Local fitness landscape of the green fluorescent protein.


Abstract

Fitness landscapes depict how genotypes manifest at the phenotypic level and form the basis of our understanding of many areas of biology, yet their properties remain elusive. Previous studies have analysed specific genes, often using their function as a proxy for fitness, experimentally assessing the effect on function of single mutations and their combinations in a specific sequence or in different sequences. However, systematic high-throughput studies of the local fitness landscape of an entire protein have not yet been reported. Here we visualize an extensive region of the local fitness landscape of the green fluorescent protein from Aequorea victoria (avGFP) by measuring the native function (fluorescence) of tens of thousands of derivative genotypes of avGFP. We show that the fitness landscape of avGFP is narrow, with 3/4 of the derivatives with a single mutation showing reduced fluorescence and half of the derivatives with four mutations being completely non-fluorescent. The narrowness is enhanced by epistasis, which was detected in up to 30% of genotypes with multiple mutations and mostly occurred through the cumulative effect of slightly deleterious mutations causing a threshold-like decrease in protein stability and a concomitant loss of fluorescence. A model of orthologous sequence divergence spanning hundreds of millions of years predicted the extent of epistasis in our data, indicating congruence between the fitness landscape properties at the local and global scales. The characterization of the local fitness landscape of avGFP has important implications for several fields including molecular evolution, population genetics and protein design.

Submission Details

ID: MPSZ3Nkk3

Submitter: Connie Wang

Submission Date: May 8, 2017, 9:52 a.m.

Version: 1

Publication Details
Sarkisyan KS;Bolotin DA;Meer MV;Usmanova DR;Mishin AS;Sharonov GV;Ivankov DN;Bozhanova NG;Baranov MS;Soylemez O;Bogatyreva NS;Vlasov PK;Egorov ES;Logacheva MD;Kondrashov AS;Chudakov DM;Putintseva EV;Mamedov IZ;Tawfik DS;Lukyanov KA;Kondrashov FA,Nature (2016) Local fitness landscape of the green fluorescent protein. PMID:27193686
Additional Information

Study Summary

Structure view and single mutant data analysis

Study data

No weblogo for data of varying length.
Colors: D E R H K S T N Q A V I L M F Y W C G P
 

Data Distribution

Studies with similar sequences (approximate matches)

Correlation with other assays (exact sequence matches)


