Mispacking and the Fitness Landscape of the Green Fluorescent Protein Chromophore Milieu.

Abstract

The autocatalytic maturation of the chromophore in green fluorescent protein (GFP) was thought to require the precise positioning of the side chains surrounding it in the core of the protein, many of which are strongly conserved among homologous fluorescent proteins. In this study, we screened for green fluorescence in an exhaustive set of point mutations of seven residues that make up the chromophore microenvironment, excluding R96 and E222 because mutations at these positions have been previously characterized. Contrary to expectations, nearly all amino acids were tolerated at all seven positions. Only four point mutations knocked out fluorescence entirely. However, chromophore maturation was found to be slower and/or fluorescence reduced in several cases. Selected combinations of mutations showed nonadditive effects, including cooperativity and rescue. The results provide guidelines for the computational engineering of GFPs.

Submission Details

ID: B7Nwc2uR

Submitter: Connie Wang

Submission Date: May 12, 2017, 12:50 p.m.

Version: 2

Publication Details
Banerjee S;Schenkelberg CD;Jordan TB;Reimertz JM;Crone EE;Crone DE;Bystroff C,Biochemistry (2017) Mispacking and the Fitness Landscape of the Green Fluorescent Protein Chromophore Milieu. PMID:28074648
Additional Information

Study Summary

Number of data points 134
Proteins Green fluorescent protein (mutated to super folder GFP, see PDB)
Unique complexes 134
Assays/Quantities/Protocols Experimental Assay: In Vivo Relative Fluorescence
Libraries Single point mutants of sfGFP at 7 positions near the chromophore

