Devlin TA, Fleming PJ, Loza, N, Fleming KG

Generation of Unfolded Outer Membrane Protein Ensembles Targeted by Hydrodynamic Properties

(2023) Eur. Biophys. J. Online Mar 11


Devlin TA, Marx, DC, Roskopf MA, Bubb, QR, Plummer, AM, Fleming KG

FkpA Enhances Membrane Protein Folding using an Extensive Interaction Surface

(2023) Protein Science Online February 12


Fleming PJ, Correia JJ, Fleming KG

Revisiting Macromolecular Hydration with HullRadSAS

(2023) Eur. Biophys. J. Online Jan 5.



To, P., Xia, Y. Devlin TA, Fleming KG, Fried, SD

A Proteome-Wide Map of Chaperone-Assisted Protein Refolding in the Cytosol

(2022) bioRxiv Mar 7
(In Press: Proc. of the Nat. Acad. of Sci., Nov. 23, 2022 | 119(48): e2210536119 | https://doi.org/10.1073/pnas.221053611)



Marx DC, Fleming KG

Local Bilayer Hydrophobicity Modulates Membrane Protein Stability

(2021) J. Am. Chem. Soc. Jan 20: 143(2):764-772


Vorobieva AA, White P, Liang B, Horne JE, Bera AK, Chow CM, Gerben S, Marx S, Kang A, Stiving AQ, Harvey SR, Marx DC, Khan GN, Fleming KG, Wysocki VH, Brockwell DJ, Tamm LK, Radford SE, Baker D

De novo design of transmembrane Beta-barrels

(2021) Science Feb 19: 371:eabc8182



Huysmans GHM, Marx DC, Radford SE, Fleming KG

Determining the Free Energies of Outer Membrane Proteins in Lipid Bilayers

(2020) Biophysics of Membrane Proteins. Methods in Molecular Biology Humana, New York, NY


Marx DC, Plummer AM, Faustino, AM, Devlin, T, Roskopf, MA, Leblanc, MJ, Lessen, HJ, Amann, BT, Fleming, PJ, Krueger S, Fried, SD, Fleming KG.

SurA is a cryptically grooved chaperone that expands unfolded outer membrane proteins

(2020) Proc. Natl. Acd. Sci. USA Nov 10: 117(45):28026-28035.


Marx DC, Leblanc MJ, Plummer AM, Krueger S, Fleming KG.

Domain interactions determine the conformational ensemble of the periplasmic chaperone SurA

(2020) Protein Sci. 29(10):2043-2053.


O'Brien ES, Fuglestad B, Lessen HJ, Stetz MA, Lin DW, Marques BS, Gupta K, Fleming KG, Wand AJ.

Membrane Proteins Have Distinct Fast Internal Motion and Residual Conformational Entropy

(2020) Angew. Chem. Int. Ed. Engl. 59(27):11108-11114.


Alford RF, Fleming PJ, Fleming KG, Gray JJ.

Protein Structure Prediction and Design in a Biologically Realistic Implicit Membrane

(2020) Biophys. J. 118(8):2042-2055


Lessen HJ, Majumdar A, Fleming KG.

Backbone Hydrogen Bond Energies in Membrane Proteins Are Insensitive to Large Changes in Local Water Concentration

(2020) J. Am. Chem. Soc. 142(13):6227-6235



Chum AP, Shoemaker SR, Fleming PJ, Fleming KG.

Plasticity and transient binding are key ingredients of the Periplasmic Chaperone Network

(2019) Protein Science 28(7):1340-1349.



Fleming PJ, Fleming KG.

HullRad: Fast Calculations of Folded and Disordered Protein and Nucleic Acid Hydrodynamic Properties

(2018) Biophys J. 114(4):856-869.


Lessen HJ, Fleming PJ, Fleming KG, Sodt AJ.

Building Blocks of the Outer Membrane: Calculating a General Elastic Energy Model for β-Barrel Membrane Proteins

(2018) J. Chem. Theory Comput. 14(8):4487-4497.


Fleming KG.

Taking deterministic control of membrane protein monomer-dimer measurements

(2018) J Gen Physiol 150(2)181-183.



Peterson JH, Plummer AM, Fleming KG, Bernstein HD.

Selective pressure for rapid membrane integration constrains the sequence of bacterial outer membrane proteins

(2017) Mol Microbiol 106(5)777-792.


Marx DC, Fleming KG.

Influence of Protein Scaffold on Side-Chain Transfer Free Energies

(2017) Biophys J. 113(3)597-604.


