Keller Group Publications

Keller Group Publications, Press, and Open Source Code
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Publications and Press in Reverse Chronological Order

Jan 2020 - present

K.J. Wilson, H.Q. Nguyen, J. Gervay-Hague, and S.L. Keller, Sterol-lipids enable large-scale, liquid-liquid phase separation in membranes of only 2 components, 2024, bioRxiv.

H.M.J. Weakly, and S.L. Keller, Coupling liquid phases in 3D condensates and 2D membranes: Successes, challenges, and tools, Biophys. J., 2024, 123, 1-13.

J. Reinhard*, C.L. Leveille*, C.E. Cornell, A.J. Merz, C. Klose, R. Ernst**, and S.L. Keller** Remodeling of yeast vacuole membrane lipidomes from the log (one phase) to stationary stage (two phases), Biophys. J., 2023, 122, 1043-1057. (Open access) *co-first authors, **co-corresponding authors (Cover article)

Z.R. Cohen, Z.R. Todd, N. Wogan, R.A. Black, S.L. Keller, and D.C. Catling, Plausible sources of membrane-forming fatty acids on the early Earth: A review of the literature and an estimation of amounts., ACS Earth Space Chem, 2023, 17, 11-27.

Z.R. Todd, Z.R. Cohen, D.C. Catling, S.L. Keller, and R.A. Black, Growth of prebiotically plausible fatty acid vesicles proceeds in the presence of prebiotic amino acids, dipeptides, sugars, and nucleic acid components, Langmuir, 2022, 38, 15106-15112.

Z.R. Cohen, Z.R. Todd, D.C. Catling, R.A. Black, and S.L. Keller, Prebiotic vesicles retain solutes and grow by micelle addition after brief cooling below the membrane melting temperature, Langmuir, 2022, 38, 13407-13413.

T. Portet, Z.R. Cohen, G.J. Goetz, N. Panek, P.N. Holmes, S.A. Stephens, T. Varga, and S.L. Keller, Ripples at edges of blooming lilies and torn plastic sheets, Biophys. J., 2022, 121, 2389-2397. (Cover article)

C.L. Leveille, C.E. Cornell, A.J. Merz, and S.L. Keller, Yeast actively tune their membranes to phase separate at temperatures that scale with growth temperatures, PNAS, 2022, 119, e2116007119.

Z.R. Cohen, C.E Cornell, D.C. Catling, R.A. Black, and S.L. Keller, Prebiotic protocell membranes retain encapsulated contents during flocculation, and phospholipids preserve encapsulation during dehydration, Langmuir, 2022, 38 1304-1310.

Z.R. Cohen, B.L. Kessenich, A.Hazra, J. Nguyen, R.S. Johnson, M.J. MacCoss, G. Lalic, R.A. Black, and S.L. Keller, Prebiotic membranes and micelles do not inhibit peptide formation during dehydration, ChemBioChem, 2021, e2021000614.

L.-P. Bergeron-Sandoval, S. Kumar, H. Khadivi Heris, C.L.A. Chang, C.E. Cornell, S.L. Keller, P. Francois, A.G. Hendricks, A.J. Ehrlicher, R.V. Pappu, and S.W. Michnick, Endocytic proteins with prion-like domains form viscoelastic condensates that enable membrane remodeling, PNAS, 2021, 118 e2113789118.

M. Xue, R.A. Black, Z.R. Cohen, A. Roehrich, G.P. Drobny, and S.L. Keller, Binding of dipeptides to fatty acid membranes explains their colocalization in protocells but does not select for them relative to unjoined amino acids, J. Phys. Chem. B, 2021, 125 7933-7939.

C.E. Cornell, A. Mileant, N. Thakkar, K.K. Lee, and S.L. Keller, Direct imaging of liquid domains in membranes by cryo-electron tomography PNAS, 2020, 117 19713-19719. (Cover article)

M. Xue, R. Black, C.E. Cornell, G.P. Drobny, and S.L. Keller, A step toward molecular evolution of RNA: Ribose binds to prebiotic fatty acid membranes, and nucleobases bind better than individual fatty acids do, ChemBioChem, 2020, 21 2764-2767. (Cover article)

Press, Write-ups, and Online Presentations:

BPS Blog "What Makes Membranes of Yeast Vacuoles Phase Separate?" - the people behind the image, 2023.
BPS Blog "Birth of the Bloom" - the human story behind the science story, 2022.
UW News Hungry yeast are tiny, living thermometers, 2022.
NASA Astrobiology Workshop presentation by Zack Cohen, 2022.
Physics Today "Why does biophysics attract a disproportionate number of women?", 2021.
BPS Bulletin "Society Names 2021 Fellows", 2021.
PBS Press release "Biophysical Society Announces 2021 Society Fellows", 2021.
UW Astrobiology Panel on "What is life and how did it begin?" presentation by Zack Cohen, 2021.
UW Astrobiology Colloquium presentation by Sarah Keller, 2020
BPS Blog "A 17-Year-Old BJ article explored the ground rules of phase separation in lipid bilayer", by Andrea Gohlke, 2020.