Relevant PDB Entries

Structure ID Release Date Resolution Structure Title
6L27 2020-04-01 0.77 X-ray crystal structure of the mutant green fluorescent protein
6JGJ 2019-04-17 0.78 Crystal structure of the F99S/M153T/V163A/E222Q variant of GFP at 0.78 A
6KL0 2019-12-11 0.8 Crystal structure of the S65T/F99S/M153T/V163A variant of perdeuterated GFP at pD 7.0
6JGI 2019-04-17 0.85 Crystal structure of the S65T/F99S/M153T/V163A variant of GFP at 0.85 A
6KL1 2019-12-11 0.85 Crystal structure of the S65T/F99S/M153T/V163A variant of non-deuterated GFP at pD 8.5
6KKZ 2019-12-11 0.9 Crystal structure of the S65T/F99S/M153T/V163A variant of perdeuterated GFP at pD 8.5
2WUR 2010-05-12 0.9 Atomic resolution structure of GFP measured on a rotating anode
6JGH 2019-04-17 0.94 Crystal structure of the F99S/M153T/V163A/T203I variant of GFP at 0.94 A
4AS8 2012-10-31 1.02 X-ray structure of the cyan fluorescent protein Cerulean cryoprotected with ethylene glycol
5OXC 2017-11-29 1.02 Structure of Cerulean Fluorescent Protein at 1.02 Angstrom resolution
4LQT 2013-12-18 1.1 1.10A resolution crystal structure of a superfolder green fluorescent protein (W57A) mutant
4GF6 2012-08-29 1.1 crystal structure of GFP with cuprum bound at the Incorporated metal Chelating Amino Acid PYZ151
4KAG 2014-08-06 1.12 Crystal structure analysis of a single amino acid deletion mutation in EGFP
6FWW 2018-08-01 1.13 GFP/KKK. A redesigned GFP with improved solubility
5DRF 2016-07-27 1.14 Green/cyan WasCFP-pH5.5 at pH 5.5
5DRG 2016-07-27 1.14 Green/cyan WasCFP at pH 10.0
6OFK 2019-07-10 1.15 Crystal structure of green fluorescent protein (GFP); S65T; ih circular permutant (50-51)
2AWK 2006-04-18 1.15 GFP R96M mature chromophore
2WSO 2009-09-29 1.15 Structure of Cerulean Fluorescent Protein at physiological pH
3SRY 2012-07-11 1.16 Engineered high-affinity halide-binding protein derived from YFP: halide-free
5OXA 2017-11-29 1.16 Structure of the S205A mutant of the Cyan Fluorescent Protein Cerulean at pH 7.0
6OGC 2020-04-08 1.18 Crystal structure of Green Fluorescent Protein (GFP); S65T, Y66(2,3-F2Y), H148D; circular permutant (50-51)
3ST0 2012-07-11 1.19 Engineered medium-affinity halide-binding protein derived from YFP: halide-free
4L1I 2013-12-11 1.2 Crystal structure of EGFP-based Calcium Sensor CatchER complexed with Ca
4XOV 2015-09-09 1.2 Structure of rsGreen0.7 in the green-off-state
2G2S 2006-04-18 1.2 Structure of S65G Y66S GFP variant after spontaneous peptide hydrolysis
2HQZ 2008-04-08 1.2 Crystal structure of L42H design intermediate for GFP metal ion reporter
2QZ0 2009-04-21 1.2 Mature Q183E variant of Green Fluorescent Protein Chromophore
2DUH 2007-07-10 1.2 crystal structure of a green fluorescent protein variant S65T/H148N at pH 9.5
2HFC 2007-03-27 1.2 Structure of S65T Y66F R96A GFP variant in precursor state
4GES 2012-08-29 1.23 crystal structure of GFP-TYR151PYZ with an unnatural amino acid incorporation
3DQ7 2008-09-23 1.23 Structure of the Yellow Fluorescent Protein Citrine Frozen at 1920 Atmospheres Number 1: Structure 17 in a Series of 26 High Pressure Structures
4AR7 2012-08-08 1.23 X-ray structure of the cyan fluorescent protein mTurquoise
2DUE 2007-07-10 1.24 crystal structure of a green fluorescent protein variant S65T/H148D at pH 10
2FZU 2006-03-14 1.25 Reduced enolate chromophore intermediate for GFP variant
2HJO 2008-04-08 1.25 Crystal structure of V224H design intermediate for GFP metal ion reporter
6OFL 2019-07-10 1.25 Crystal structure of green fluorescent protein (GFP); S65T, Y66(3-ClY); ih circular permutant (50-51)
5DQB 2016-07-27 1.25 Green/cyan WasCFP at pH 8.0
4XOW 2015-09-09 1.25 Structure of rsGreen0.7 in the green-on-state
4J8A 2013-05-15 1.26 Irradiated-state structure of sfGFP containing the unnatural amino acid p-azido-phenylalanine at residue 145
6IR7 2019-04-03 1.28 Green fluorescent protein variant GFPuv with the modification to 6-hydroxynorleucine at the C-terminus
5NI3 2018-09-12 1.28 sfGFP 204-204 mutant dimer
3SSH 2012-07-11 1.28 Engineered high-affinity halide-binding protein derived from YFP: chloride complex
3LA1 2011-01-12 1.29 High resolution crystal structure of CyPet mutant A167I
5OX8 2017-11-29 1.29 Structure of Enhanced Cyan Fluorescent Protein at pH 5.0
6HUT 2020-01-22 1.29 GFP8 - a stabilized variant of cycle-3 GFP
5DQM 2016-07-27 1.3 Green/cyan WasCFP at pH 2.0
2G6E 2006-04-18 1.3 Structure of cyclized F64L S65A Y66S GFP variant
3W1C 2013-10-02 1.3 Structure of a pressure sensitive YFP variant YFP-G1
3SVB 2012-07-18 1.3 Engineered medium-affinity halide-binding protein derived from YFP: fluoride complex
3CD1 2008-08-19 1.31 Development of a family of redox-sensitive green fluorescent protein indicators for use in relatively oxidizing subcellular environments
3ZTF 2012-03-21 1.31 X-ray Structure of the Cyan Fluorescent Protein mTurquoise2 (K206A mutant)
3CB9 2008-08-19 1.31 Development of a family of redox-sensitive green fluorescent protein indicators for use in relatively oxidizing subcellular environments
2QT2 2009-04-21 1.31 Cyclized-Dehydrated Intermediate of GFP Variant Q183E in Chromophore Maturation
3SVC 2012-07-18 1.31 Engineered medium-affinity halide-binding protein derived from YFP: chloride complex