Mo GC, Ross B, Hertel F, Manna P, Yang X, Greenwald E, Booth C, Plummer AM, Tenner B, Chen Z, Wang Y, Kennedy EJ, Cole PA, Fleming KG, Palmer A, Jimenez R, Xiao J, Dedecker P, Zhang J.

Genetically encoded biosensors for visualizing live-cell biochemical activity at super-resolution

(2017) Nat Meth 14(4)427-434.


Danoff EJ, Fleming KG.

Novel Kinetic Intermediates Populated along the Folding Pathway of the Transmembrane β-Barrel OmpA

(2017) Biochemistry 56(1)47-60.



McDonald SK, Fleming KG

Negative charge neutralization in the loops and turns of outer membrane phospholipase A impacts folding hysteresis at neutral pH.

(2016) Biochemistry 55(44)6133-6137.


Costello SM, Plummer AM, Fleming PJ, Fleming KG

Dynamic periplasmic chaperone reservoir facilitates biogenesis of outer membrane proteins

(2016) PNAS 113(33)E4794-800.


Plummer AM, Fleming KG

From Chaperones to the Membrane with a BAM!

(2016) Trends Biochem Sci. 41(10):872-82.


Fleming PJ, Patel DS, Wu EL, Qi Y, Yeom MS, Sousa MC, Fleming KG, Im W

BamA POTRA Domain Interacts with a Native Lipid Membrane Surface

(2016) Biophys J 110:2698-709.


McDonald SK, Fleming KG

Aromatic Side Chain Water-to-Lipid Transfer Free Energies Show a Depth Dependence across the Membrane Normal

(2016) J Am Chem Soc. 138(25):7946<80><93>7950.


Zaccai NR, Sandlin CW, Hoopes JT, Curtis JE, Fleming PJ, Fleming KG and Krueger S

Deuterium labeling together with contrast variation small-angle neutron scattering suggests how Skp captures and releases unfolded outer membrane proteins

(2016) Methods Enzymol 566:159-210.



Sandlin, CW, Zaccai NR and Fleming KG

Skp trimer formation is insensitive to ionic strength

(2015) Biochemistry 54:7059-62.


Plummer AM, Gessmann D, and KG Fleming

The role of a destabilized membrane for OMP Insertion

(2015) Methods Mol Biol 1329:57-65.


Plummer AM and KG Fleming

BamA Alone Accelerates Outer Membrane Protein Folding In Vitro through a Catalytic Mechanism

(2015) Biochemistry 54:6009-11.


KG Fleming

A kinetic push and thermodynamic pull as driving forces for outer membrane protien sorting and folding in bacteria

(2015) Philos Trans R Soc Lond B Biol Sci 370:20150026.


Danoff EJ and KG Fleming

Aqueous unfolded OmpA forms amyloid-like fibrils upon self-association

(2015) PLos One 10:e0132301.


Danoff EJ and KG Fleming

Membrane defects accelerate outer membrane beta-barrel protein folding

(2015) Biochemistry 54:97-99.



Wu EL, Fleming PJ, Yeom MS, Widmalm G, Klauda JB, Fleming KG and W Im

E. coli outer membrane and interactions with OmpLA

(2014) Biophys J 106:2493-2502.


Gessmann D, Chung YH, Danoff EJ, Plummer AM, Sandlin CW, Zaccai NR and KG Fleming

Outer membrane beta-barrel protein folding is physically controlled by periplasmic lipid head groups and BamA

(2014) PNAS 111:5878-5883.



Moon CP, Zaccai NR, Fleming PJ, Gessmann D and KG Fleming

Membrane protein thermodynamic stability may serve as the energy sink for sorting in the periplasm

(2013) PNAS 110:4285-4290.



Lees JP, Manlandro CM, Picton LK, Tan AZ, Casares S, Flanagan JM, Fleming KG and RB Hill

A designed point mutant in Fis1 disrupts dimerization and mitochondrial fission

(2012) J Mol Biol 423:143-158.


O'Neill MJ, Bhakta MN, Fleming KG and A Wilks

Induced fit on heme binding to the Pseudomonas aeruginosa cytoplasmic protein (PhuS) drives interaction with heme oxygenase (HemO)

(2012) PNAS 109:5639-5644.


Fleming PJ, Freites JA, Moon CP, Tobias DJ and KG Fleming

Outer membrane phospholipase A in phospholipid bilayers: A model system for concerted computational and experimental investigations of amino acid side chain partitioning into lipid bilayers

(2012) Biochim Biophys Acta 1818:126-134.



