56. Super-resolution imaging of proteins inside live mammalian cells with mLIVE-PAINT

Bhaskar, H., Gidden, Z., Virdi, G., Kleinjan, D., Rosser, S.J., Gandhi, S., Regan, L., Horrocks, M.H.

bioRxiv, Pre-print, 2024.

55. Sequential replacement of PSD95 subunits in postsynaptic supercomplexes is slowest in the cortex

Morris, K., Bulovaite, E., Kaizuka, T., Schnorrenberg, S., Adams, C., Komiyama, N., Mendive-Tapia, L., Grant, S. G. N., Horrocks, M. H.

eLife, Reviewed Preprint, 2024.

54. Astrocytes modulate Aquaporin-4 array size in response to adrenergic signaling and osmolarity changes.

Zepernick, A., Metodieva, V., Pelegrina-Hidalgo, N., Lippert, A.H., Horrocks, M.H., Varela, J.A.

Nanoscale, 16, 9576-9582, 2024.

53. RNA aptamer reveals nuclear TDP-43 pathology is an early aggregation event that coincides with STMN-2 cryptic splicing and precedes clinical manifestation in ALS.

Spence, H., Waldron, F. M., Saleeb, R. S., Brown, A.-L., Rifai, O. M., Gilodi, M., Read, F., Roberts, K., Milne, G., Wilkinson, D., O'Shaughnessy, J., Pastore, A., Fratta, P., Shneider, N., Tartaglia, G. G., Zacco, E., Horrocks, M. H., Gregory, J. M.

Acta Neuropathologica, 147, 50, 2024.

52. GLP1 Receptor Agonism Ameliorates Parkinson's Disease through Modulation of Neuronal Insulin Signalling and Glial Suppression

Athauda, D., Evans, J. R., Mahoney-Sanchez, L., Virdi, G. S., Lopez-Garcia, P., Wernick, A., Wagen, A., D'Sa, K., Lachica, J., Strohbuecker, S., Vecchi, G., Leighton, C., Saleeb, R. S., O'Shaughnessy, J., Toomey, C. E., Vijiaratnam, N., Girges, C., Li, Y., Mustapic, M., Ismail, K., Davies, M., Kapogiannis, D., Choi, M. L., Ryten, M., Horrocks, M. H., Greig, N., Foltynie, T., Gandhi, S.

bioRxiv, Pre-print, 2024.


51. Two-Color Coincidence Single-Molecule Pull-down for the Specific Detection of Disease-Associated Protein Aggregates

Saleeb, R.S., Leighton, C., Lee, J., O'Shaughnessy, J., Jeacock, K., Chappard, A., Cumberland, R., Zhao, T., Ball, S.R., Sunde, M., Clarke, D.J., Piche, K., McPhail, J.A., Louwrier, A., Angers, R., Gandhi, S., Downey, P., Kunath, T., Horrocks, M.H.

Science Advances, 9, eadi7359, 2023.

50. Imaging Proteins Sensitive to Direct Fusions Using Transient Peptide-Peptide Interactions.

Gidden, Z., Oi, C., Johnston, E.J., Konieczna, Z., Bhaskar, H., Mendive-Tapia, L., de Moliner, F., Rosser, S.J., Mochrie, S. G. J., Vendrell, M., Horrocks, M. H., Regan, L.

ACS Nano Letters, 23, 22, 10633-10641, 2023.

49. Synaptic Expression of TAR-DNA-Binding Protein 43 in the Mouse Spinal Cord Determined Using Super-Resolution Microscopy

Broadhead, M. J., Ayvazian-Hancock, A., Doucet, K., Kantelberg, O. G., Zhu, F., Motherwell, L., Grant, S. G. N., Horrocks, M. H., Miles, G. B.

Frontiers in Molecular Neuroscience, 16, 2023.

48. Prediction of mechanistic subtypes of Parkinson's using patient derived stem cell models

D, Karishma, Evans, J.R., Virdi, G.S., Vecchi, G., Adam, A., Bertolli, O., Fleming, J., Chang, H., Leighton, C., Horrocks, M.H., Athauda, D., Choi, M.L., Gandhi, S.

Nature Machine Intelligence, 5, 933-946, 2023.