4N3D 2014-08-27 1.34 Crystal structure of the dimeric variant EGFP-K162Q in P61 space group
4EUL 2012-10-03 1.35 Crystal structure of enhanced Green Fluorescent Protein to 1.35A resolution reveals alternative conformations for Glu222
2G3D 2006-04-18 1.35 Structure of S65G Y66A GFP variant after spontaneous peptide hydrolysis
1KYP 2002-04-10 1.35 Crystal Structure of an Apo Green Fluorescent Protein Zn Biosensor
1YJF 2005-02-15 1.35 Cyclized post-translational product for S65A Y66S (GFPhal) green fluorescent protein variant
5WJ3 2018-01-17 1.35 Crystal structure of green fluorescent protein Clover mutant S147C/Q204C
2HCG 2007-03-27 1.35 Structure of S65T Y66F GFP variant after cyclization, carbon-carbon bond cleavage, and oxygen incorporation reactions
6UN5 2020-10-14 1.36 Crystal structure of green fluorescent protein (GFP); S65T, Y66(2,3,5-F3Y); ih circular permutant (50-51)
3SSK 2012-07-11 1.36 Engineered high-affinity halide-binding protein derived from YFP: bromide complex
2DUI 2007-07-10 1.36 crystal structure of a green fluorescent protein variant H148D at pH 9
3I19 2010-06-30 1.36 1.4 Angstrom Crystal Structure of Fluorescent Protein Cypet
6OA8 2020-08-05 1.37 Superfolder Green Fluorescent Protein with 4-cyano-L-phenylalanine at the chromophore (position 66)
5AQB 2016-03-23 1.37 DARPin-based Crystallization Chaperones exploit Molecular Geometry as a Screening Dimension in Protein Crystallography
2WSN 2009-09-29 1.37 Structure of Enhanced Cyan Fluorescent Protein at physiological pH
5OXB 2017-11-29 1.38 Structure of blue-light irradiated Cerulean
1Q4E 2004-02-03 1.38 S65T Q80R Y145C Green Fluorescent Protein (GFP) pH 8.5
3DQD 2008-09-23 1.4 Structure of the Yellow Fluorescent Protein Citrine Frozen at 1250 Atmospheres Number 2: Structure 12 in a Series of 26 High Pressure Structures
6KRG 2020-09-09 1.4 Crystal structure of sfGFP Y182TMSiPhe
3SST 2012-07-11 1.4 Engineered low-affinity halide-binding protein derived from YFP: chloride complex
2FWQ 2006-03-14 1.4 Reduced enolate chromophore intermediate for Y66H GFP variant
3AI5 2010-09-29 1.4 Crystal structure of yeast enhanced green fluorescent protein-ubiquitin fusion protein
2HRS 2008-04-08 1.4 Crystal structure of L42H V224H design intermediate for GFP metal ion reporter
5MA4 2017-11-08 1.4 GFP-binding DARPin fusion gc_K7
3DQK 2008-09-23 1.4 Structure of the Yellow Fluorescent Protein Citrine Frozen at 1000 Atmospheres Number 2: Structure 6 in a Series of 26 High Pressure Structures
2B3P 2005-11-08 1.4 Crystal structure of a superfolder green fluorescent protein
3DQ9 2008-09-23 1.4 Structure of the Yellow Fluorescent Protein Citrine Frozen at 1500 Atmospheres Number 1: Structure 15 in a Series of 26 High Pressure Structures
2DUG 2007-07-10 1.4 crystal structure of a green fluorescent protein S65T/H148N at pH 5
3DQI 2008-09-23 1.42 Structure of the Yellow Fluorescent Protein Citrine Frozen at 1000 Atmospheres Number 4: Structure 8 in a Series of 26 High Pressure Structures
3DQU 2008-09-23 1.42 Structure of the Yellow Fluorescent Protein Citrine Frozen at 2000 Atmospheres Number 1: Structure 20 in a Series of 26 High Pressure Structures
5DPH 2016-01-13 1.42 sfGFP mutant - 149 p-cyano-L-phenylalanine
3DQE 2008-09-23 1.43 Structure of the Yellow Fluorescent Protein Citrine Frozen at 1250 Atmospheres Number 1: Structure 11 in a Series of 26 High Pressure Structures
3DQN 2008-09-23 1.44 Structure of the Yellow Fluorescent Protein Citrine Frozen at 500 Atmospheres: Structure 3 in a Series of 26 High Pressure Structures
3DQM 2008-09-23 1.44 Structure of the Yellow Fluorescent Protein Citrine Frozen at 750 Atmospheres: Structure 4 in a Series of 26 High Pressure Structures
1KYS 2002-04-10 1.44 Crystal Structure of a Zn-bound Green Fluorescent Protein Biosensor
3DQA 2008-09-23 1.44 Structure of the Yellow Fluorescent Protein Citrine Frozen at 1250 Atmospheres Number 4: Structure 14 in a Series of 26 High Pressure Structures
4H48 2013-09-18 1.45 1.45 angstrom CyPet Structure at pH7.0
3EVP 2008-12-09 1.45 crystal structure of circular-permutated EGFP
1Q4A 2004-02-03 1.45 S65T Q80R Green Fluorescent Protein (GFP) pH 8.5
5DTX 2016-01-20 1.45 Crystal structure of rsEGFP2 in the fluorescent on-state
5EHU 2015-12-23 1.45 sfGFP mutant with unnatural amino acid 4-azidoethoxy-L-phenylalanine incorporated at the 149 site
4B5Y 2013-08-28 1.45 X-ray structure of the cyan fluorescent protein mTurquoise-GL (K206A mutant) in space group C222(1)
3DQH 2008-09-23 1.45 Structure of the Yellow Fluorescent Protein Citrine Frozen at 1000 Atmospheres Number 5: Structure 9 in a Series of 26 High Pressure Structures
6PFT 2019-08-07 1.45 rsEGFP2 with a chlorinated chromophore in the non-fluorescent off-state
2O24 2007-05-15 1.45 Spectroscopic and Structural Study of the Heterotropic Linkage between Halide and Proton Ion Binding to Gfp Proteins: E2(GFP)-Cl Complex
3SSL 2012-07-11 1.45 Engineered high-affinity halide-binding protein derived from YFP: iodide complex
3DQF 2008-09-23 1.46 Structure of the Yellow Fluorescent Protein Citrine Frozen at 1000 Atmospheres Number 6: Structure 10 in a Series of 26 High Pressure Structures
6QQ8 2019-06-19 1.46 Cryogenic temperature structure of the fluorescent protein Cerulean recorded after an accumulated dose of 290 kGy
