Faculty Appointments
Assistant Professor of Biochemistry
Education
Ph.D., Cell Biology, Molecular Biology, Vanderbilt University, Nashville, TennesseeB.S., Biochemistry, University of Iowa, Iowa City, Iowa
Research Description
How does a cell organize biochemistry within a crowded, dynamic molecular environment? Our laboratory approaches this question through the lens of soft-matter biophysics. The cytoplasm and nucleoplasm are not uniform solutions, but heterogeneous materials in which proteins and nucleic acids interact collectively to generate distinct molecular environments. Biomolecular condensates are a form of biological soft matter, in which many weak molecular interactions collectively give rise to emergent properties such as phase behavior, interfaces, material states, and dynamic molecular transport. These concentrated assemblies of proteins and nucleic acids can organize cellular components without a surrounding lipid membrane. Unlike membrane-bound organelles such as the nucleus or mitochondria, condensates establish compartments through networks of weak, multivalent molecular interactions. These interactions can drive phase separation and other forms of mesoscale organization, producing dynamic structures with distinct compositions, interfaces, material properties, and molecular dynamics. Because molecules can continuously exchange across condensate boundaries, these assemblies provide an opportunity to understand how cells organize biochemical reactions without relying on physical barriers.
We are particularly interested in the relationship between the physical properties of condensates and their biological functions. How do interaction networks determine which molecules enter, leave, or become enriched at an interface? How do material properties such as viscosity, elasticity, molecular mobility, and interfacial organization influence biochemical activity? And how do cells reshape these properties through environmental changes, post-translational modifications, and cell-cycle regulation? To address these questions, we combine experiments in the nematode C. elegans, cultured cells, and minimal in vitro reconstituted systems. Reconstitution allows us to systematically perturb molecular interactions and measure the resulting phase behavior and material properties, while experiments in living cells allow us to determine how these physical principles are exploited and regulated in a biological context.
Our laboratory integrates quantitative soft-matter approaches with cell and molecular biology, including in vitro reconstitution, single-molecule tracking, super-resolution microscopy, quantitative imaging, biophysical measurements, CRISPR genome editing, and computational modeling. By connecting molecular interactions to mesoscale material properties and ultimately to cellular behavior, our long-term goal is to uncover the physical principles that allow cells to organize and regulate biochemical activity across scales, from molecules, to condensates, to cells, and through animal development.
We are particularly interested in the relationship between the physical properties of condensates and their biological functions. How do interaction networks determine which molecules enter, leave, or become enriched at an interface? How do material properties such as viscosity, elasticity, molecular mobility, and interfacial organization influence biochemical activity? And how do cells reshape these properties through environmental changes, post-translational modifications, and cell-cycle regulation? To address these questions, we combine experiments in the nematode C. elegans, cultured cells, and minimal in vitro reconstituted systems. Reconstitution allows us to systematically perturb molecular interactions and measure the resulting phase behavior and material properties, while experiments in living cells allow us to determine how these physical principles are exploited and regulated in a biological context.
Our laboratory integrates quantitative soft-matter approaches with cell and molecular biology, including in vitro reconstitution, single-molecule tracking, super-resolution microscopy, quantitative imaging, biophysical measurements, CRISPR genome editing, and computational modeling. By connecting molecular interactions to mesoscale material properties and ultimately to cellular behavior, our long-term goal is to uncover the physical principles that allow cells to organize and regulate biochemical activity across scales, from molecules, to condensates, to cells, and through animal development.
Research Keywords
Biochemistry, Biophysics, Cell Biology, Developmental Biology, Genetics, Single-molecule microscopy, Super-resolution microscopy, Molecular biology, Molecular genetics, RNA biology, Quantitative Biology, Genome editing, Cell polarity
Publications
Gilbert AE, Bomber ML, Ellis JD, Bartlett LN, Chakraborty J, Folkmann AW, Stengel KR, Hiebert SW. Mtg16/Eto2 tumor suppressor maintains and establishes repression by distinct mechanisms. Mol Cell [print-electronic]. 2026 Jul 7/2/2026; 86(13): 2492-2508.e6-2508.e6. PMID: 42309063, PII: S1097-2765(26)00349-7, DOI: 10.1016/j.molcel.2026.05.025, ISSN: 1097-4164.