Moon CP, Kwon S and KG Fleming

Overcoming hysteresis to attain reversible equilibrium folding for outer membrane phospholipase A in phospholipid bilayers

(2011) J Mol Biol 413:484-494.


Danoff EJ and KG Fleming

The soluble, periplasmic domain of OmpA folds as an independent unit and displays chaperone activity by reducing the self-association propensity of the unfolded OmpA transmembrane beta-barrel

(2011) Biophys Chem 159:194-204.


Moon CP and KG Fleming

Side-chain hydrophobicity scale derived from transmembrane protein folding into lipid bilayers

(2011) PNAS 108:10174-10177.


Moon CP and KG Fleming

Using tryptophan fluorescence to measure the stability of membrane proteins folded in liposomes

(2011) Methods Enzymol 492:189-211.



Tan AE, Burgess NK, DeAndrade DS, Marold JD and KG Fleming

Self-association of unfolded outer membrane proteins

(2010) Macromol Biosci 10:763-767.



Pang T, Savva CG, Fleming KG, Struck DK and R Young

Structure of the lethal phage pinhole

(2009) PNAS 106:18966-18971.



Tan AE and KG Fleming

Outer membrane phospholipase A dimer stability does not correlate to occluded surface area

(2008) Biochemistry 47:12095-12103.


Burgess NK, Dao TP, Stanley AM and KG Fleming

Beta-barrel proteins that reside in the Escherichia coli outer membrane in vivo demonstrate varied folding behavior in vitro

(2008) J Biol Chem 283:26748-26758.


MacKenzie KR and KG Fleming

Association energetics of membrane spanning alpha-helices

(2008) Curr Opin Struct Biol. 18:412-419.


Stanley AM and KG Fleming

The process of folding proteins into membranes: Challenges and progress

(2008) Arch Biochem Biophys 469:46-66.


Burgess NK, Stanley AM and KG Fleming

Determination of membrane protein molecular weights and association equilibrium constants using sedimentation equilibrium and sedimentation velocity

(2008) Chapter 7 in Methods in Cell Biology (JJ Correia, ed) 84:181-211.



Duong MT, Jaszewski TM, Fleming KG, and KR MacKenzie

Changes in apparent free energy of helix-helix dimerization in a biological membrane due to point mutations

(2007) J Mol Biol 371:422-434.


Stanley AM and KG Fleming

The role of a hydrogen bonding network in the transmembrane beta-barrel OMPLA

(2007) J Mol Biol 370:912-924.


Stanley AM, Treubrodt AM, Chuawong P, Hendrickson TL and KG Fleming

Lipid chain selectivity by outer membrane phospholipase A

(2007) J Mol Biol 366:461-468.


Ebie AZ and KG Fleming

Dimerization of the erythropoietin receptor transmembrane domain in micelles

(2007) J Mol Biol 366:517-524.



Stanley AM, Chuawong P, Hendrickson TL and KG Fleming

Energetics of outer membrane phospholipase A (OMPLA) dimerization

(2006) J Mol Biol 358:120-131.


Kroch AE and KG Fleming

Alternate interfaces may mediate homomeric and heteromeric assembly in the transmembrane domains of SNARE proteins

(2006) J Mol Biol 357:184-194.



Fleming KG

Analysis of membrane proteins using analytical ultracentrifugation

(2005) Analytical Ultracentrifugation, Techniques and Methods (Scott DJ, Harding SE & Rowe AJ, eds) Royal Society of Chemistry Publishing, Cambridge, UK.

Stanley AM and KG Fleming

The transmembrane domains of ErbB receptors do not dimerize strongly in micelles

(2005) J Mol Biol 347:759-772.


Kobus FJ and KG Fleming

The GxxxG-containing transmembrane domain of the CCK4 oncogene does not encode preferential self-interactions

(2005) Biochemistry 44:1464-1470.



Doura AK and KG Fleming

Complex interactions at the helix-helix interface stabilize the glycophorin A transmembrane dimer

(2004) J Mol Biol 343:1487-1497.


Raasi S, Orlov I, Fleming KG and CM Pickart

Binding of polyubiquitin chains to ubiquitin-associated (UBA) domains of HHR23A

(2004) J Mol Biol 341:1367-1379.


Doura AK, Kobus FJ, Dubrovsky L, Hibbard E and KG Fleming

Sequence context modulates the stability of a GxxxG-mediated transmembrane helix-helix dimer

(2004) J Mol Biol 341:991-998.


