Bibliography
Publications
71 papers from 2004 to today. The full list also is found in Stirling's NCBI bibliography and on Google Scholar.
*equal authorship; #co-corresponding authorship; Churchman Lab members in bold.
Preprints
- 2026
Mitochondrial RNA processing promotes translation by resolving structured precursor RNAs
Nuessmeier CH, Prakash G, Hansen LN, Couvillion MT, Churchman LS
bioRxiv
- 2026
Analog intrinsic recoding measures RNA dynamics without chemical conversion
Hansen LN, Couvillion MT, McShane E, Azbukina N, Treutlein B, Churchman LS
bioRxiv
- 2026
- 2025
Research articles
- 2026
Translational activators align mRNAs at the small mitoribosomal subunit for translation initiation
Bridgers JB*, Carlström A*, Sherpa D*, Couvillion MT, Rovšnik U, Gao J, Wan B, Shao S#, Ott M#, Churchman LS#
Nature Structural & Molecular Biology 33(2), 245–258

- 2026
Copper depletion boosts CNS leukemia therapy by inhibiting nucleotide synthesis through impairment of mitochondrial complex IV activity
Wong AYL, Myers JW, Prakash G, Ma A, Brook JR, Petrova B, Bulut B, White R 3rd, Merz C, de Souza M, Maynard AG, Wang P, Yu A, Pohl NK, Weitman M, Silverman LB, Stegmaier K, Churchman LS, Cole PD, Brady DC, Kanarek N
Nature Cancer

- 2025
Transcription arrest induces formation of RNA granules in mitochondria
Hansen KG, Baxter-Koenigs A, Weiss CA, McShane E, Churchman LS
Life Science Alliance 8(9), e202403082

- 2025
A molecular switch at the yeast mitoribosomal tunnel exit controls cytochrome b synthesis
Carlström A, Bridgers JB, Couvillion M, Singh AP, Forné I, Imhof A, Churchman LS#, Ott M#
Nucleic Acids Research 53(13)

- 2025
HIF regulates multiple translated endogenous retroviruses: Implications for cancer immunotherapy
Jiang Q, Braun DA, Clauser KR, Ramesh V, Shirole NH, Duke-Cohan JE, Nabilsi N, Kramer NJ, Forman C, Lippincott IE, Klaeger S, Phulphagar KM, Chea V, Kim N, Vanasse AP, Saad E, Parsons T, Carr-Reynolds M, Carulli I, Pinjusic K, Jiang Y, Li R, Syamala S, Rachimi S, Verzani EK, Stevens JD, Lane WJ, Camp SY, Meli K, Pappalardi MB, Herbert ZT, Qiu X, Cejas P, Long HW, Shukla SA, Van Allen EM, Choueiri TK, Churchman LS, Abelin JG, Gurer C, MacBeath G, Childs RW, Carr SA, Keskin DB, Wu CJ, Kaelin WG Jr
Cell 188(7), 1807–1827.e34
- 2025
Genetic regulation of nascent RNA maturation revealed by direct RNA nanopore sequencing
Choquet K, Chaumont LP, Bache S, Baxter-Koenigs AR, Churchman LS
Genome Research 35(4), 712–724

- 2025
Context-specific inhibition of mitochondrial ribosomes by phenicol and oxazolidinone antibiotics
Bibel B, Raskar T, Couvillion M, Lee M, Kleinman JI, Takeuchi-Tomita N, Churchman LS, Fraser JS, Fujimori DG
Nucleic Acids Research 53(3)

- 2024
RNA polymerases reshape chromatin architecture and couple transcription on individual fibers
Tullius TW, Isaac RS, Ranchalis J, Dubocanin D, Churchman LS#, Stergachis AB#
Molecular Cell 84(17), 3209–3222.e5

- 2024
Genome-wide quantification of RNA flow across subcellular compartments reveals determinants of the mammalian transcript life cycle
Ietswaart R#*, Smalec BM*, Xu A*, Choquet K, McShane E, Jowhar ZM, Guegler CK, Baxter-Koenigs AR, West ER, Fu BXH, Gilbert L, Floor SN#, Churchman LS#
Molecular Cell 84, 2765–2784.e16