Donahue EKF, Hepowit NL, Ruark EM, Mulligan AG, Keuchel B, Urban ND, Peng L, Stephens S, Johnson DJ, Wallace NS, Jackson LP, Ellisman MH, Arrojo E Drigo R, Folkmann AW, Truttmann MC, MacGurn JA, Burkewitz K. ER remodelling is a feature of ageing and depends on ER-phagy. Nat Cell Biol [print-electronic]. 2026 Mar; 28(3): 449-64. PMID: 41629400, PMCID: PMC12992112, PII: 10.1038/s41556-025-01860-1, DOI: 10.1038/s41556-025-01860-1, ISSN: 1476-4679.
Thomas L, Putnam A, Folkmann A. Germ granules in development. Development [print-electronic]. 2023 Jan 1/15/2023; 150(2): PMID: 36715566, PMCID: PMC10165536, PII: 286764, DOI: 10.1242/dev.201037, ISSN: 1477-9129.
Folkmann AW, Putnam A, Lee CF, Seydoux G. Regulation of biomolecular condensates by interfacial protein clusters. Science [print-electronic]. 2021 Sep 9/10/2021; 373(6560): 1218-24. PMID: 34516789, PMCID: PMC8627561, DOI: 10.1126/science.abg7071, ISSN: 1095-9203.
Folkmann AW, Collier SE, Zhan X, Aditi, Ohi MD, Wente SR. Gle1 functions during mRNA export in an oligomeric complex that is altered in human disease. Cell [print-electronic]. 2013 Oct 10/24/2013; 155(3): 582-93. PMID: 24243016, PMCID: PMC3855398, PII: S0092-8674(13)01160-4, DOI: 10.1016/j.cell.2013.09.023, ISSN: 1097-4172.
Donahue EKF, Hepowit NL, Ruark EM, Mulligan AG, Keuchel B, Urban ND, Peng L, Stephens S, Johnson DJ, Wallace NS, Jackson LP, Ellisman MH, Arrojo E Drigo R, Folkmann AW, Truttmann MC, MacGurn JA, Burkewitz K. ER remodelling is a feature of ageing and depends on ER-phagy. Nat Cell Biol [print-electronic]. 2026 Mar; 28(3): 449-64. PMID: 41629400, PMCID: PMC12992112, PII: 10.1038/s41556-025-01860-1, DOI: 10.1038/s41556-025-01860-1, ISSN: 1476-4679.
Thomas L, Putnam A, Folkmann A. Germ granules in development. Development [print-electronic]. 2023 Jan 1/15/2023; 150(2): PMID: 36715566, PMCID: PMC10165536, PII: 286764, DOI: 10.1242/dev.201037, ISSN: 1477-9129.
Folkmann AW, Putnam A, Lee CF, Seydoux G. Regulation of biomolecular condensates by interfacial protein clusters. Science [print-electronic]. 2021 Sep 9/10/2021; 373(6560): 1218-24. PMID: 34516789, PMCID: PMC8627561, DOI: 10.1126/science.abg7071, ISSN: 1095-9203.
Folkmann AW, Collier SE, Zhan X, Aditi, Ohi MD, Wente SR. Gle1 functions during mRNA export in an oligomeric complex that is altered in human disease. Cell [print-electronic]. 2013 Oct 10/24/2013; 155(3): 582-93. PMID: 24243016, PMCID: PMC3855398, PII: S0092-8674(13)01160-4, DOI: 10.1016/j.cell.2013.09.023, ISSN: 1097-4172.