Milka Doktorova

Group Leader, DDLS Fellow, Stockholm University

Key Publications

Doktorova M, Symons JL, Zhang X, Wang HY, Schlegel J, Lorent JH, Heberle FA, Sezgin E, Lyman E, Levental KR, Levental I. 2025. Cell membranes sustain phospholipid imbalance via cholesterol asymmetry. Cell. 188(10):2586 – 2602.E24.

Sharma K, Heberle FA, Doktorova M. 2026. From molecular dynamics to cryo-EM: Imaging liposomes in silico. In Methods in Enzymology (MIE) Volume 727: Lipids and Membranes: Dynamics and Interorganelle Lipid Transport. Pages 291-319. Baskin J editor. Academic Press. Cambridge, MA.

Heberle FA, Doktorova M. 2025. Exploring the sensitivities of experimental techniques to various types of membrane asymmetry using atomistic simulations. Faraday Discuss. 259:300-320.

Doktorova M, Daum S, Reagle TR, Cannon HI, Ebenhan J, Neudorf S, Han B, Sharma S, Kasson P, Levental KR, Bacia K, Kenworthy AK, Levental I. 2025. Caveolin assemblies displace one bilayer leaflet to organize and bend membranes. Proc. Natl. Acad. Sci. U.S.A. 122(20), e2417024122.

Doktorova M, and Weinstein H. 2018. Accurate in silico modeling of asymmetric bilayers based on biophysical principles. Biophys. J. 115:1638-1643.

Doktorova M*, Heberle FA*, Geier B, Standaert RF, Katsaras J, London E, Pabst G, and Marquardt D*. 2018. Preparation of asymmetric phospholipid vesicles for use as cell membrane models. Nat. Protoc. 13:2086-2101.

Research interest

Lipid membranes are among the most fundamental structures in biology. They consist of two layers (or leaflets) of lipid molecules and form thin, flexible sheets that separate and organize biological environments. Despite their simple appearance, they possess a rich and complex internal structure. Different lipids can mix non-ideally and distribute asymmetrically between the two membrane leaflets, creating local variations that influence membrane behavior. Understanding this organization is a long-standing challenge because it occurs on extremely small length scales that are difficult to measure directly.

The Cell Membrane Biophysics Lab (CMBL) tackles this challenge by combining various computational, theoretical and experimental tools to bridge time and length scales and uncover molecular mechanisms driving observed phenomena. Our main approach thus involves identifying fundamental principles in well-designed simpler models and translating them to more complex biological environments.

We work with both cellular and synthetic membranes, primarily focusing on their biophysical properties. We study how collective lipid organization and dynamics—including lipid distributions in and between leaflets—affect membrane structure (lipid packing, thickness, curvature), elasticity (tension, bending, compression), and interactions with membrane proteins. Understanding these lipid-based structure-function relationships at different scales and contexts is essential for uncovering the often-hidden roles of lipid organization in biological phenomena.

Group members

  • Frederick Heberle, Senior researcher
  • Karan Sharma, Postdoctoral researcher
  • Shambhavi Pandey, Postdoctoral researcher
  • Julian Wagner, PhD student
  • Leon Bergner, PhD student

Last updated: 2026-07-20

Content Responsible: Sofia Falorni(sofia.falorni@scilifelab.se)