Heberle FA. 2019. With lipid rafts, context is everything. Biophysical Journal (in press).

Scott HL, Skinkle A, Kelley EG, Waxham MN, Levental I, Heberle FA. 2019. On the mechanism of bilayer separation by extrusion: or, why your LUVs are not really unilamellar. Biophysical Journal 117:1381-1386.

FAH’s previous publications


Nguyen MHL, DiPasquale M, Rickeard BW, Doktorova M, Heberle FA, Scott HL, Barrera FN, Taylor G, Collier CP, Stanley CB, Katsaras J, Marquardt D. 2019. Peptide-Induced Lipid Flip-Flop in Asymmetric Liposomes Measured by Small Angle Neutron Scattering. Langmuir 35:11735-11744.

Scott HL, Heberle FA, Katsaras J, Barrera FN. 2019. Phosphatidylserine asymmetry promotes the membrane insertion of a transmembrane helix. Biophysical Journal 116:1495-1506.

Doktorova M, Heberle FA, Marquardt D, Rusinova R, Sanford RL, Peyear TA, Katsaras J, Feigenson GW, Weinstein H, Anderson OS. 2019. Gramicidin Increases Lipid Flip-Flop in Symmetric and Asymmetric Lipid Vesicles. Biophysical Journal 116:860-873.


Doktorova M, Heberle FA, Eicher B, Standaert RF, Katsaras J, London E, Pabst G, Marquardt D. 2018. Preparation of asymmetric phospholipid vesicles: The next generation of cell membrane models. Nature Protocols 13:2086-2101.

Wassall SR, Leng X, Canner SW, Pennington ER, Kinnun JJ, Cavazos AT, Dadoo S, Johnson D, DeSantis A, Zeczycki T, Heberle FA, Katsaras J, Shaikh SR. 2018. Docosahexaenoic acid regulates the formation of lipid rafts: A unified view from experiment and simulation. Biochimica et Biophysica Acta 1860:1985-1993.

Enoki TA, Heberle FA, Feigenson GW. 2018. FRET Detects the Size of Nanodomains for Coexisting Liquid-Disordered + Liquid-Ordered Phases. Biophysical Journal 114:1921-1935.

Eicher B, Marquardt D, Heberle FA, Letofsky-Papst I, Rechberger GN, Appavou M-S, Katsaras J, Pabst G. 2018. Intrinsic Curvature-Mediated Transbilayer Coupling in Asymmetric Lipid Vesicles. Biophysical Journal 114:146-157.


Doktorova M, Heberle FA, Kingston RL, Khelashvili G, Wen Y, Katsaras J, Feigenson GW, Vogt VM, Dick RD. 2017. Cholesterol Promotes Protein Binding by Affecting Membrane Electrostatics and Solvation Properties. Biophysical Journal 113:2004-2015.

Taylor GJ, Heberle FA, Seinfeld JS, Katsaras J, Collier CP, Sarles SA. 2017. Capacitive Detection of Low-Enthalpy, Higher Order Phase Transitions in Synthetic and Natural Composition Lipid Membranes. Langmuir 33:10016-10026.

Usery RD, Enoki TA, Wickramasinghe SP, Weiner MD, Tsai W-C, Kim MB, Wang S, Torng TL, Ackerman DG, Heberle FA, Katsaras J, Feigenson GW. 2017. Line tension controls liquid-disordered + liquid-ordered domain size transition in lipid bilayers. Biophysical Journal 112:1431-1443 (Biophys. J. New and Notable).

Marquardt D, Heberle FA, Miti T, Eicher B, London E, Katsaras J, Pabst G. 2017. 1H NMR Shows Slow Phospholipid Flip-Flop in Gel and Fluid Bilayers. Langmuir 33:3731-3741.

Eicher B, Heberle FA, Marquardt D, Rechberger GN, Katsaras J, Pabst G. 2017. Joint small-angle X-ray and neutron scattering data analysis of asymmetric lipid vesicles. Journal of Applied Crystallography 50:419-429.

Heberle FA, Pabst G. 2017. Complex Biomembrane Mimetics on the Sub-Nanometer Scale. Biophysical Reviews 9:353-373.


Marquardt D, Heberle FA, Greathouse DV, Koeppe RE III, Standaert RF, Van Oosten BJ, Harroun TA, Kučerka N, Kinnun JJ, Williams JA, Wassall SR, Katsaras J. 2016. Lipid bilayer thickness determines cholesterol’s location in model membranes. Soft Matter 12:9393-9594.