Jan 2015 - Dec 2019

C.E. Cornell, R.A. Black, M. Xue, H.E. Litz, A. Ramsay, M. Gordon, A. Mileant, Z.R. Cohen, J.A. Williams, K.K. Lee, G.P. Drobny, and S.L. Keller, Prebiotic amino acids bind to and stabilize prebiotic fatty acid membranes, PNAS, 2019, 116, 17239-17244. Postprint, as allowed by PNAS author licenses.
C.E. Cornell, A.D. Skinkle, S. He, I. Levental, K.R. Levental, and S.L. Keller, Tuning length scales of small domains in cell-derived membranes and synthetic model membranes, Biophys. J., 2018, 115, 690-701.
S.P. Rayermann, G.E. Rayermann, C.E. Cornell, A.J. Merz, and S.L. Keller, Hallmarks of reversible separation of living, unperturbed cell membranes into two liquid phases, Biophys. J., 2017, 113, 2425-2432.
C.E. Cornell, N.L.C. McCarthy, K.R. Levental, I. Levental, N.J. Brooks, and S.L. Keller, Lengths of n-alcohols govern how Lo-Ld mixing temperatures shift in synthetic and cell-derived membranes, Biophys. J., 2017, 113, 1200-1211. (Open access)
M.C. Blosser, C.E. Cornell, S.P. Rayermann, and S.L. Keller, Phase Diagrams and Tie Lines in GUVs, Chapter 18 in "The Giant Vesicle Book" edited by R. Dimova and C. Marques, in press, 2017.
M.C. Blosser, B.G. Horst, and S.L. Keller, cDICE Method Produces Giant Lipid Vesicles under Physiological Conditions of Charged Lipids and Ionic Solutions, 2016, Soft Matter, 2016, 12, 7364-7371. (Open access)
R.A. Black and M.C.Blosser, A self-assembled aggregate composed of a fatty acid membrane and the building blocks of biological polymers provides a first step in the emergence of protocells, Life, 2016, 6, life6030033. (doi:10.3390/life6030033)
J.V. Bleecker, P.A. Cox, and S.L. Keller, Mixing Temperatures of Bilayers Not Simply Related to Thickness Differences Between Lo and Ld Phases, Biophys. J., 2016, 110, 2305-2308.
J.V. Bleecker, P.A. Cox, R.N. Foster, J.P. Litz, M.C. Blosser, D.G. Castner, and S.L. Keller, Thickness Mismatch of Coexisting Liquid Phases in Non-Canonical Lipid Bilayers, J. Phys. Chem. B, 2016, 120, 2761-2770.
J.P. Litz, N. Thakkar, T. Portet, and S.L. Keller, Depletion with Cyclodextrin Reveals Two Populations of Cholesterol in Model Membranes, Biophys. J., 2016, 110, 635-645.
M.C. Blosser, A.R. Honerkamp-Smith, T. Han, M. Haataja, and S.L. Keller, Transbilayer Colocalization of Lipid Domains Explained via Measurement of Strong Coupling Parameters, Biophys. J, 2015, 109, 2317-2327. (Cover article)
S.L. Keller and A.L. Kwiram, Harden Marsden McConnell (obituary), Physics Today, 2015, 68, 64.
Press, Write-ups, and Stories:
- The Daily "The Bond Behind Origins of Life", Nov. 4, 2019.
- The Stranger "My Brother's Passing, God, and the Origins of Life", Sept. 11, 2019.
- Hank Green - SciShow News "How Cells Got Their Membranes (Maybe)", Aug. 16, 2019.
- NASA Research Highlight "Did Amino Acids Stabilize Prebiotic Membranes?", Aug. 14, 2019, (with a tweet by Thomas Zurbuchen, Assoc. Admin. NASA Science Mission Directorate)
- Discover "How Cell Membranes May Have Kicked Off the Origin of Life", Aug. 12, 2019.
- Sofie Bates - Inside Science "The Origin of Life May Not Be as Coincidental as Scientists Once Thought", Aug. 12, 2019.
- Ed Yong - The Atlantic "A New Clue to How Life Originated", Aug. 12, 2019.
- NASA's comic book "Astrobiology - The Story of our Search for Life in the Universe, Issue #7, 2019. (See page 23)
- Physics Today article "Membrane phase demixing seen in living cells", Feb. 2018.
- Physics Today update "Demixing in cell membranes", Dec. 2017.
- UW News "Living cell membranes can self-sort their components by demixing", Dec. 2017.
- Alex's Twitter Tutorial on the history and implications of domains in yeast vacuoles.
- NSF: News from the field, regarding our observation of domains in yeast vacuoles, Dec. 2017.
- Biophysical Society Press Release of 2017 Avanti Award.
- Biophysical Journal Spotlight, "Two Phases on Two Faces", Dec. 2015.