47. Lipid-Induced Polymorphic Amyloid Fibrils Formation by alpha-Synuclein.

Singh, B. P., Morris, R. J., Kunath, T., MacPhee, C. E., Horrocks, M. H.

Protein Science, 32:e4736, 2023.

46. Synaptic Oligomeric Tau in Alzheimer's Disease - a Potential Culprit in the Spread of Tau Pathology through the Brain

Colom-Cadena, M., Davies, C., Sirisi, S., Lee, J.-E., Simzer, E., Tzioras, M., Querol-Vilaseca, M., Sanchez-Aced, E., Chang, Y. Y., Holt, K., McGeachan, R., Rose, J., Tulloch, J., Wilkins, L., Smith, C., Andrian, T., Belbin, O., Pujals, S., Horrocks, M. H., Llee, A., Spires-Jones, T.

Neuron, 111, 1-14, 2023.

45. Determining the Location of the alpha-Synuclein Dimer Interface Using Native Top-Down Fragmentation and Isotope Depletion-Mass Spectrometry

Jeacock, K., Chappard, A., Gallagher, K. J., Mackay, C. L., Kilgour, D. P. A., Horrocks, M. H., Kunath, T., Clarke, D. J.

Journal of the American Society for Mass Spectrometry, 34, 5, 847-856, 2023.

44. Single-Molecule Two-Color Coincidence Detection of Unlabeled alpha-Synuclein Aggregates

Chappard, A., Leighton, C., Saleeb, R.S., Jeacock, K., Ball, S.R., Morris, K., Kantelberg, O., Lee, J., Zacco, E., Pastore, A., Sunde, M., Clarke, D.J., Downey, P., Kunath, T., Horrocks, M.H.

Angewandte Chemie International Edition, e202216771, 2023.


43. Live-cell super-resolution imaging of actin using LifeAct-14 with a PAINT-based approach

Bhaskar, H., Kleinjan, D.J., Oi, C., Gidden, Z., Rosser, S., Horrocks, M.H.*, Regan, L.*, *Joint corresponding authors

Protein Science, e4558, 2022.

42. Small Fluorogenic Amino Acids for Peptide-Guided Background-Free Imaging

de Moliner, F., Konieczna, Z., Mendive-Tapia, L., Saleeb, R.S., Morris, K., Gonzalez-Vera, J.A., Kaizuka, T., Grant, S.G.N., Horrocks, M.H.*, Vendrell, M.*, *Joint corresponding authors

Angewandte Chemie International Edition, e202216231, 2022.

41. Protein Aggregation and Calcium Dysregulation Are the Earliest Hallmarks of Synucleinopathy in Human Midbrain Dopaminergic Neurons.

Virdi, G. S., Choi, M. L., Evans, J. R., Yao, Z., Athauda, D., Strohbuecker, S., Nirujogi, R. S., Wernick, A. I., Pelegrina-Hidalgo, N., Leighton, C., Saleeb, R. S., Kopach, O., Alrashidi, H., Melandri, D., Perez-Lloret, J., Angelova, P. R., Sylantyev, S., Eaton, S., Heales, S., Rusakov, D. A., Alessi, D. R., Kunath, T., Horrocks, M. H., Abramov, A. Y., Patani, R., Gandhi, S.

npj Parkinson's Disease, 8, 1, 44562, 2022.

40. pTDP-43 aggregates accumulate in non-central nervous system tissues prior to symptom onset in amyotrophic lateral sclerosis: a case series linking archival surgical biopsies with clinical phenotypic data.

Pattle, S.B., O'Shaughnessy, J., Kantelberg, O., Rifai, O.M., Pate, J., Nellany, K., Hays, N., Arends, M.J., Horrocks, M.H., Waldron, F.M., Gregory, J.M.

The Journal of Pathology: Clinical Research, 9, 44-55, 2022.