Fleming KG*, Ren CC, Doura AK, Eisley ME, Kobus FJ and AM Stanley
*Corresponding author

Thermodynamics of glycophorin A transmembrane helix dimerization in C14 betaine micelles

(2004) Biophys Chem 108:43-49.



Fleming KG

Standardizing the free energy change of transmembrane helix-helix interactions

(2002) J Mol Biol 323:563-571.



Fleming KG* and DM Engelman

*Corresponding author

Specificity in transmembrane helix-helix interactions can define a hierarchy of stability for sequence variants

(2001) PNAS 98:14340-14344.


Fleming KG* and DM Engelman
*Corresponding author

Computation and mutagenesis suggest a right-handed structure for the synaptobrevin transmembrane dimer

(2001) Proteins 45:313-317.


Trombetta ES, Fleming KG and A Helenius

Quaternary and domain structure of glycoprotein processing glucosidase II

(2001) Biochemistry 40:10717-10722.


Vergis JM, Bulock KG, Fleming KG and GP Beardsley

Human AICAR transformylase/IMP cyclohydrolase: A bifunctional protein requiring dimerization for transformylase activity but not for cyclohydrolase activity

(2001) J Biol Chem 276:7727-7733.



Fleming KG

Riding the wave: structural and energetic principles of helical membrane proteins

(2000) Current Opinion in Biotechnology (P Hensley & D Myszka, eds) 11:67-71.

Abstract PDF Reprint

Fleming KG

Probing stability of helical transmembrane proteins

(2000) Energetics of Biological Macromolecules, Part C Methods Enzymol (ML Johnson & G Ackers, eds) Academic Press 323:63-77.

Abstract PDF Reprint


Biswas I, Ban C, Fleming KG, Qin J, Lary JW, Yphantis DA, Yang W and P Hsieh

Oligomerization of a MutS mismatch repair protein from Thermus aquaticus

(1999) J Biol Chem 274:23673-23678.

Abstract PDF Reprint

Schubert C, Hirsch JA, Gurevich VV, Engelman DM, Sigler PB and KG Fleming*
*Corresponding author

Visual arrestin activity may be regulated by self-association

(1999) J Biol Chem 274:21186-21190.

Abstract PDF Reprint


Fleming KG

Measuring transmembrane alpha-helix energies using analytical ultracentrifugation

(1998) ChemTracts: Biological Applications of the Analytical Ultracentrifuge (JC Hanson, ed) 11:985-990.

Fleming KG, Hohl TM, Yu RC, Müller SA, Wolpensinger BA, Engel A, Engelhardt H, Brünger AT, Söllner TH, and PI Hanson

A revised model for the oligomeric state of the N-ethylmaleimide-sensitive fusion protein, NSF

(1998) J Biol Chem 273:15675-15681.

Abstract PDF Reprint

Fleming KG

Analysis of Membrane Protein Dimerization by Sedimentation Equilibrium

(1998) Application Information Bulletin (A Furst, ed) Beckman Coulter, Inc.

Engelman DM, Fleming KG and KR MacKenzie

Helix Stability and Interactions in Membrane Proteins

(1998) Biol Skr Dan Vid Selsk 49:83-86.

O'Brien R, DeDecker B, Fleming KG, Sigler PB and JE Ladbury

The effects of salt on the TATA binding protein-DNA interaction of a hyperthermophilic archaeon

(1998) J Mol Biol 279:117-125.

Abstract PDF Reprint


Rodgers KK and KG Fleming

Metal-dependent structure and self-association of the RAG1 zinc-binding domain

(1997) Techniques in Protein Chemistry VIII (DK Marshak, ed) 573-584.

Fleming KG, Ackerman AL and DM Engelman

The effect of point mutations on the free energy of transmembrane alpha-helix dimerization

(1997) J Mol Biol 272:266-275.

Abstract PDF Reprint


Munson M, Balasubramanian S, Fleming KG, Nagi AD, O'Brien R, Sturtevant JM and L Regan

What makes a protein a protein? Hydrophobic core designs that specify stability and structural properties

(1996) Protein Sci 5:1584-1593.

Abstract PDF Reprint

Rodgers KK, Bu Z, Fleming KG, Schatz DG, Engelman DM and JE Coleman

A zinc-binding domain involved in the dimerization of RAG1

(1996) J Mol Biol 260:70-84.

Abstract PDF Reprint