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
2WUR 2010-05-12 0.9 Atomic resolution structure of GFP measured on a rotating anode
6KKZ 2019-12-11 0.9 Crystal structure of the S65T/F99S/M153T/V163A variant of perdeuterated GFP at pD 8.5
6JGH 2019-04-17 0.94 Crystal structure of the F99S/M153T/V163A/T203I variant of GFP at 0.94 A
5OXC 2017-11-29 1.02 Structure of Cerulean Fluorescent Protein at 1.02 Angstrom resolution
4AS8 2012-10-31 1.02 X-ray structure of the cyan fluorescent protein Cerulean cryoprotected with ethylene glycol
4GF6 2012-08-29 1.1 crystal structure of GFP with cuprum bound at the Incorporated metal Chelating Amino Acid PYZ151
4LQT 2013-12-18 1.1 1.10A resolution crystal structure of a superfolder green fluorescent protein (W57A) mutant
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
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
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
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
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
4XOW 2015-09-09 1.25 Structure of rsGreen0.7 in the green-on-state
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
5DQB 2016-07-27 1.25 Green/cyan WasCFP at pH 8.0
6OFL 2019-07-10 1.25 Crystal structure of green fluorescent protein (GFP); S65T, Y66(3-ClY); ih circular permutant (50-51)
4J8A 2013-05-15 1.26 Irradiated-state structure of sfGFP containing the unnatural amino acid p-azido-phenylalanine at residue 145
5NI3 2018-09-12 1.28 sfGFP 204-204 mutant dimer
6IR7 2019-04-03 1.28 Green fluorescent protein variant GFPuv with the modification to 6-hydroxynorleucine at the C-terminus
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
3SVB 2012-07-18 1.3 Engineered medium-affinity halide-binding protein derived from YFP: fluoride complex
3W1C 2013-10-02 1.3 Structure of a pressure sensitive YFP variant YFP-G1
2G6E 2006-04-18 1.3 Structure of cyclized F64L S65A Y66S GFP variant
5DQM 2016-07-27 1.3 Green/cyan WasCFP at pH 2.0
3SVC 2012-07-18 1.31 Engineered medium-affinity halide-binding protein derived from YFP: chloride 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)
2QT2 2009-04-21 1.31 Cyclized-Dehydrated Intermediate of GFP Variant Q183E in Chromophore Maturation
3CB9 2008-08-19 1.31 Development of a family of redox-sensitive green fluorescent protein indicators for use in relatively oxidizing subcellular environments
4N3D 2014-08-27 1.34 Crystal structure of the dimeric variant EGFP-K162Q in P61 space group
1YJF 2005-02-15 1.35 Cyclized post-translational product for S65A Y66S (GFPhal) green fluorescent protein variant
1KYP 2002-04-10 1.35 Crystal Structure of an Apo Green Fluorescent Protein Zn Biosensor
2HCG 2007-03-27 1.35 Structure of S65T Y66F GFP variant after cyclization, carbon-carbon bond cleavage, and oxygen incorporation reactions
5WJ3 2018-01-17 1.35 Crystal structure of green fluorescent protein Clover mutant S147C/Q204C
2G3D 2006-04-18 1.35 Structure of S65G Y66A GFP variant after spontaneous peptide hydrolysis
4EUL 2012-10-03 1.35 Crystal structure of enhanced Green Fluorescent Protein to 1.35A resolution reveals alternative conformations for Glu222
2DUI 2007-07-10 1.36 crystal structure of a green fluorescent protein variant H148D at pH 9
3SSK 2012-07-11 1.36 Engineered high-affinity halide-binding protein derived from YFP: bromide complex
3I19 2010-06-30 1.36 1.4 Angstrom Crystal Structure of Fluorescent Protein Cypet
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
3AI5 2010-09-29 1.4 Crystal structure of yeast enhanced green fluorescent protein-ubiquitin fusion protein
2DUG 2007-07-10 1.4 crystal structure of a green fluorescent protein S65T/H148N at pH 5
2FWQ 2006-03-14 1.4 Reduced enolate chromophore intermediate for Y66H GFP variant
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
3SST 2012-07-11 1.4 Engineered low-affinity halide-binding protein derived from YFP: chloride complex
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
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
2B3P 2005-11-08 1.4 Crystal structure of a superfolder green fluorescent protein
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