- 2024
A kinetic dichotomy between mitochondrial and nuclear gene expression processes
McShane E, Couvillion M, Ietswaart R, Prakash G, Smalec BM, Soto I, Baxter-Koenigs AR, Choquet K, Churchman LS
Molecular Cell 84(8), 1541–1555.e11

- 2024
Post-transcriptional splicing can occur in a slow-moving zone around the gene
Coté A, O'Farrell A, Dardani I, Dunagin M, Coté C, Wan Y, Bayatpour S, Drexler HL, Alexander KA, Chen F, Wassie AT, Patel R, Pham K, Boyden ES, Berger S, Phillips-Cremins J, Churchman LS, Raj A
eLife 12

- 2024
Proteome-scale movements and compartment connectivity during the eukaryotic cell cycle
Litsios A, Grys BT, Kraus OZ, Friesen H, Ross C, Masinas MPD, Forster DT, Couvillion MT, Timmermann S, Billmann M, Myers C, Johnsson N, Churchman LS, Boone C, Andrews BJ
Cell 187, 1490–1507.e21

- 2024
Single-nucleoid architecture reveals heterogeneous packaging of mitochondrial DNA
Isaac RS, Tullius TW*, Hansen KG*, Dubocanin D, Couvillion M, Stergachis AB#, Churchman LS#
Nature Structural & Molecular Biology 31, 568–577

- 2024
Structural basis of LRPPRC-SLIRP-dependent translation by the mitoribosome
Singh V, Moran JC, Itoh Y, Soto IC, Fontanesi F, Couvillion M, Huynen MA, Churchman LS, Barrientos A, Amunts A
Nature Structural & Molecular Biology 31(12), 1838–1847

- 2023
Regulators of mitonuclear balance link mitochondrial metabolism to mtDNA expression
Kramer NJ*, Prakash G*, Choquet K, Soto I, Petrova B, Merens HE, Kanarek N, Churchman LS
Nature Cell Biology 25, 1575–1589

- 2023
RNA Polymerase II pausing temporally coordinates cell cycle progression and erythroid differentiation
Martell DJ, Merens HE, Caulier A, Fiorini C, Ulirsch J, Ietswaart R, Choquet K, Graziadei G, Brancaleoni V, Cappellini MD, Scott C, Roberts N, Proven M, Roy NBA, Babbs C, Higgs DR, Sankaran VG#, Churchman LS#
Developmental Cell 58, 2112–2127.e4
- 2023
- 2023
Intronic small nucleolar RNAs regulate host gene splicing through base pairing with their adjacent intronic sequences
Bergeron D, Faucher-Giguère L, Emmerichs A-K, Choquet K, Song KS, Deschamps-Francoeur G, Fafard-Couture É, Rivera A, Couture S, Churchman LS, Heyd F, Abou Elela S, Scott MS
Genome Biology 24, 160

- 2023
Elongation rate of RNA polymerase II affects pausing patterns across 3' UTRs
Khitun A, Brion C, Moqtaderi Z, Geisberg JV, Churchman LS, Struhl K
Journal of Biological Chemistry 299(11), 105289

- 2022
Balanced mitochondrial and cytosolic translatomes underlie the biogenesis of human respiratory complexes
Soto I*, Couvillion M*, Hansen KG, McShane E, Moran JC, Barrientos A, Churchman LS
Genome Biology 23(1), 170

- 2022
Transcription elongation is finely tuned by dozens of regulatory factors
Couvillion M*, Harlen KM*, Lachance KC*, Trotta KL, Smith E, Brion C, Smalec BM, Churchman LS
eLife 11

- 2021
- 2021
Essential histone chaperones collaborate to regulate transcription and chromatin integrity
Viktorovskaya O, Chuang J, Jain D, Reim NI, López-Rivera F, Murawska M, Spatt D, Churchman LS, Park PJ, Winston F
Genes & Development 35(9-10), 698–712

- 2021
GeneWalk identifies relevant gene functions for a biological context using network representation learning
Ietswaart R, Gyori BM, Bachman JA, Sorger PK, Churchman LS
Genome Biology 22(1), 55

- 2020
Single-molecule regulatory architectures captured by chromatin fiber sequencing
Stergachis AB, Debo BM, Haugen E, Churchman LS, Stamatoyannopoulos JA
Science 368, 1449–1454