3DQ4 2008-09-23 1.47 Structure of the Yellow Fluorescent Protein Citrine Frozen at 2000 Atmospheres Number 2: Structure 20 in a Series of 26 High Pressure Structures
3DQL 2008-09-23 1.47 Structure of the Yellow Fluorescent Protein Citrine Frozen at 1000 Atmospheres Number 1: Structure 5 in a Series of 26 High Pressure Structures
2YE0 2012-03-21 1.47 X-ray structure of the cyan fluorescent protein mTurquoise (K206A mutant)
2H6V 2007-05-15 1.47 Spectroscopic and structural study of the heterotropic linkage between halide and proton ion binding to GFP proteins- E2(GFP) APO FORM
1Q4B 2004-02-03 1.48 S65T Q80R Green Fluorescent Protein (GFP) pH 5.5
6OFM 2019-07-10 1.48 Crystal structure of green fluorescent protein (GFP); S65T, Y66(3-CH3Y); ih circular permutant (50-51)
3DQC 2008-09-23 1.49 Structure of the Yellow Fluorescent Protein Citrine Frozen at 1250 Atmospheres Number 3: Structure 13 in a Series of 26 High Pressure Structures
3CBE 2008-08-19 1.49 Development of a family of redox-sensitive green fluorescent protein indicators for use in relatively oxidizing subcellular environments
3SVE 2012-07-18 1.49 Engineered low-affinity halide-binding protein derived from YFP: bromide complex
3SS0 2012-07-11 1.49 Engineered high-affinity halide-binding protein derived from YFP: fluoride complex
4P1Q 2014-07-16 1.5 GREEN FLUORESCENT PROTEIN E222H VARIANT
1KYR 2002-04-10 1.5 Crystal Structure of a Cu-bound Green Fluorescent Protein Zn Biosensor
6UN6 2020-10-14 1.5 Crystal structure of green fluorescent protein (GFP); S65T, Y66(3-NO2Y); ih circular permutant (50-51)
6QUH 2020-05-27 1.5 GHK tagged GFP variant crystal form II at 1.34A wavelength
3DQO 2008-09-23 1.5 Structure of the Yellow Fluorescent Protein Citrine Frozen at 1 Atmosphere Number 2: Structure 2 in a Series of 26 High Pressure Structures
1QYF 2003-09-30 1.5 Crystal structure of matured green fluorescent protein R96A variant
1H6R 2001-11-15 1.5 The oxidized state of a redox sensitive variant of green fluorescent protein
3CD9 2008-08-19 1.5 Development of a family of redox-sensitive green fluorescent protein indicators for use in relatively oxidizing subcellular environments
2QRF 2009-04-21 1.5 Green Fluorescent Protein: Cyclized-only Intermediate of Chromophore Maturation in the Q183E variant
1Z1Q 2005-06-21 1.5 Y66L Variant of Enhanced Green Fluorescent Protein with 374-nm Absorbing Chromophore
1S6Z 2004-05-04 1.5 Enhanced Green Fluorescent Protein Containing the Y66L Substitution
3DPX 2008-09-23 1.5 Structure of the Yellow Fluorescent Protein Citrine Frozen at 5000 Atmospheres: Structure 26 in a Series of 26 High Pressure Structures
5J2O 2016-04-13 1.5 Crystal structure of the cyan fluorescence protein Cerulean S175G mutant
2Y0G 2011-04-27 1.5 X-ray structure of Enhanced Green Fluorescent Protein (EGFP)
3DQ5 2008-09-23 1.5 Structure of the Yellow Fluorescent Protein Citrine Frozen at 1960 Atmospheres: Structure 19 in a Series of 26 High Pressure Structures
5DTZ 2016-01-27 1.5 Crystal structure of rsFolder in the fluorescent on-state
6UN4 2020-07-15 1.5 Crystal structure of rsEGFP2, Y67(3-ClY), Y107(3-ClY)
1YJ2 2005-02-15 1.5 Cyclized, non-dehydrated post-translational product for S65A Y66S H148G GFP variant
6UHJ 2020-03-18 1.5 X-ray Structure of C148 mGFP
1JBZ 2003-01-07 1.5 CRYSTAL STRUCTURE ANALYSIS OF A DUAL-WAVELENGTH EMISSION GREEN FLUORESCENT PROTEIN VARIANT AT HIGH PH
6UN7 2020-10-14 1.5 Crystal structure of green fluorescent protein (GFP); S65T, Y66(3-OMeY); ih circular permutant (50-51)
5DTY 2016-01-20 1.5 Crystal structure of rsEGFP2 in the non-fluorescent off-state
3W1D 2013-10-02 1.5 Structure of a pressure sensitive YFP variant YFP-G3
1YHH 2005-02-15 1.5 Uncyclized precursor structure of S65A Y66S G67A GFP variant
2DUF 2007-07-10 1.5 crystal structure of a green fluorescent protein variant S65T/H148D at pH 5.6
3DQJ 2008-09-23 1.51 Structure of the Yellow Fluorescent Protein Citrine Frozen at 1000 Atmospheres Number 3: Structure 7 in a Series of 26 High Pressure Structures
6PFR 2019-08-07 1.51 rsEGFP2 with a chlorinated chromophore in the fluorescent on-state
3DQ8 2008-09-23 1.51 Structure of the Yellow Fluorescent Protein Citrine Frozen at 1500 Atmospheres Number 2: Structure 16 in a Series of 26 High Pressure Structures
5N9O 2018-03-21 1.53 EGFP(enhanced green fluorescent protein) mutant - L232H
3SV5 2012-07-18 1.53 Engineered medium-affinity halide-binding protein derived from YFP: iodide complex
5MAD 2017-12-06 1.53 GFP-binding DARPin 3G61
4IK8 2014-02-05 1.55 High resolution structure of GCaMP3 dimer form 1 at pH 7.5
1Q4C 2004-02-03 1.55 S65T Q80R T203C Green Fluorescent Protein (GFP) pH 8.5
5OX9 2017-11-29 1.56 Structure of the Cyan Fluorescent Protein SCFP3A at pH 4.5
5MA9 2017-11-08 1.57 GFP-binding DARPin fusion gc_R11
4KA9 2014-05-07 1.58 Crystal structure analysis of single amino acid deletion mutations in EGFP
1Q4D 2004-02-03 1.58 S65T Q80R T203C Green Fluorescent Protein (GFP) pH 5.5
3DPW 2008-09-23 1.59 Structure of the Yellow Fluorescent Protein Citrine Frozen at 1 Atmosphere Number 1: Structure 1 in a Series of 26 High Pressure Structures
2YDZ 2012-03-21 1.59 X-ray structure of the cyan fluorescent protein SCFP3A (K206A mutant)