Jan 2010 - Dec 2014

R.A. Black, M.C. Blosser, B.L. Stottrup, R. Tavakley, D.W. Deamer, and S.L. Keller, Nucleobases Bind to and Stabilize Aggregates of a Prebiotic Amphiphile, Providing a Viable Mechanism for the Emergence of Protocells, Proc. Natl. Acad. Sci, 2013, 110, 13272.
C.A. Stanich, A.R. Honerkamp-Smith, G.G. Putzel, C.S. Warth, A.K. Lamprecht, P. Mandal, E. Mann, T.D. Hua, and S.L. Keller, Coarsening Dynamics of Domains in Lipid Membranes, Biophys. J., 2013, 105, 444. (cover article)
M.C. Blosser, J.B. Starr, C.W. Turtle, J. Ashcraft, and S.L. Keller, Minimal Effect of Lipid Charge on Membrane Miscibility Phase Behavior in Three Ternary Systems, Biophys. J., 2013, 104, 2629.
T. Portet, S.E. Gordon, and S.L. Keller, Increasing Membrane Tension Decreases Miscibility Temperatures; an Experimental Demonstration via Micropipette Aspiration, Biophys. J., 2012, 103, L35-L37. (featured article; free download)
A.R. Honerkamp-Smith and S.L. Keller Experimental Observations of Dynamic Critical Phenomena in a Lipid Membrane, Phys. Rev. Lett, 2012, 108, 265702.
M.M. Stevens, A.R. Honerkamp-Smith and S.L. Keller, Solubility limits of cholesterol, lanosterol, ergosterol, stigmasterol, and beta-sitosterol in electroformed lipid vesicles, Soft Matter, 2010, 6, 5882-5890. (cover article)
Press and Write-ups:
- Biophysical Journal Spotlight, "Giant Unilamellar Vesicle Dominates BJ Cover", May 2014.
- F1000Prime Apr. 2014.
- The Economist: Babbage, the Science and Technology Blog
- Aggregators: Slashdot, Science Daily, Kurzweil Accelerating Intelligence, R&D Magazine, Science World Report, Teknoids, Yahoo India, Zee News India, myScience, and others, July 29-31 2013.
- The Home Team: UW Today July 29 2013, UW Daily Aug. 6 2013, UW Arts & Sciences Perspectives Dec. 2013.
- Biophysical Society, July 25 2013.
- ASBMB today, June 2010, p. 32.
- ASBMB today, Jan. 2010, p. 19.