- 2020
- 2020
Splicing kinetics and coordination revealed by direct nascent RNA sequencing through nanopores
Drexler HL, Choquet K, Churchman LS
Molecular Cell 77(5), 985–998.e8

- 2019
Proteotoxicity from aberrant ribosome biogenesis compromises cell fitness
Tye BW, Commins N, Ryazanova LV, Wühr M, Springer M, Pincus D, Churchman LS
eLife 8

- 2018
Spt6 is required for the fidelity of promoter selection
Doris SM, Chuang J, Viktorovskaya O, Murawska M, Spatt D, Churchman LS, Winston F
Molecular Cell 72(4), 687–699.e6

- 2018
Cell-cycle modulation of transcription termination factor Sen1
Mischo HE, Chun Y, Harlen KM, Smalec BM, Dhir S, Churchman LS, Buratowski S
Molecular Cell 70(2), 312–326.e7

- 2018
Set2 methyltransferase facilitates cell cycle progression by maintaining transcriptional fidelity
Dronamraju R, Jha DK, Eser U, Adams AT, Dominguez D, Choudhury R, Chiang Y-C, Rathmell WK, Emanuele MJ, Churchman LS, Strahl BD
Nucleic Acids Research 46(3), 1331–1344

- 2017
The ground state and evolution of promoter region directionality
Jin Y*, Eser U*, Struhl K#, Churchman LS#
Cell 170(5), 889–898.e10

- 2017
BET bromodomain proteins function as master transcription elongation factors independent of CDK9 recruitment
Winter GE*, Mayer A*, Buckley DL*, Erb MA, Roderick JE, Vittori S, Reyes JM, di Iulio J, Souza A, Ott CJ, Roberts JM, Zeid R, Scott TG, Paulk J, Lachance K, Olson CM, Dastjerdi S, Bauer S, Lin CY, Gray NS, Kelliher MA, Churchman LS#, Bradner JE#
Molecular Cell 67(1), 5–18.e19

- 2017
Total RNA-seq to identify pharmacological effects on specific stages of mRNA synthesis
Boswell SA, Snavely A, Landry HM, Churchman LS, Gray JM, Springer M
Nature Chemical Biology 13(5), 501–507

- 2017
Subgenic Pol II interactomes identify region-specific transcription elongation regulators
Harlen KM, Churchman LS
Molecular Systems Biology 13(1), 900

- 2016
Comprehensive RNA polymerase II interactomes reveal distinct and varied roles for each phospho-CTD residue
Harlen KM, Trotta KL, Smith EE, Mosaheb MM, Fuchs SM, Churchman LS
Cell Reports 15(10), 2147–2158

- 2016
Synchronized mitochondrial and cytosolic translation programs
Couvillion MT, Soto IC, Shipkovenska G, Churchman LS
Nature 533(7604), 499–503

- 2015
Native elongating transcript sequencing reveals human transcriptional activity at nucleotide resolution
Mayer A*, di Iulio J*, Maleri S, Eser U, Vierstra J, Reynolds A, Sandstrom R, Stamatoyannopoulos JA, Churchman LS
Cell 161(3), 541–554

- 2015
Single mammalian cells compensate for differences in cellular volume and DNA copy number through independent global transcriptional mechanisms
Padovan-Merhar O, Nair GP, Biaesch AG, Mayer A, Scarfone S, Foley SW, Wu AR, Churchman LS, Singh A, Raj A
Molecular Cell 58(2), 339–352

- 2014
Reviews, commentaries and protocols
- 2026
- 2026
A splicing factor's unexpected detour to the mitochondrial surface
Raicu AM, Churchman LS
Molecular Cell 86(7), 1195–1196

- 2025
The regulation and function of post-transcriptional RNA splicing
Choquet K, Patop IL, Churchman LS
Nature Reviews Genetics 26(6), 378–394

- 2024
Central dogma rates in human mitochondria
McShane E, Churchman LS
Human Molecular Genetics 33(R1), R34–R41

- 2024
- 2024
Human mitoribosome profiling: a re-engineered approach tailored to study mitochondrial translation
Soto I, Couvillion M, Churchman LS
Methods in Molecular Biology 2661, 257–280

- 2021
Revealing nascent RNA processing dynamics with nano-COP
Drexler HL, Choquet K, Merens HE, Tang PS, Simpson JT, Churchman LS
Nature Protocols 16(3), 1343–1375