Xia Y, Charubin K, Marquardt D, Heberle FA, Katsaras J, Tian J, Cheng X, Liu Y, Nieh M-P. 2016. Morphology-Induced Defects Enhance Lipid Transfer Rates. Langmuir 32:9757-9764.

Heberle FA, Marquardt D, Doktorova M, Geier B, Standaert RF, Heftberger P, Kollmitzer B, Nickels JD, Feigenson GW, Katsaras J, London E, Pabst G. 2016. Subnanometer Structure of an Asymmetric Model Membrane: Interleaflet Coupling Influences Domain Properties. Langmuir 32:5195-5200.


Xia Y, Li M, Charubin K, Liu Y, Heberle FA, Katsaras J, Jing B, Zhu Y, Nieh M-P. 2015. The Effects of Nanoparticle Morphology and Chain Length on Spontaneous Lipid Transfer Rates. Langmuir 31:12920-12928.

Nickels JD, Cheng X, Mostofian B, Stanley C, Lindner B, Heberle FA, Perticaroli S, Feygenson M, Egami T, Standaert RF, Smith JC, Myles DAA, Ohl M, Katsaras J. 2015. Mechanical properties of nanoscopic lipid domains. Journal of the American Chemical Society 137:15772-15780.

Heberle FA, Anghel VNP, Katsaras J. 2015. Scattering from phase-separated vesicles I. An analytical form factor for multiple static domains. Journal of Applied Crystallography 48:1391-1404.

Kučerka N, Van Oosten B, Pan J, Heberle FA, Harroun TA, Katsaras J. 2015. Molecular Structures of Fluid Phosphatidylethanolamine Bilayers Obtained from Simulation-to-Experiment Comparisons and Experimental Scattering Density Profiles. The Journal of Physical Chemistry B 119:1947-1956.

Kučerka N, Heberle FA, Pan J, Katsaras J. 2015. Structural Significance of Lipid Diversity as Studied by Small Angle Neutron and X-ray Scattering. Membranes 5:454-472.

Marquardt D, Heberle FA, Nickels JD, Pabst G, Katsaras J. 2015. On scattered waves and lipid domains: Detecting rafts with X-rays and neutrons. Soft Matter 11:9055-9072.

Heberle FA, Myles DAA, Katsaras J. 2015. Biomembranes Research Using Thermal and Cold Neutrons. Chemistry and Physics of Lipids 192:41-50.


Pan J, Marquardt D, Heberle FA, Kučerka N, Katsaras J. 2014. Revisiting the Bilayer Structures of Fluid Phase Phosphatidylglycerol Lipids: Accounting for Exchangeable Hydrogens. Biochimica et Biophysica Acta 1838:2966-2969.

Pan J, Cheng X, Monticelli L, Heberle FA, Kučerka N, Tieleman P, Katsaras J. 2014. The Molecular Structure of a Phosphatidylserine Bilayer Determined by Scattering and Molecular Dynamics Simulations. Soft Matter 10:3716-3725 (Soft Matter Hot Papers).

Heftberger P, Kollmitzer B, Heberle FA, Pan J, Rappolt M, Amenitsch H, Kučerka N, Katsaras J, Pabst G. 2014. Global SAXS data analysis for multilamellar vesicles: The evolution of the scattering density profile (SDP) model. Journal of Applied Crystallography 47:173-180.

Heberle FA, Petruzielo RS, Goh SL, Konyakhina TM, Ackerman DG, Amazon JJ, Feigenson GW. 2014. Liposome-Based Models for Membrane Rafts: Methodology and Applications. In Liposomes, Lipid Bilayers and Model Membranes: From Basic Research to Application. G. Pabst, editor. CRC Press. Boca Raton, FL.

Pan J, Kučerka N, Nieh M-P, Heberle FA, Drazba P, Katsaras J. 2014. Lipid Diversity and Its Implications For Membrane Organization. In Liposomes, Lipid Bilayers and Model Membranes: From Basic Research to Application. G. Pabst, editor. CRC Press. Boca Raton, FL.


Heberle FA, Doktorova M, Goh SL, Standaert RF, Katsaras J, Feigenson GW. 2013. Hybrid and Nonhybrid Lipids Exert Common Effects on Membrane Raft Size and Morphology. Journal of the American Chemical Society 135:14932-14935.

Konyakhina TM, Wu J, Mastroianni JD, Heberle FA, Feigenson GW. 2013. Phase Diagram of a 4-Component Lipid Mixture: DSPC/DOPC/POPC/chol. Biochimica et Biophysica Acta 1828:2204-2214.