Jan 2005 - Dec 2009

A.R. Honerkamp-Smith, S.L. Veatch, and S.L. Keller, An Introduction to Critical Points for Biophysicists: Observations of Compositional Heterogeneity in Lipid Membranes, Biochim. Biophys. Acta - Biomembranes (Invited), 1788, 53-63, 2009. (cover article)
A.R. Honerkamp-Smith, P. Cicuta, M.D. Collins, S.L. Veatch, M. den Nijs, M. Schick, and S.L. Keller, Line Tensions, Correlation Lengths, and Critical Exponents in Lipid Membranes near Critical Points, Biophys. J., 95, 236-246, 2008.
M.D. Collins and S.L. Keller, Tuning Lipid Mixtures to Induce Domains across Leaflets of Unsupported Asymmetric Bilayers, Proc. Natl. Acad. Sci., 105, 124-128, 2008.
M. Halter, Y. Liao, R.M. Plocinik, D.C. Coffey, S. Bhattacharjee, U. Mazur, G.J. Simpson, B.H. Robinson, and S.L. Keller Molecular Self-Assembly of Mixed High-beta Zwitterionic and Neutral Ground State NLO Chromophores, Chem. Mater., 20, 1778-1787, 2008.
S.L. Veatch, O. Soubias, S.L. Keller, and K. Gawrisch, Critical Fluctuations in Domain-Forming Lipid Mixtures, Proc. Natl. Acad. Sci., 104, 17650-17655, 2007.
P. Cicuta, S.L. Keller, and S.L. Veatch, Diffusion of Liquid Domains in Lipid Bilayer Membranes, J. Phys. Chem. B., 111, 3328-3331, 2007.
S.L. Veatch, K. Gawrisch, and S.L. Keller, Closed-loop Miscibility Gap and Quantitative Tie-lines in Ternary Membranes Containing DiphytanoylPC", Biophys. J., 90, 4428-4436 2006.
B.L. Stottrup and S.L. Keller, Phase Behavior of Lipid Monolayers Containing DPPC and Cholesterol Analogs, Biophys. J., 90, 3176-3183, 2006.
S.L. Keller and A.L. Smith, Advice for New Faculty Teaching Undergraduate Science J. Chem. Educ., 83, 401-406, 2006.
S.L. Veatch and S.L. Keller, Seeing Spots: Complex Phase Behavior in Simple Membranes, Biochim. Biophys. Acta - Molecular Cell Research (Invited), 1746, 172-185, 2005. (cover article)
M.E. Beattie, S.L. Veatch, B.L. Stottrup, and S.L. Keller, Sterol Structure Determines Miscibility vs. Melting Transitions in Lipid Vesicles, Biophys. J., 89, 1760-1768, 2005.
S.L. Veatch and S.L. Keller, Miscibility Phase Diagrams of Giant Vesicles Containing Sphingomyelin, Phys. Rev. Lett., 94, 148101, 2005.
B.L. Stottrup, D.S. Stevens, and S.L. Keller, Miscibility of Ternary Mixtures of Phospholipids and Cholesterol in Monolayers, and Application to Bilayer Systems, Biophys. J., 88, 269-276, 2005.
Press and Write-ups:
- "Simulating Life's Envelopes" Chemical and Engineering News, Feb. 9 2009, p. 31-33.

Jan 2000 - Dec 2004

S.L. Veatch, I.V. Polozov, K. Gawrisch, and S.L. Keller Liquid Domains in Vesicles Investigated by NMR and Fluorescence Microscopy Biophys. J. 2004 86 2910-2922.
B.L. Stottrup, S.L. Veatch and S.L. Keller Nonequilibrium Behavior in Supported Lipid Membranes Containing Cholesterol Biophys. J. 2004 86 2942-2950.
S.L. Keller Sequential Folding of a Rigid Wire into Three-Dimensional Structures Am. J. Phys. 2004 72 599-604.
S.L. Veatch and S.L. Keller Separation of Liquid Phases in Giant Vesicles of Ternary Mixtures of Phospholipids and Cholesterol Biophys. J. 2003 85 3074-3083.
S.L. Veatch and S.L. Keller A Closer Look at the Canonical Raft Mixture in Model Membrane Studies Biophys. J. 2003 84, 725-726.
S.L. Keller Miscibility Transitions and Lateral Compressibility in Liquid Phases of Lipid Monolayers Langmuir 2003 19, 1451-1456.
S.L. Veatch and S.L. Keller Organization in Lipid Membranes Containing Cholesterol Phys. Rev. Lett 2002 89 268101.
S. Bezzine, J.G. Bollinger, S.L. Veatch, S.L. Keller and M.H. Gelb, On the Binding Preference of Secreted Phospholipases A2 for Membranes with Anionic Phospholipids J. Biol. Chem. 2002 277, 48523-48534.
S.L. Keller Coexisting Liquid Phases in Lipid Monolayers and Bilayers J. Phys.: Condens. Matt. (Special Issue Article), 2002, 14, 4763-4766.
W.H. Pitcher III, S.L. Keller, and W.H. Huestis Interaction of Nominally Soluble Proteins with Phospholipid Monolayers at the Air-Water Interface Biochim. Biophys. Acta (Biomembranes) 2002 1564, 107-113.
S.L. Keller, T.G. Anderson, and H.M. McConnell, Miscibility Critical Pressures in Monolayers of Ternary Lipid Mixtures Biophys. J., 2000, 79, 2033-2042.
S.L. Keller, A. Radhakrishnan, and H.M. McConnell, Saturated Phospholipids with High Melting Temperature Form Complexes with Cholesterol in Monolayers J. Phys. Chem. B, 2000, 104, 7522-7527.
Press and Write-ups:
- "Order Out of Disorder" 2000 Yearbook of Science and the Future, Encyclopedia Brittanica, p. 247.