- 2019
The long and the short of the RNA polymerase C-terminal domain and phase separation
Isaac RS, Churchman LS
Molecular Cell 73(6), 1087–1088

- 2018
The multiple levels of mitonuclear coregulation
Isaac RS, McShane E, Churchman LS
Annual Review of Genetics 52, 511–533

- 2017
Pause & go: from the discovery of RNA polymerase pausing to its functional implications
Mayer A, Landry HM, Churchman LS
Current Opinion in Cell Biology 46, 72–80

- 2017
A detailed protocol for subcellular RNA sequencing (subRNA-seq)
Mayer A, Churchman LS
Current Protocols in Molecular Biology 120, 4.29.1–4.29.18

- 2017
The code and beyond: transcription regulation by the RNA polymerase II carboxy-terminal domain
Harlen KM, Churchman LS
Nature Reviews Molecular Cell Biology 18(4), 263–273

- 2017
Not just noise: genomics and genetics bring long noncoding RNAs into focus
Churchman LS
Molecular Cell 65(1), 1–2

- 2017
Mitochondrial ribosome (mitoribosome) profiling for monitoring mitochondrial translation in vivo
Couvillion MT, Churchman LS
Current Protocols in Molecular Biology 119, 4.28.1–4.28.25

- 2016
Genome-wide profiling of RNA polymerase transcription at nucleotide resolution in human cells with native elongating transcript sequencing
Mayer A, Churchman LS
Nature Protocols 11(4), 813–833

- 2012
Native elongating transcript sequencing (NET-seq)
Churchman LS, Weissman JS
Current Protocols in Molecular Biology Chapter 4, Unit 4.14.1–17

Earlier publications
- 2013
Single-molecule fluorescence imaging of processive myosin with enhanced background suppression using linear zero-mode waveguides (ZMWs) and convex lens induced confinement (CLIC)
Elting MW, Leslie SR, Churchman LS, Korlach J, McFaul CMJ, Leith JS, Levene MJ, Cohen AE, Spudich JA
Optics Express 21(1), 1189–1202

- 2012
Colocalization of fluorescent probes: accurate and precise registration with nanometer resolution
Churchman LS, Spudich JA
Cold Spring Harbor Protocols 2012(2), 141–149

- 2012
Single-molecule high-resolution colocalization of single probes
Churchman LS, Spudich JA
Cold Spring Harbor Protocols 2012(2), 242–245

- 2011
H3K4 trimethylation by Set1 promotes efficient termination by the Nrd1-Nab3-Sen1 pathway
Terzi N, Churchman LS, Vasiljeva L, Weissman J, Buratowski S
Molecular and Cellular Biology 31(17), 3569–3583

- 2011
Nascent transcript sequencing visualizes transcription at nucleotide resolution
Churchman LS, Weissman JS
Nature 469(7330), 368–373

- 2010
Optimized localization analysis for single-molecule tracking and super-resolution microscopy
Mortensen KI*, Churchman LS*, Spudich JA, Flyvbjerg H
Nature Methods 7(5), 377–381

- 2009
Rapid creation and quantitative monitoring of high coverage shRNA libraries
Bassik MC, Lebbink RJ, Churchman LS, Ingolia NT, Patena W, LeProust EM, Schuldiner M, Weissman JS, McManus MT
Nature Methods 6(6), 443–445

- 2007
Colocalization of fluorescent probes: accurate and precise registration with nanometer resolution
Churchman LS, Spudich JA
Single-Molecule Techniques: A Laboratory Manual, 73–84
- 2006
A non-Gaussian distribution quantifies distances measured with fluorescence localization techniques
Churchman LS, Flyvbjerg H, Spudich JA
Biophysical Journal 90(2), 668–671

- 2005
Single molecule high-resolution colocalization of Cy3 and Cy5 attached to macromolecules measures intramolecular distances through time
Churchman LS, Ökten Z, Rock RS, Dawson JF, Spudich JA
Proceedings of the National Academy of Sciences 102(5), 1419–1423

- 2004
Myosin VI walks hand-over-hand along actin
Ökten Z, Churchman LS, Rock RS, Spudich JA
Nature Structural & Molecular Biology 11(9), 884–887

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