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
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
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
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
1KYS 2002-04-10 1.44 Crystal Structure of a Zn-bound Green Fluorescent Protein Biosensor
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
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
4B5Y 2013-08-28 1.45 X-ray structure of the cyan fluorescent protein mTurquoise-GL (K206A mutant) in space group C222(1)
3EVP 2008-12-09 1.45 crystal structure of circular-permutated EGFP
3SSL 2012-07-11 1.45 Engineered high-affinity halide-binding protein derived from YFP: iodide complex
5EHU 2015-12-23 1.45 sfGFP mutant with unnatural amino acid 4-azidoethoxy-L-phenylalanine incorporated at the 149 site
4H48 2013-09-18 1.45 1.45 angstrom CyPet Structure at pH7.0
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
1Q4A 2004-02-03 1.45 S65T Q80R Green Fluorescent Protein (GFP) pH 8.5
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
5DTX 2016-01-20 1.45 Crystal structure of rsEGFP2 in the fluorescent on-state
6PFT 2019-08-07 1.45 rsEGFP2 with a chlorinated chromophore in the non-fluorescent off-state
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
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
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
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)
3SS0 2012-07-11 1.49 Engineered high-affinity halide-binding protein derived from YFP: fluoride complex
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
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
3W1D 2013-10-02 1.5 Structure of a pressure sensitive YFP variant YFP-G3
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
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
5DTZ 2016-01-27 1.5 Crystal structure of rsFolder in the fluorescent on-state
2DUF 2007-07-10 1.5 crystal structure of a green fluorescent protein variant S65T/H148D at pH 5.6
1YHH 2005-02-15 1.5 Uncyclized precursor structure of S65A Y66S G67A GFP variant
1JBZ 2003-01-07 1.5 CRYSTAL STRUCTURE ANALYSIS OF A DUAL-WAVELENGTH EMISSION GREEN FLUORESCENT PROTEIN VARIANT AT HIGH PH
6QUH 2020-05-27 1.5 GHK tagged GFP variant crystal form II at 1.34A wavelength
1H6R 2001-11-15 1.5 The oxidized state of a redox sensitive variant of green fluorescent protein
5DTY 2016-01-20 1.5 Crystal structure of rsEGFP2 in the non-fluorescent off-state
1Z1Q 2005-06-21 1.5 Y66L Variant of Enhanced Green Fluorescent Protein with 374-nm Absorbing Chromophore
4P1Q 2014-07-16 1.5 GREEN FLUORESCENT PROTEIN E222H VARIANT
1QYF 2003-09-30 1.5 Crystal structure of matured green fluorescent protein R96A variant
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
2Y0G 2011-04-27 1.5 X-ray structure of Enhanced Green Fluorescent Protein (EGFP)
1KYR 2002-04-10 1.5 Crystal Structure of a Cu-bound Green Fluorescent Protein Zn Biosensor
3CD9 2008-08-19 1.5 Development of a family of redox-sensitive green fluorescent protein indicators for use in relatively oxidizing subcellular environments
5J2O 2016-04-13 1.5 Crystal structure of the cyan fluorescence protein Cerulean S175G mutant
2QRF 2009-04-21 1.5 Green Fluorescent Protein: Cyclized-only Intermediate of Chromophore Maturation in the Q183E variant
1S6Z 2004-05-04 1.5 Enhanced Green Fluorescent Protein Containing the Y66L Substitution
1YJ2 2005-02-15 1.5 Cyclized, non-dehydrated post-translational product for S65A Y66S H148G GFP variant
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
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
5N9O 2018-03-21 1.53 EGFP(enhanced green fluorescent protein) mutant - L232H
5MAD 2017-12-06 1.53 GFP-binding DARPin 3G61
3SV5 2012-07-18 1.53 Engineered medium-affinity halide-binding protein derived from YFP: iodide complex
1Q4C 2004-02-03 1.55 S65T Q80R T203C Green Fluorescent Protein (GFP) pH 8.5
4IK8 2014-02-05 1.55 High resolution structure of GCaMP3 dimer form 1 at pH 7.5
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
1Q4D 2004-02-03 1.58 S65T Q80R T203C Green Fluorescent Protein (GFP) pH 5.5