39. Structural Conversion of alpha-Synuclein at the Mitochondria Induces Neuronal Toxicity

Choi, M. L., Chappard, A., Singh, B. P., Maclachlan, C., Rodrigues, M., Fedotova, E., Berezhnov, A. V., De, S., Peddie, C., Athauda, D., Virdi, G. S., Zhang, W., Evans, J. R., Wernick, A., Zanjani, Z. S., Angelova, P. R., Esteras, N., Vinikurov, A., Morris, K., Jeacock, K., Tosatto, L., Little, D., Gissen, P., Clarke, D. J., Kunath, T., Collinson, L., Klenerman, D., Abramov, A. Y.*, Horrocks, M. H.*, Gandhi, S.*, *Joint corresponding authors

Nature Neuroscience, 25, 9, 1134-1148, 2022.

38. Probing TDP-43 Condensation Using an in Silico Designed Aptamer.

Zacco, E., Kantelberg, O., Milanetti, E., Armaos, A., Panei, F. P., Gregory, J., Jeacock, K., Clarke, D. J., Chandran, S., Ruocco, G., Gustincich, S., Horrocks, M. H.*, Pastore, A.*, Tartaglia, G. G.*, *Joint corresponding authors

Nature Communications, 13, 1, 41275, 2022.

37. Self-Assembling Protein Surfaces for In Situ Capture of Cell-Free-Synthesized Proteins.

Thornton, E. L., Paterson, S. M., Gidden, Z., Horrocks, M. H., Laohakunakorn, N., Regan, L.

Frontiers in Bioengineering and Biotechnology, 1193, 2022.


36. PAINT using proteins: A new brush for super-resolution artists

Oi, C., Mochrie, S.J., Horrocks, M.H., Regan, L.

Protein Science, 29, 11, 2142-2149, 2020.

35. A step-by-step protocol for performing LIVE-PAINT super-resolution imaging of proteins in live cells using reversible peptide-protein interactions

Oi, C., Gidden, Z., Holyoake, L., Kantelberg, O., Mochrie, S., Horrocks, M.H.*, Regan, L.*, *Joint corresponding authors

Nature Protocols, Protocol, 2020.

34. LIVE-PAINT allows super-resolution microscopy inside living cells using reversible peptide-protein interactions

Oi, C., Gidden, Z., Holyoake, L., Kantelberg, O., Mochrie, S., Horrocks, M.H.*, Regan, L.*, *Joint corresponding authors

Communications Biology, 3, 1, 45200, 2020.

33. Alpha synuclein aggregation drives ferroptosis: an interplay of iron, calcium and lipid peroxidation

Angelova, P.R., Choi, M-L., Berezhnov, A.V., Horrocks, M.H., Hughes, C.D., De, S., Rodrigues, M., Yapom, R., Little, D., Dolt, K.S., Kunath, T., Devine, M.J., Gissen, P., Shchepinov, M.S., Sylantyev, S., Pavlov, E.V., Klenerman, D., Abramov, A.Y., Gandhi, S.

Cell Death and Differentiation, 27, 10, 2781-2796, 2020.


32. ASYN-CONA, a novel bead-based assay for detecting early stage alpha-synuclein aggregation.

Pérez-Pi, I., Evans, D.A., Horrocks, M.H., Pham, N.A., Dolt, K.S., Koszela, J., Kunath, T., Auer, M.

Analytical Chemistry, 91, 9, 5582-5590, 2019.

31. SCOTfluors: Small, Conjugatable, Orthogonal and Tunable Fluorophores for in vivo Imaging of Cell Metabolism

Benson, S., Fernandez, A., Barth, N., de Moliner, F., Horrocks, M.H., Herrington, S., Abad, J.L., Delgado, A., Kelly, L., Chang, Z., Feng, Y., Nishiura, M., Hori, Y., Kikuchi, K., Vendrell, M.

Angewandte Chemie International Edition, 131, 21, 6985-6989, 2019.


30. Mapping Surface Hydrophobicity of alpha-Synuclein Oligomers at the Nanoscale

Lee, J., Sang J.C., Rodrigues M., Carr, A.R., Horrocks, M.H., De, S., Bongiovanni, M.N., Flagmeier, P.,  Dobson, C.M., Wales D.J., Lee, S.F., Klenerman, D.

Nano Letters, 18, 12, 7494-7501, 2018.

29. Nanoscopic characterization of individual endogenous protein aggregates in human neuronal cells

Whiten, D.R., Zuo, Y., Calo, L., Choi, M., De, S., Flagmeier, P., Wirthensohn, D.C., Kundel, F., Ranasinghe, R.T., Sanchez, S.E., Athauda, D., Lee, S.F., Dobson, C.M., Gandhi, S., Spillantini, M., Klenerman, D., Horrocks, M.H.