3AI4 2010-09-29 1.6 Crystal structure of yeast enhanced green fluorescent protein - mouse polymerase iota ubiquitin binding motif fusion protein
6OGA 2020-04-08 1.6 Crystal structure of Green Fluorescent Protein (GFP); S65T, Y66(3-Br1Y), H148D; circular permutant (50-51)
5T3I 2016-11-30 1.6 cyan fluorescence protein soaked with selenourea for 5 min
6T39 2020-02-19 1.6 Crystal structure of rsEGFP2 in its off-state determined by SFX
1Q73 2004-02-03 1.6 S65T Q80R Y145C T203C Green Fluorescent Protein (GFP) pH 8.5
2HGD 2007-03-27 1.6 Structure of S65A Y66F GFP variant with an oxidized chromophore
3DQ2 2008-09-23 1.6 Structure of the Yellow Fluorescent Protein Citrine Frozen at 4000 Atmospheres Number 1: Structure 23 in a Series of 26 High Pressure Structures
2AWJ 2006-04-18 1.6 GFP R96M pre-cyclized intermediate in chromophore formation
4LQU 2013-12-18 1.6 1.60A resolution crystal structure of a superfolder green fluorescent protein (W57G) mutant
6OG8 2020-04-08 1.6 Crystal structure of Green Fluorescent Protein (GFP); S65T, H148D with globally incorporated 3-F1Y; circular permutant (50-51)
6DQ1 2018-10-17 1.6 sfGFP N149 mutated to 4-nitro-L-phenylalanine
4KEX 2014-08-06 1.6 Crystal structure analysis of a single amino acid deletion mutation in EGFP
3DQ6 2008-09-23 1.6 Structure of the Yellow Fluorescent Protein Citrine Frozen at 1920 Atmospheres Number 2: Structure 18 in a Series of 26 High Pressure Structures
3GEX 2010-03-09 1.6 1.6 angstrom crystal structure of fluorescent protein Cypet
3G9A 2009-12-08 1.61 Green fluorescent protein bound to minimizer nanobody
6PFU 2019-08-07 1.62 rsEGFP2 with a chlorinated chromophore in the non-fluorescent off-state in a contracted unit cell
4EN1 2013-04-24 1.62 The 1.62A structure of a FRET-optimized Cerulean Fluorescent Protein
5WJ4 2018-01-17 1.63 Crystal structure of redox-sensitive green fluorescent protein Clover mutant roClover1
3SSP 2012-07-11 1.63 Engineered low-affinity halide-binding protein derived from YFP: halide-free
2YE1 2012-04-11 1.63 X-ray structure of the cyan fluorescent proteinmTurquoise-GL (K206A mutant)
6IR6 2019-04-03 1.64 Green fluorescent protein variant GFPuv with the native lysine residue at the C-terminus
6GO8 2018-12-19 1.65 Structure of GFPmut2 crystallized at pH 6
6OGB 2020-04-08 1.65 Crystal structure of Green Fluorescent Protein (GFP); S65T, Y66(3-I1Y), H148D; circular permutant (50-51)
5HBD 2017-01-04 1.65 Filamentous Assembly of Green Fluorescent Protein Supported by a C-terminal fusion of 18-residues, viewed in space group C2
6OFN 2019-07-10 1.65 Crystal structure of green fluorescent protein (GFP); S65T, T203(3-OMeY); ih circular permutant (50-51)
5MSE 2018-05-02 1.66 GFP nuclear transport receptor mimic 3B8
6QQA 2019-06-19 1.66 Room temperature structure of the fluorescent protein Cerulean recorded after an accumulated dose of 21 kGy
4L13 2013-12-11 1.66 Crystal structure of Ligand Free EGFP-based Calcium Sensor CatchER
6LR7 2020-04-29 1.67 Crystal structure of GFPuv complexed with the nanobody LaG16 at 1.67 Angstron resolution
6GO9 2018-12-19 1.67 Structure of GFPmut2 crystallized at pH 6 and transferred to pH 7
6QUJ 2020-05-27 1.68 GHK tagged GFP variant
5O8B 2018-01-24 1.7 Difference-refined excited-state structure of rsEGFP2 1ps following 400nm-laser irradiation of the off-state.
5O8C 2017-12-06 1.7 Composite structure of rsEGFP2 1ps following 400nm-laser irradiation of the off-state.
5O89 2017-12-13 1.7 Crystal Structure of rsEGFP2 in the fluorescent on-state determined by SFX
2QU1 2007-09-11 1.7 Crystal Structure of a Cyclized GFP Variant
6B9C 2018-10-17 1.7 Superfolder Green Fluorescent Protein with 4-nitro-L-phenylalanine at the chromophore (position 66)
3DQ1 2008-09-23 1.7 Structure of the Yellow Fluorescent Protein Citrine Frozen at 4000 Atmospheres Number 2: Structure 24 in a Series of 26 High Pressure Structures
2AWM 2006-04-18 1.7 GFP R96A chromophore maturation recovery mutant R96A Q183R
3DPZ 2008-09-23 1.7 Structure of the Yellow Fluorescent Protein Citrine Frozen at 4000 Atmospheres Number 3: Structure 25 in a Series of 26 High Pressure Structures
3DQ3 2008-09-23 1.7 Structure of the Yellow Fluorescent Protein Citrine Frozen at 2500 Atmospheres: Structure 22 in a Series of 26 High Pressure Structures
3SG6 2012-06-20 1.7 Crystal Structure of Dimeric GCaMP2-LIA(linker 1)
5MA3 2017-11-08 1.7 GFP-binding DARPin fusion gc_R11
4ZF5 2015-06-10 1.7 Crystal structure of Green Fluorescent Protein (GFP); S65T, Y66(Cl2Y), H148D; circular permutant ( 50-51)
4ORN 2015-02-11 1.71 Blue Fluorescent Protein mKalama1
6B7R 2017-12-27 1.73 Truncated strand 11-less green fluorescent protein
6AS9 2018-05-30 1.75 Filamentous Assembly of Green Fluorescent Protein Supported by a C-terminal fusion of 18-residues, viewed in space group P212121 form 2
4KW4 2014-04-09 1.75 Crystal Structure of Green Fluorescent Protein
3EVU 2008-12-09 1.75 Crystal structure of Calcium bound dimeric GCAMP2
6PFS 2019-08-07 1.76 rsEGFP2 with a chlorinated chromophore in the fluorescent on-state in a contracted unit cell
3SSY 2012-07-11 1.77 Engineered low-affinity halide-binding protein derived from YFP: iodide complex
3SVD 2012-07-18 1.78 Engineered medium-affinity halide-binding protein derived from YFP: bromide complex