4KA9 2014-05-07 1.58 Crystal structure analysis of single amino acid deletion mutations in EGFP
2YDZ 2012-03-21 1.59 X-ray structure of the cyan fluorescent protein SCFP3A (K206A mutant)
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
6DQ1 2018-10-17 1.6 sfGFP N149 mutated to 4-nitro-L-phenylalanine
4LQU 2013-12-18 1.6 1.60A resolution crystal structure of a superfolder green fluorescent protein (W57G) mutant
2HGD 2007-03-27 1.6 Structure of S65A Y66F GFP variant with an oxidized chromophore
3AI4 2010-09-29 1.6 Crystal structure of yeast enhanced green fluorescent protein - mouse polymerase iota ubiquitin binding motif fusion protein
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
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
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
2AWJ 2006-04-18 1.6 GFP R96M pre-cyclized intermediate in chromophore formation
3GEX 2010-03-09 1.6 1.6 angstrom crystal structure of fluorescent protein Cypet
4KEX 2014-08-06 1.6 Crystal structure analysis of a single amino acid deletion mutation in EGFP
5T3I 2016-11-30 1.6 cyan fluorescence protein soaked with selenourea for 5 min
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
2YE1 2012-04-11 1.63 X-ray structure of the cyan fluorescent proteinmTurquoise-GL (K206A mutant)
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
6IR6 2019-04-03 1.64 Green fluorescent protein variant GFPuv with the native lysine residue at the C-terminus
6OFN 2019-07-10 1.65 Crystal structure of green fluorescent protein (GFP); S65T, T203(3-OMeY); ih 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
6GO8 2018-12-19 1.65 Structure of GFPmut2 crystallized at pH 6
6QQA 2019-06-19 1.66 Room temperature structure of the fluorescent protein Cerulean recorded after an accumulated dose of 21 kGy
5MSE 2018-05-02 1.66 GFP nuclear transport receptor mimic 3B8
4L13 2013-12-11 1.66 Crystal structure of Ligand Free EGFP-based Calcium Sensor CatchER
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
2AWM 2006-04-18 1.7 GFP R96A chromophore maturation recovery mutant R96A Q183R
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
5O8C 2017-12-06 1.7 Composite structure of rsEGFP2 1ps following 400nm-laser irradiation of the off-state.
3SG6 2012-06-20 1.7 Crystal Structure of Dimeric GCaMP2-LIA(linker 1)
6B9C 2018-10-17 1.7 Superfolder Green Fluorescent Protein with 4-nitro-L-phenylalanine at the chromophore (position 66)
5O89 2017-12-13 1.7 Crystal Structure of rsEGFP2 in the fluorescent on-state determined by SFX
4ZF5 2015-06-10 1.7 Crystal structure of Green Fluorescent Protein (GFP); S65T, Y66(Cl2Y), H148D; circular permutant ( 50-51)
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
5MA3 2017-11-08 1.7 GFP-binding DARPin fusion gc_R11
5O8B 2018-01-24 1.7 Difference-refined excited-state structure of rsEGFP2 1ps following 400nm-laser irradiation of the off-state.
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
2QU1 2007-09-11 1.7 Crystal Structure of a Cyclized GFP Variant
4ORN 2015-02-11 1.71 Blue Fluorescent Protein mKalama1
6B7R 2017-12-27 1.73 Truncated strand 11-less green fluorescent protein
3EVU 2008-12-09 1.75 Crystal structure of Calcium bound dimeric GCAMP2
4KW4 2014-04-09 1.75 Crystal Structure of 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
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
1QYQ 2003-09-30 1.8 Crystal Structure of the cyclized S65G Y66G GFP variant
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
3P28 2011-10-05 1.8 Structure of a Circular Permutant of Green Fluorescent Protein
4IK9 2014-01-29 1.8 High resolution structure of GCaMP3 dimer form 2 at pH 7.5
1QYO 2003-09-30 1.8 Anaerobic precylization intermediate crystal structure for S65G Y66G GFP variant
1RMO 2004-06-08 1.8 Probing the Role of Tryptophans in Aequorea Victoria Green Fluorescent Proteins with an Expanded Genetic Code
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
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
1HCJ 2001-12-07 1.8 Photoproduct of the wild-type Aequorea victoria Green Fluorescent Protein
3GJ1 2009-03-24 1.8 Non photoactivated state of PA-GFP