Ackerman DG, Heberle FA, Feigenson GW. 2013. Limited Perturbation of a DPPC Bilayer by Fluorescent Lipid Probes: A Molecular Dynamics Study. The Journal of Physical Chemistry B 117:4844-4852.

Heberle FA, Petruzielo RS, Pan J, Drazba P, Kučerka N, Standaert RF, Feigenson GW, Katsaras J. 2013. Bilayer Thickness Mismatch Controls Domain Size in Model Membranes. Journal of the American Chemical Society 135:6853-6859 (JACS Spotlight).

Petruzielo RS, Heberle FA, Drazba P, Katsaras J, Feigenson GW. 2013. Phase Behavior and Domain Size in Sphingomyelin-Containing Lipid Bilayers. Biochimica et Biophysica Acta 1828:1302-1313.

Pan J, Heberle FA, Petruzielo RS, Katsaras J. 2013. The Use of Small-Angle Neutron Scattering to Detect Nanoscopic Lipid Domains. Chemistry and Physics of Lipids 170-171:19-32.

Pabst G, Heberle FA, Katsaras J. Encyclopedia of Biophysics, 1st ed., s.v. “X-ray Scattering of Lipid Membranes.” New York: Springer, 2013, DOI:10.1007/978-3-642-16712-6_554.


Pan J, Cheng X, Heberle FA, Mostofian B, Kučerka N, Drazba P, Katsaras J. 2012. Interactions between Ether Phospholipids and Cholesterol as Determined by Scattering and Molecular Dynamics Simulations. The Journal of Physical Chemistry B 116:14829-14838.

Pan J, Heberle FA, Carmichael JR, Ankner JF, Katsaras J. 2012. Time-of-Flight Bragg Scattering from Aligned Stacks of Lipid Bilayers using the Liquids Reflectometer at the Spallation Neutron Source. Journal of Applied Crystallography 45:1219-1227.

Pan J, Heberle FA, Tristram-Nagle S, Szymanski M, Koepfinger M, Katsaras J, Kučerka N. 2012. Molecular structures of fluid phase phosphatidylglycerol bilayers as determined by small-angle neutron and X-ray scattering. Biochimica et Biophysica Acta 1818:2135-2148.

Heberle FA, Pan J, Standaert RF, Drazba P, Kučerka N, Katsaras J. 2012. Model-based Approaches for the Determination of Lipid Bilayer Structure from Small-Angle neutron and X-ray Scattering Data. European Biophysical Journal 41:875-890.


Konyakhina TM, Goh SL, Amazon JA, Heberle FA, Wu J, Feigenson GW. 2011. Control of a Nanoscopic-to-Macroscopic Transition: Modulated Phases in 4-Component DSPC/DOPC/POPC/chol Giant Unilamellar Vesicles. Biophysical Journal 101:L08-L10.

Heberle FA, Feigenson GW. 2011. Phase Separation in Lipid Membranes. In The Biology of Lipids: Trafficking, Regulation, and Function. K. Simons, editor. Cold Spring Harbor Laboratory Press. New York. 49-61.

Heberle FA, Wu J, Goh SL, Petruzielo RS, Feigenson GW. 2010. Comparison of three ternary bilayer mixtures: FRET and ESR reveal nanodomains. Biophysical Journal 99:3309-3318

Mills TT, Tristram-Nagle S, Heberle FA, Morales NF, Zhao J, Wu J, Toombes GES, Nagle JF, Feigenson GW. 2008. Liquid-Liquid Domains in Bilayers Detected by Wide Angle X-Ray Scattering. Biophysical Journal 95:682-690.

Zhao J, Wu J, Heberle FA, Mills TT, Klawitter P, Huang G, Costanza G, Feigenson GW. 2007. Phase studies of model biomembranes: Complex behavior of DSPC/DOPC/Cholesterol. Biochimica et Biophysica Acta 1768:2764-2776.

Hammond AT, Heberle FA, Baumgart T, Holowka D, Baird B, Feigenson GW. 2005. Crosslinking a lipid raft component triggers liquid ordered-liquid disordered phase separation in model plasma membranes. Proceedings of the National Academy of Sciences of the USA 102(18):6320-6325.

Heberle FA, Buboltz JT, Stringer D, Feigenson GW. 2005. Fluorescence methods to detect phase boundaries in lipid bilayer mixtures. Biochimica et Biophysica Acta 1746:186-192.