4L12 2013-12-11 1.78 Crystal structure of EGFP-based Calcium Sensor CatchER complexed with Gd
6GQG 2018-12-19 1.79 Structure of GFPmut2 crystallized at pH 8.5
6OG9 2020-04-08 1.8 Crystal structure of Green Fluorescent Protein (GFP); S65T, Y66(3,5-F2Y), H148D; circular permutant (50-51)
3GJ1 2009-03-24 1.8 Non photoactivated state of PA-GFP
1HCJ 2001-12-07 1.8 Photoproduct of the wild-type Aequorea victoria Green Fluorescent Protein
1RMO 2004-06-08 1.8 Probing the Role of Tryptophans in Aequorea Victoria Green Fluorescent Proteins with an Expanded Genetic Code
1QYO 2003-09-30 1.8 Anaerobic precylization intermediate crystal structure for S65G Y66G GFP variant
2O29 2007-05-15 1.8 Spectroscopic and Structural Study of the Heterotropic Linkage between Halide and Proton Ion Binding to Gfp Proteins: E2(GFP)-BR Complex
3P28 2011-10-05 1.8 Structure of a Circular Permutant of Green Fluorescent Protein
5JZL 2017-12-06 1.8 The Structure of Monomeric Ultra Stable Green Fluorescent Protein
2G5Z 2006-04-18 1.8 Structure of S65G Y66S GFP variant after spontaneous peptide hydrolysis and decarboxylation
4IK9 2014-01-29 1.8 High resolution structure of GCaMP3 dimer form 2 at pH 7.5
4KW9 2014-04-09 1.8 Crystal Structure of Green Fluorescent Protein
1JBY 2003-01-07 1.8 CRYSTAL STRUCTURE ANALYSIS OF A DUAL-WAVELENGTH EMISSION GREEN FLUORESCENT PROTEIN VARIANT AT LOW PH
1QYQ 2003-09-30 1.8 Crystal Structure of the cyclized S65G Y66G GFP variant
4ZF4 2015-06-10 1.82 Crystal structure of Green Fluorescent Protein (GFP); S65T, Y66(Cl1Y), H148D; circular permutant (50-51)
6QQ9 2019-06-19 1.82 Cryogenic temperature structure of the fluorescent protein Cerulean recorded after an accumulated dose of 5.8 MGy
2H9W 2007-04-24 1.82 Green fluorescent protein ground states: the influence of a second protonation site near the chromophore
1W7U 2005-07-20 1.85 Photoproduct of the Wild-Type Aequorea victoria Green Fluorescent Protein after structural annealing at 170K
3EK7 2008-12-16 1.85 Calcium-saturated GCaMP2 dimer
5DPG 2016-01-13 1.85 sfGFP mutant - 133 p-cyano-L-phenylalanine
2AWL 2006-04-18 1.85 Mature R96K GFP mutant
6T3A 2020-02-19 1.85 Difference-refined structure of rsEGFP2 10 ns following 400-nm laser irradiation of the off-state determined by SFX
3UFZ 2012-10-17 1.85 Crystal structure of a Trp-less green fluorescent protein translated by the universal genetic code
5MA5 2017-11-08 1.85 GFP-binding DARPin fusion gc_K11
1W7S 2005-07-20 1.85 Wild-Type Aequorea victoria Green Fluorescent Protein
1W7T 2005-07-20 1.85 Photoproduct of the Wild-Type Aequorea victoria Green Fluorescent Protein at 100 K
5HGE 2017-01-11 1.86 Filamentous Assembly of Green Fluorescent Protein Supported by a C-terminal fusion of 18-residues, viewed in space group P212121
3SSV 2012-07-11 1.86 Engineered low-affinity halide-binding protein derived from YFP: fluoride complex
5Z6Y 2019-06-12 1.87 Structure of sfYFP48S95C66BPA
4H47 2013-09-18 1.9 1.9 angstrom CyPet structure at pH5.2
6UHK 2020-03-18 1.9 Crystal Structure of C176 mGFP
5FGU 2016-03-30 1.9 Structure of Sda1 nuclease apoprotein as an EGFP fixed-arm fusion
1EMA 1996-11-08 1.9 GREEN FLUORESCENT PROTEIN FROM AEQUOREA VICTORIA
3SG5 2012-06-20 1.9 Crystal Structure of Dimeric GCaMP3-D380Y, QP(linker 1), LP(linker 2)
6HR1 2020-04-08 1.9 Crystal structure of the YFPnano fusion protein
2OKW 2007-12-04 1.9 A non-invasive GFP-based biosensor for mercury ions
3GJ2 2009-03-24 1.9 Photoactivated state of PA-GFP
5JZK 2017-12-06 1.9 The Structure of Ultra Stable Green Fluorescent Protein
3SG7 2012-06-20 1.9 Crystal Structure of GCaMP3-KF(linker 1)
3OSQ 2011-10-26 1.9 Maltose-bound maltose sensor engineered by insertion of circularly permuted green fluorescent protein into E. coli maltose binding protein at position 175
1JC1 2003-09-09 1.9 CRYSTAL STRUCTURE ANALYSIS OF A REDOX-SENSITIVE GREEN FLUORESCENT PROTEIN VARIANT IN A OXIDIZED FORM
1GFL 1997-01-11 1.9 STRUCTURE OF GREEN FLUORESCENT PROTEIN
1RMM 2004-06-08 1.9 Probing the Role of Tryptophans in Aequorea Victoria Green Fluorescent Proteins with an Expanded Genetic Code
1YHI 2005-02-15 1.9 Uncyclized precursor structure of S65A Y66S R96A GFP variant
6B7T 2017-12-27 1.91 Truncated strand 10-less green fluorescent protein
6UHL 2020-03-18 1.91 Crystal Structure of C148 mGFP-scDNA-1
6UHN 2020-03-18 1.92 Crystal Structure of C148 mGFP-cDNA-1
6QUI 2020-05-27 1.94 GHK tagged GFP variant at 17Kev
2O2B 2007-05-15 1.94 Spectroscopic and Structural Study of the Heterotropic Linkage between Halide and Proton Ion Binding to Gfp Proteins: E2(GFP)-I Complex
3V3D 2012-12-26 1.95 Crystal Structure of an eYFP single mutant
6UHO 2020-03-18 1.95 Crystal Structure of C148 mGFP-cDNA-2
5NHN 2018-09-26 1.96 Super-Folder Green Fluorescent Protein Artificiall dimer linked via 148 position
2AHA 2006-03-21 1.98 Crystal structure analysis of a rate-enhanced variant of redox-sensitive green fluorescent protein in the reduced form, roGFP1-R8.
1Z1P 2005-06-21 2.0 Y66L variant of Enhanced Green Fluorescent Protein with 412-nm Absorbing Chromophore
2Q57 2007-11-27 2.0 X-ray structure of Cerulean GFP: A tryptophan-based chromophore useful for fluorescence lifetime imaging