6QQ9 2019-06-19 1.82 Cryogenic temperature structure of the fluorescent protein Cerulean recorded after an accumulated dose of 5.8 MGy
4ZF4 2015-06-10 1.82 Crystal structure of Green Fluorescent Protein (GFP); S65T, Y66(Cl1Y), H148D; circular permutant (50-51)
2H9W 2007-04-24 1.82 Green fluorescent protein ground states: the influence of a second protonation site near the chromophore
2AWL 2006-04-18 1.85 Mature R96K GFP mutant
3UFZ 2012-10-17 1.85 Crystal structure of a Trp-less green fluorescent protein translated by the universal genetic code
3EK7 2008-12-16 1.85 Calcium-saturated GCaMP2 dimer
1W7U 2005-07-20 1.85 Photoproduct of the Wild-Type Aequorea victoria Green Fluorescent Protein after structural annealing at 170K
5MA5 2017-11-08 1.85 GFP-binding DARPin fusion gc_K11
1W7T 2005-07-20 1.85 Photoproduct of the Wild-Type Aequorea victoria Green Fluorescent Protein at 100 K
1W7S 2005-07-20 1.85 Wild-Type Aequorea victoria Green Fluorescent Protein
5DPG 2016-01-13 1.85 sfGFP mutant - 133 p-cyano-L-phenylalanine
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
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
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
4H47 2013-09-18 1.9 1.9 angstrom CyPet structure at pH5.2
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
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
3SG7 2012-06-20 1.9 Crystal Structure of GCaMP3-KF(linker 1)
1EMA 1996-11-08 1.9 GREEN FLUORESCENT PROTEIN FROM AEQUOREA VICTORIA
5JZK 2017-12-06 1.9 The Structure of Ultra Stable 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
3GJ2 2009-03-24 1.9 Photoactivated state of PA-GFP
5FGU 2016-03-30 1.9 Structure of Sda1 nuclease apoprotein as an EGFP fixed-arm fusion
6B7T 2017-12-27 1.91 Truncated strand 10-less green fluorescent protein
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
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.
1QY3 2003-09-23 2.0 Crystal structure of precyclized intermediate for the green fluorescent protein R96A variant (B)
2Q57 2007-11-27 2.0 X-ray structure of Cerulean GFP: A tryptophan-based chromophore useful for fluorescence lifetime imaging
2EMD 1997-08-20 2.0 GREEN FLUORESCENT PROTEIN FROM AEQUOREA VICTORIA, MUTANT
1EMG 1999-05-12 2.0 GREEN FLUORESCENT PROTEIN (65-67 REPLACED BY CRO, S65T SUBSTITUTION, Q80R)
1JC0 2003-09-09 2.0 CRYSTAL STRUCTURE ANALYSIS OF A REDOX-SENSITIVE GREEN FLUORESCENT PROTEIN VARIANT IN A REDUCED FORM
1Z1P 2005-06-21 2.0 Y66L variant of Enhanced Green Fluorescent Protein with 412-nm Absorbing Chromophore
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
4XL5 2015-08-19 2.0 X-ray structure of bGFP-A / EGFP complex
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
4IK1 2014-02-05 2.0 High resolution structure of GCaMPJ at pH 8.5
3EKH 2008-12-16 2.0 Calcium-saturated GCaMP2 T116V/K378W mutant monomer
3EVR 2008-12-09 2.0 Crystal structure of Calcium bound monomeric GCAMP2
4IK4 2014-02-05 2.01 High resolution structure of GCaMP3 at pH 5.0
4IK3 2014-01-29 2.01 High resolution structure of GCaMP3 at pH 8.5
4XBI 2015-07-29 2.01 Structure Of A Malarial Protein Involved in Proteostasis
6DQ0 2018-10-17 2.05 sfGFP D133 mutated to 4-nitro-L-phenylalanine
2HGY 2007-03-27 2.05 Structure of S65A Y66F E222A GFP variant after cyclization and carbon-carbon bond cleavage
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
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
2Q6P 2007-06-19 2.1 The Chemical Control of Protein Folding: Engineering a Superfolder Green Fluorescent Protein
4J89 2013-06-26 2.1 Different photochemical events of a genetically encoded aryl azide define and modulate GFP fluorescence
1F0B 2000-11-17 2.1 CRYSTAL STRUCTURE OF THE GREEN FLUORESCENT PROTEIN (GFP) VARIANT YFP-H148Q
1RRX 2004-06-08 2.1 Crystallographic Evidence for Isomeric Chromophores in 3-Fluorotyrosyl-Green Fluorescent Protein
3SG3 2012-06-20 2.1 Crystal Structure of GCaMP3-D380Y
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
3K1K 2009-12-08 2.15 Green fluorescent protein bound to enhancer nanobody
4Z4M 2016-04-06 2.15 Crystal structure of GFP-TAX1BP1 UBZ2 domain fusion protein
