence optical signals", "authors": "Milicevic KD, et al.", "journal": "Journal of Integrative Neuroscience", "year": 2023, "sample": "Mouse, cortex, brain slices", "url": "https://doi.org/10.31083/j.jin2206160" }, { "title": "Cortical networks relating to arousal are differentially coupled to neural activity and hemodynamics", "authors": "Meyer-Baese L, et al.", "journal": "Journal of Neuroscience", "year": 2024, "sample": "Mouse, cortex, in vivo, awake", "url": "https://doi.org/10.1523/JNEUROSCI.0298-23.2024" }, { "title": "Ex vivo propagation of synaptically-evoked cortical depolarizations in a mouse model of Alzheimer's disease at 20 Hz, 40 Hz, or 83 Hz", "authors": "Patel AA, Zhu MH, Yan R, Antic SD", "journal": "Scientific Reports", "year": 2024, "sample": "Mouse, cortex, brain slices, Alzheimer's disease model (5xFAD), chi-VSFP transgenic", "url": "https://doi.org/10.1038/s41598-024-74262-2" }, { "title": "Psychedelic 5-HT2A agonist increases spontaneous and evoked 5-Hz oscillations in visual and retrosplenial cortex", "authors": "White CM, Azimi Z, Staadt R, Song C, Knöpfel T, Jancke D", "journal": "Communications Biology", "year": 2026, "sample": "Mouse, visual cortex; retrosplenial cortex, in vivo, awake", "url": "https://doi.org/10.1038/s42003-025-09492-9" } ] }, { "id": "vsfp1", "name": "VSFP1", "url": "https://gevibench.org/gevi/vsfp1", "description": "The first genetically encoded voltage indicator, establishing the VSFP architecture of pairing a voltage-sensing domain with a FRET fluorescent protein pair.", "category": "VSD-FRET GEVI", "tags": [ "CFP/YFP", "FRET" ], "year": 2001, "date": "2001-12-20", "paper": "European Journal of Neuroscience 2001", "paperUrl": "https://doi.org/10.1046/j.0953-816x.2001.01617.x", "parent": null, "crossBranchParent": null, "lineagePath": [ "GEVI", "VSD", "VSD-FRET", "vsfp1" ], "polarity": "positive", "spectrum": { "peakExcitationNm": 435, "peakEmissionNm": 529, "fluorophore": "VSFP1" }, "twoPhotonCompatible": null, "bioluminescent": false, "addgene": null, "lastUpdated": "2026-09-19", "display": { "brightness": null, "kinetics": "0.74 / 0.74", "dynamicRange": "+1.8%", "subthreshold": "0.018%", "sensitivity": null, "apWidth": null, "photostability": null, "papers": "1" }, "values": { "brightnessRelEGFP": null, "tauOnMs": 0.74, "tauOffMs": 0.74, "dffPer100mV": 1.8, "dffPer100mVMedianAbs": 1.8, "dffPerMV": 0.018, "dffPerMVEstimatedFromFV": false, "dffPerAP": null, "dffPerAPMedianAbs": null, "apWidthMs": null, "t75At100mWmm2Sec": null, "papersUsingIt": 1 }, "measurements": { "kinetics": [ { "tauOnMs": 0.74, "tauOffMs": 0.74, "temperature": "25-28°C", "dye": null, "source": { "label": "Sakai 2001, Fig. 3E", "url": "https://doi.org/10.1046/j.0953-816x.2001.01617.x" } } ], "dffPer100mV": [ { "dff": 1.8, "responseType": "steady-state", "modality": "1P", "dye": null, "source": { "label": "Sakai 2001, Fig. 3B", "url": "https://doi.org/10.1046/j.0953-816x.2001.01617.x" } } ], "dffPerMV": [ { "slopePercentPerMV": 0.018, "modality": null, "note": "Linear F-V slope = 1.8 ± 0.1%/100 mV = 0.018 %/mV (Sakai 2001 Fig. 3B, r=0.99, n=11). 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