1JC0 2003-09-09 2.0 CRYSTAL STRUCTURE ANALYSIS OF A REDOX-SENSITIVE GREEN FLUORESCENT PROTEIN VARIANT IN A REDUCED FORM
1QY3 2003-09-23 2.0 Crystal structure of precyclized intermediate for the green fluorescent protein R96A variant (B)
1QXT 2003-09-23 2.0 Crystal structure of precyclized intermediate for the green fluorescent protein R96A variant (A)
2G16 2006-04-18 2.0 Structure of S65A Y66S GFP variant after backbone fragmentation
3OSR 2011-10-26 2.0 Maltose-bound maltose sensor engineered by insertion of circularly permuted green fluorescent protein into E. coli maltose binding protein at position 311
3SG2 2012-06-20 2.0 Crystal Structure of GCaMP2-T116V,D381Y
3EKH 2008-12-16 2.0 Calcium-saturated GCaMP2 T116V/K378W mutant monomer
4IK1 2014-02-05 2.0 High resolution structure of GCaMPJ at pH 8.5
3EVR 2008-12-09 2.0 Crystal structure of Calcium bound monomeric GCAMP2
1EMG 1999-05-12 2.0 GREEN FLUORESCENT PROTEIN (65-67 REPLACED BY CRO, S65T SUBSTITUTION, Q80R)
2EMD 1997-08-20 2.0 GREEN FLUORESCENT PROTEIN FROM AEQUOREA VICTORIA, MUTANT
4XL5 2015-08-19 2.0 X-ray structure of bGFP-A / EGFP complex
4XBI 2015-07-29 2.01 Structure Of A Malarial Protein Involved in Proteostasis
4IK3 2014-01-29 2.01 High resolution structure of GCaMP3 at pH 8.5
4IK4 2014-02-05 2.01 High resolution structure of GCaMP3 at pH 5.0
2HGY 2007-03-27 2.05 Structure of S65A Y66F E222A GFP variant after cyclization and carbon-carbon bond cleavage
6DQ0 2018-10-17 2.05 sfGFP D133 mutated to 4-nitro-L-phenylalanine
4J88 2013-06-26 2.08 Dark-state structure of sfGFP containing the unnatural amino acid p-azido-phenylalanine at residue 66
3UG0 2012-10-17 2.09 Crystal structure of a Trp-less green fluorescent protein translated by the simplified genetic code
4J89 2013-06-26 2.1 Different photochemical events of a genetically encoded aryl azide define and modulate GFP fluorescence
3SG3 2012-06-20 2.1 Crystal Structure of GCaMP3-D380Y
1F0B 2000-11-17 2.1 CRYSTAL STRUCTURE OF THE GREEN FLUORESCENT PROTEIN (GFP) VARIANT YFP-H148Q
6UHM 2020-03-18 2.1 Crystal Structure of a Physical Mixture of C148 mGFP and scDNA-1
1EMK 1997-08-20 2.1 GREEN FLUORESCENT PROTEIN FROM AEQUOREA VICTORIA, MUTANT
1BFP 1997-07-07 2.1 BLUE VARIANT OF GREEN FLUORESCENT PROTEIN
1RRX 2004-06-08 2.1 Crystallographic Evidence for Isomeric Chromophores in 3-Fluorotyrosyl-Green Fluorescent Protein
2Q6P 2007-06-19 2.1 The Chemical Control of Protein Folding: Engineering a Superfolder Green Fluorescent Protein
1EMB 1997-06-16 2.13 GREEN FLUORESCENT PROTEIN (GFP) FROM AEQUOREA VICTORIA, GLN 80 REPLACED WITH ARG
1F09 2000-11-17 2.14 CRYSTAL STRUCTURE OF THE GREEN FLUORESCENT PROTEIN (GFP) VARIANT YFP-H148Q WITH TWO BOUND IODIDES
4Z4M 2016-04-06 2.15 Crystal structure of GFP-TAX1BP1 UBZ2 domain fusion protein
3K1K 2009-12-08 2.15 Green fluorescent protein bound to enhancer nanobody
1MYW 2003-01-14 2.2 CRYSTAL STRUCTURE OF A YELLOW FLUORESCENT PROTEIN WITH IMPROVED MATURATION AND REDUCED ENVIRONMENTAL SENSITIVITY
1HUY 2001-07-04 2.2 CRYSTAL STRUCTURE OF CITRINE, AN IMPROVED YELLOW VARIANT OF GREEN FLUORESCENT PROTEIN
4OGS 2014-05-14 2.21 Crystal structure of GFP S205A/T203V at 2.2 A resolution
2AH8 2006-03-21 2.24 roGFP1-R7. Cystal structure analysis of a rate-enhanced variant of redox-sensitive green fluorescent protein in the oxidized form.
1C4F 1999-08-31 2.25 GREEN FLUORESCENT PROTEIN S65T AT PH 4.6
4PA0 2015-07-08 2.25 Omecamtiv Mercarbil binding site on the Human Beta-Cardiac Myosin Motor Domain
6GRM 2018-12-19 2.3 Structure of GFPmut2 crystallized at pH 6 and transferred to pH 9
4U2V 2014-09-10 2.3 Bak BH3-in-Groove dimer (GFP)
1EMC 1997-08-20 2.3 GREEN FLUORESCENT PROTEIN FROM AEQUOREA VICTORIA, MUTANT
6AA2 2019-05-29 2.3 X-ray structure of ReQy1 (oxidized form)
2EMN 1997-08-20 2.3 GREEN FLUORESCENT PROTEIN FROM AEQUOREA VICTORIA, MUTANT
5MA6 2017-12-06 2.3 GFP-binding DARPin 3G124nc
2B3Q 2005-11-08 2.3 Crystal structure of a well-folded variant of green fluorescent protein
1EMM 1997-08-20 2.3 GREEN FLUORESCENT PROTEIN FROM AEQUOREA VICTORIA, MUTANT
1EML 1997-08-20 2.3 GREEN FLUORESCENT PROTEIN FROM AEQUOREA VICTORIA, MUTANT
5DU0 2016-01-27 2.35 Crystal structure of rsFolder in the non-fluorescent off-state
5MA8 2017-12-06 2.35 GFP-binding DARPin 3G124nc
3O77 2010-09-29 2.35 The structure of Ca2+ Sensor (Case-16)
6AA6 2019-05-29 2.39 X-ray structure of ReQy1 (reduced form)
1B9C 2000-11-17 2.4 Green Fluorescent Protein Mutant F99S, M153T and V163A
6GQH 2018-12-19 2.4 Structure of GFPmut2 crystallized at pH 8.5 and transferred to pH 6
3VHT 2012-10-10 2.4 Crystal structure of GFP-Wrnip1 UBZ domain fusion protein in complex with ubiquitin
2OKY 2007-12-04 2.4 A non-invasive GFP-based biosensor for mercury ions
3SG4 2012-06-20 2.4 Crystal Structure of GCaMP3-D380Y, LP(linker 2)
7BWN 2020-06-17 2.4 Crystal Structure of a Designed Protein Heterocatenane
6EFR 2019-01-23 2.4 Crystal Structure of iNicSnFR 1.0
1EMF 1997-08-20 2.4 GREEN FLUORESCENT PROTEIN FROM AEQUOREA VICTORIA, MUTANT
4JRB 2013-07-24 2.41 Structure of Cockroach Allergen Bla g 1 Tandem Repeat as a EGFP fusion
5WJ2 2018-01-17 2.41 Crystal structure of the green fluorescent protein Clover
6QQB 2019-06-19 2.45 Room temperature structure of the fluorescent protein Cerulean recorded after an accumulated dose of 147 kGy