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
5MA6 2017-12-06 2.3 GFP-binding DARPin 3G124nc
4U2V 2014-09-10 2.3 Bak BH3-in-Groove dimer (GFP)
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
2EMN 1997-08-20 2.3 GREEN FLUORESCENT PROTEIN FROM AEQUOREA VICTORIA, MUTANT
1EMC 1997-08-20 2.3 GREEN FLUORESCENT PROTEIN FROM AEQUOREA VICTORIA, MUTANT
6GRM 2018-12-19 2.3 Structure of GFPmut2 crystallized at pH 6 and transferred to pH 9
2B3Q 2005-11-08 2.3 Crystal structure of a well-folded variant of green fluorescent protein
6AA2 2019-05-29 2.3 X-ray structure of ReQy1 (oxidized form)
5MA8 2017-12-06 2.35 GFP-binding DARPin 3G124nc
3O77 2010-09-29 2.35 The structure of Ca2+ Sensor (Case-16)
5DU0 2016-01-27 2.35 Crystal structure of rsFolder in the non-fluorescent off-state
6AA6 2019-05-29 2.39 X-ray structure of ReQy1 (reduced form)
3VHT 2012-10-10 2.4 Crystal structure of GFP-Wrnip1 UBZ domain fusion protein in complex with ubiquitin
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
6GQH 2018-12-19 2.4 Structure of GFPmut2 crystallized at pH 8.5 and transferred to pH 6
1B9C 2000-11-17 2.4 Green Fluorescent Protein Mutant F99S, M153T and V163A
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)
5WJ2 2018-01-17 2.41 Crystal structure of the green fluorescent protein Clover
4JRB 2013-07-24 2.41 Structure of Cockroach Allergen Bla g 1 Tandem Repeat as a EGFP fusion
5J3N 2017-04-12 2.45 C-terminal domain of EcoR124I HsdR subunit fused with the pH-sensitive GFP variant ratiometric pHluorin
6QQB 2019-06-19 2.45 Room temperature structure of the fluorescent protein Cerulean recorded after an accumulated dose of 147 kGy
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
5MAK 2017-11-08 2.5 GFP-binding DARPin fusion gc_R7
1CV7 2003-09-16 2.5 Crystal structure of enhanced cyan-emission variant of GFP
1YFP 1998-10-28 2.5 STRUCTURE OF YELLOW-EMISSION VARIANT OF GFP
5DPJ 2016-02-03 2.5 sfGFP double mutant - 133/149 p-ethynyl-L-phenylalanine
4IK5 2014-01-29 2.5 High resolution structure of Delta-REST-GCaMP3
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
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
3O78 2010-09-29 2.6 The structure of Ca2+ Sensor (Case-12)
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)
2EMO 1997-08-20 2.6 GREEN FLUORESCENT PROTEIN FROM AEQUOREA VICTORIA, MUTANT
4PFE 2015-06-24 2.6 Crystal structure of vsfGFP-0
3EVV 2008-12-09 2.6 Crystal Structure of Calcium bound dimeric GCAMP2 (#2)
2YFP 1999-01-13 2.6 STRUCTURE OF YELLOW-EMISSION VARIANT OF GFP
4KF5 2013-12-18 2.6 Crystal Structure of Split GFP complexed with engineered sfCherry with an insertion of GFP fragment
4ANJ 2012-10-17 2.6 MYOSIN VI (MDinsert2-GFP fusion) PRE-POWERSTROKE STATE (MG.ADP.AlF4)
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)
3EKJ 2008-12-16 2.8 Calcium-free GCaMP2 (calcium binding deficient mutant)
5KTG 2016-08-17 2.8 Crystal structure of mouse Bak BH3-in-groove homodimer (GFP)
3OGO 2010-08-25 2.8 Structure of the GFP:GFP-nanobody complex at 2.8 A resolution in spacegroup P21212
4Z4K 2016-04-06 2.8 Crystal structure of GFP-TAX1BP1 UBZ1+2 domain fusion protein
3EK8 2008-12-16 2.8 Calcium-saturated GCaMP2 T116V/G87R mutant monomer
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
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
6MWQ 2018-11-21 3.0 Single particle cryoEM structure of a DARPin-aldolase platform in complex with GFP
4JFG 2013-10-02 3.0 Crystal structure of sfGFP-66-HqAla
1RMP 2004-06-08 3.0 Probing the Role of Tryptophans in Aequorea Victoria Green Fluorescent Proteins with an Expanded Genetic Code
4BDU 2013-02-13 3.0 Bax BH3-in-Groove dimer (GFP)
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
5B61 2017-06-14 3.12 Extra-superfolder GFP
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
6MKS 2018-11-07 3.4 Cryo-EM structure of NLRC4-CARD filament
4XVP 2015-08-19 3.4 X-ray structure of bGFP-C / EGFP complex
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
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
95.0 Green fluorescent protein (mutated to super folder GFP, see PDB) P42212 GFP_AEQVI