Jan 1995 - Dec 1999

S.L. Keller and H.M. McConnell, Stripe Phases in Lipid Monolayers near a Miscibility Critical Point, Phys. Rev. Lett., 1999, 82, 1602.
S.L. Keller, W.H. Pitcher III, W.H. Huestis and H.M. McConnell, Red Blood Cell Lipids Form Immiscible Liquids, Phys. Rev. Lett., 1998, 81, 5019.
S.L. Keller, P. Boltenhagen, D. Pine and J.A. Zasadzinski, Direct Observation of Shear-Induced Structures in Worm-Like Micelles: a Freeze-Fracture Electron Microscopy Study, Phys. Rev. Lett., 1998, 80, 2725-2728.
M. Adams, Z. Dogic, S.L. Keller and S. Fraden, Entropically Driven Microphase Transitions in Mixtures of Colloidal Rods and Spheres, Nature, 1998, 393, 349.
H.E. Warriner, S.L. Keller, S.H.J. Idziak, N.L. Slack, P. Davidson, J.A. Zasadzinski and C.R. Safinya, The Influence of Polymer Molecular Weight in Lamellar Gels Based on PEG-Lipids, Biophys. J., 1998, 75, 272-293.
S.L. Keller, H.E. Warriner, C.R. Safinya and J.A. Zasadzinski, Direct Observation of the Defect Structure in a Lamellar Biogel of Membranes and Polymer-Lipids. Phys. Rev. Letters, 1997, 78, 4781-4785.
S.L. Keller, S.M. Gruner and K. Gawrisch, Small Concentrations of Alamethicin Induce a Cubic Phase in Bulk Phosphatidylethanolamine Mixtures, Biochim. Biophys. Acta, 1996, 1278, 241-246.
Y-S. Lee, J-Z. Yang, T.M. Sisson, D.A. Frankel, J.T. Gleeson, E. Aksay, S.L. Keller, S.M. Gruner and D.F. O'Brien, Polymerization of Nonlamellar Lipid Assemblies, J. Am. Chem. Soc., 1995, 117, no. 20, 5573-5578.
Press and Write-ups:
- Peter Olmsted, "Worm-Like Micelles Stir Up a Storm" Physics World, June 1998.
- "Entropy Drives Phase Behavior in Colloids" Chemical and Engineering News, June 1 1998, p. 43.

Jan 1990 - Dec 1994

S.L. Keller, S.M. Bezrukov, S.M. Gruner, M.W. Tate, I. Vodyanoy and V.A. Parsegian, Probability of Alamethicin Conductance States Varies with Nonlamellar Tendency of Bilayer Phospholipids, Biophys. J., 1993, 65, 23-27.

Original Open Source MATLAB and Python Code

Track_Vesicle (.docx) This library of original Matlab code tracks, centers, and crops images of vesicles. Copyright 2012, Cynthia A. Stanich, Aurelia R. Honerkamp-Smith, Pietro Cicuta, Matthew C. Blosser, and Chris S. Warth. This code is described in the 2013 publication: Coarsening Dynamics of Domains in Lipid Membranes

BilayerHeightMeasurements.py. This original Python code Canny filters cryo electron micrographs to detect the inner and outer edges of bilayers. It is available by public license at github and was written by Caitlin E. Cornell. The program is described in the 2020 publication: Direct imaging of liquid domains in membranes by cryo-electron tomography

Crop and Display Vacuoles. This original Python code crops fields of view into individual images of vacuoles and displays those images to the user. It is available by public license at github and was written by Chantelle L. Leveille.

Lily tepal analysis. This original Python code skeletonizes images of sliced lily tepals and measures tepal thicknesses along lines that are perpendicular to the skeleton. It is available by public license at github and was written by Nicole Panek and Gunnar Goetz.

The MATLAB and python code above is based upon work supported by the National Science Foundation under grant numbers MCB-0744852, MCB-1402059, and MCB-1925731. Any opinions, findings, and conclusions or recommendations expressed in this material are those of the author(s) and do not necessarily reflect the views of the National Science Foundation.

Sarah L. Keller, Ph.D.
Professor of Chemistry
Dept. of Chemistry 351700, Univ. of Washington
Seattle, WA, 98195-1700
e-mail: slkeller -at- chem.washington.edu