5J3N 2017-04-12 2.45 C-terminal domain of EcoR124I HsdR subunit fused with the pH-sensitive GFP variant ratiometric pHluorin
4KW8 2014-04-09 2.46 Crystal Structure of Green Fluorescent Protein
6GEZ 2019-08-21 2.47 THE STRUCTURE OF TWITCH-2B N532F
3WLC 2014-01-22 2.49 Crystal structure of dimeric GCaMP6m
5HZO 2016-02-17 2.49 GFP mutant S205G
1EME 1997-08-20 2.5 GREEN FLUORESCENT PROTEIN FROM AEQUOREA VICTORIA, MUTANT
1YHG 2005-02-15 2.5 Uncyclized precursor structure of S65G Y66S V68G GFP variant
4IK5 2014-01-29 2.5 High resolution structure of Delta-REST-GCaMP3
1YFP 1998-10-28 2.5 STRUCTURE OF YELLOW-EMISSION VARIANT OF GFP
1CV7 2003-09-16 2.5 Crystal structure of enhanced cyan-emission variant of GFP
5DPJ 2016-02-03 2.5 sfGFP double mutant - 133/149 p-ethynyl-L-phenylalanine
7BYL 2020-12-02 2.5 Cryo-EM structure of human KCNQ4
5MAK 2017-11-08 2.5 GFP-binding DARPin fusion gc_R7
6GEL 2019-08-21 2.51 The structure of TWITCH-2B
5DPI 2016-01-13 2.54 sfGFP double mutant - 133/149 p-cyano-L-phenylalanine
4LW5 2014-02-05 2.55 Crystal structure of all-trans green fluorescent protein
4PFE 2015-06-24 2.6 Crystal structure of vsfGFP-0
3O78 2010-09-29 2.6 The structure of Ca2+ Sensor (Case-12)
4KF5 2013-12-18 2.6 Crystal Structure of Split GFP complexed with engineered sfCherry with an insertion of GFP fragment
3EVV 2008-12-09 2.6 Crystal Structure of Calcium bound dimeric GCAMP2 (#2)
2EMO 1997-08-20 2.6 GREEN FLUORESCENT PROTEIN FROM AEQUOREA VICTORIA, MUTANT
4ANJ 2012-10-17 2.6 MYOSIN VI (MDinsert2-GFP fusion) PRE-POWERSTROKE STATE (MG.ADP.AlF4)
1KP5 2002-08-28 2.6 Cyclic Green Fluorescent Protein
6OFO 2019-07-10 2.6 Crystal structure of split green fluorescent protein (GFP); s10 circular permutant (194-195)
2YFP 1999-01-13 2.6 STRUCTURE OF YELLOW-EMISSION VARIANT OF GFP
3EK4 2008-12-16 2.65 Calcium-saturated GCaMP2 Monomer
2JAD 2006-12-20 2.7 Yellow fluorescent protein - glutaredoxin fusion protein
3WLD 2014-01-22 2.7 Crystal structure of monomeric GCaMP6m
3ED8 2009-01-20 2.7 Application of the superfolder YFP bimolecular fluorescence complementation for studying protein-protein interactions in vitro
3U8P 2012-08-29 2.75 Cytochrome b562 integral fusion with EGFP
5F9G 2016-02-03 2.77 pnGFP1.5-Y.Cro: circularly permuted green fluorescent protein (with a tyrosine-derived chromophore)
5KTG 2016-08-17 2.8 Crystal structure of mouse Bak BH3-in-groove homodimer (GFP)
3EK8 2008-12-16 2.8 Calcium-saturated GCaMP2 T116V/G87R mutant monomer
4Z4K 2016-04-06 2.8 Crystal structure of GFP-TAX1BP1 UBZ1+2 domain fusion protein
3EKJ 2008-12-16 2.8 Calcium-free GCaMP2 (calcium binding deficient mutant)
3OGO 2010-08-25 2.8 Structure of the GFP:GFP-nanobody complex at 2.8 A resolution in spacegroup P21212
6UHQ 2020-03-18 2.85 Crystal Structure of C148 mGFP-cDNA-3
6UHP 2020-03-18 2.9 Crystal Structure of C148 mGFP-ncDNA-1
6WVG 2020-09-16 2.9 human CD53
1RM9 2004-06-08 2.9 Probing the Role of Tryptophans in Aequorea Victoria Green Fluorescent Proteins with an Expanded Genetic Code
5LEM 2017-08-02 2.98 Crystal structure of DARPin-DARPin rigid fusion, variant DD_Off7_11_3G124 in complex with Maltose-binding Protein and Green Fluorescent Protein
6LNP 2020-05-06 2.99 Crystal structure of citrate Biosensor
4JFG 2013-10-02 3.0 Crystal structure of sfGFP-66-HqAla
5HW9 2017-02-01 3.0 Filamentous Assembly of Green Fluorescent Protein Supported by a C-terminal fusion of 18-residues, viewed in space group P21
6UHR 2020-03-18 3.0 Crystal Structure of C148 mGFP-scDNA-2
6MWQ 2018-11-21 3.0 Single particle cryoEM structure of a DARPin-aldolase platform in complex with GFP
4BDU 2013-02-13 3.0 Bax BH3-in-Groove dimer (GFP)
1RMP 2004-06-08 3.0 Probing the Role of Tryptophans in Aequorea Victoria Green Fluorescent Proteins with an Expanded Genetic Code
6T90 2020-05-06 3.05 OCT4-SOX2-bound nucleosome - SHL-6
5LEL 2017-11-15 3.1 Crystal structure of DARPin-DARPin rigid fusion, variant DD_Off7_10_3G124 in complex with Maltose-binding Protein and Green Fluorescent Protein
7BYM 2020-12-02 3.1 Cryo-EM structure of human KCNQ4 with retigabine
5B61 2017-06-14 3.12 Extra-superfolder GFP
6WRF 2020-11-04 3.14 ClpX-ClpP complex bound to GFP-ssrA, recognition complex
6WSG 2020-11-04 3.16 ClpX-ClpP complex bound to ssrA-tagged GFP, intermediate complex
4P7H 2014-05-21 3.2 Structure of Human beta-Cardiac Myosin Motor Domain::GFP chimera
6UZ0 2020-01-01 3.24 Cardiac sodium channel with flecainide
7BYN 2020-12-02 3.3 Cryo-EM structure of human KCNQ4 with linopirdine
4XVP 2015-08-19 3.4 X-ray structure of bGFP-C / EGFP complex
6MKS 2018-11-07 3.4 Cryo-EM structure of NLRC4-CARD filament
6YOV 2020-05-06 3.42 OCT4-SOX2-bound nucleosome - SHL+6
6ITC 2019-06-12 3.45 Structure of a substrate engaged SecA-SecY protein translocation machine
6MDR 2019-01-23 3.47 Cryo-EM structure of the Ceru+32/GFP-17 protomer
6UZ3 2020-01-01 3.5 Cardiac sodium channel
6VAM 2020-01-29 3.63 Cryo-EM structure of octameric chicken CALHM1
6VAL 2020-01-29 3.87 Cryo-EM structure of an undecameric chicken CALHM1 and human CALHM2 chimera
6MB2 2018-09-05 5.0 Cryo-EM structure of the PYD filament of AIM2

Relevant UniProtKB Entries

Percent Identity Matching Chains Protein Accession Entry Name
100.0 Green fluorescent protein P42212 GFP_AEQVI