ChemBioChem, 19, 19, 2001, 2018.

28. Extrinsic Amyloid-Binding Dyes for the Detection of Individual Protein Aggregates in Solution

Taylor, C.G., Meisl, G., Horrocks, M.H., Zetterberg, H., Knowles, T.P.J., Klenerman, D.

Analytical Chemistry, 90, 17, 10385-10393, 2018.

27. Single-Molecule Characterization of the Interactions between Extracellular Chaperones and Toxic alpha-Synuclein Oligomers

Whiten, D.R., Dezerae, D., Horrocks, M.H., Taylor, C.G., De, S., Flagmeier, P., Tosatto, L., Kumita, J.R., Ecroyd, H., Dobson, C.M., Klenerman, D.

Cell Reports, 23, 12, 3492-3500, 2018.

26. alpha-Synuclein Oligomers Interact with ATP Synthase and Open the Permeability Transition Pore in Parkinson's Disease

Ludtmann, M. H. R.*, Angelova, P. R.*, Horrocks, M. H.*, Choi, M. L., Rodrigues, M., Baev, A. Y., Berezhnov, A. V., Yao, Z., Little, D., Banushi, B., Al-Menhali, A.S., Ranasinghe, R.T., Whiten, D.R., Yapom, R., Dolt, K.S., Devine, M.J., Gissen, P., Kunath, T., Jaganjac, M., Pavlov, E.V., Klenerman, D., Abramov, A.Y., S. Gandhi, S., *Joint first authors

Nature Communications, 9, 1, 42370, 2018.

25. A sticky situation: aberrant protein-protein interactions in Parkinson's disease

Brown, J.W.P., Horrocks, M.H.

Seminars in Cell & Developmental Biology, 99, 65-77, 2018.

24. Shedding light on aberrant interactions: A review of modern tools for studying protein aggregates

Kundel, F., Tosatto, L.T., Whiten, D.R., Wirthensohn, D.C., Horrocks, M.H.*, Klenerman, D.*, *Joint corresponding authors.

The FEBS Journal, 285, 19, 3604-3630, 2018.

23. Detecting RNA base methylations in single cells by in situ hybridization

Ranasinghe, R.T., Challand, M.R., Ganzinger, K.A., Lewis, B.W., Softley, C., Schmied, W.H., Horrocks, M.H., Shivji, N., Chin, J.W, Spencer, J., Klenerman, D.

Nature Communications, 9, 1, 45200, 2018.

22. The small heat shock protein Hsp27 binds alpha-synuclein fibrils, preventing elongation and cytotoxicity

Cox, D., Whiten, D.R., Brown, J.W.P., Horrocks, M.H., San Gil, R., Dobson, C.M., Klenerman, D., van Oijen, A.M., Ecroyd, H.

Journal of Biological Chemistry, 293, 12, 4486-4497, 2018.

21. Hsp70 Inhibits the Nucleation and Elongation of Tau and Sequesters Tau Aggregates with High Affinity

Kundel, F., De, S., Flagmeier, P., Horrocks, M.H., Kjaergaard, M., Shammas, S.L., Jackson, S.E., Dobson, C.M., Klenerman, D.

ACS Chemical Biology, 13, 3, 636-646, 2018.


20. Nanobodies raised against monomeric alpha-synuclein inhibit fibril formation and destabilize toxic oligomeric species.

Iljina, M., Hong, L, Horrocks, M.H., Ludtmann, M.H., Choi, M.L., Hughes, C.D. Ruggeri, F.S. Guilliams, T., Buell, A.K., Lee, J.E., Gandhi, S., Lee, S.F, Bryant, C.E., Vendruscolo, M., Knowles, T.P.J., Dobson, C.M., De Genst, E., Klenerman, D.

BMC Biology, 15, 1, 41640, 2017.

19. PEGylated liposomes associate with Wnt3A protein and expand putative stem cells in human bone marrow populations.

Janeczek, AA, Scarpa, E., Horrocks M.H, Tare, R.S., Rowland, C.A., Jenner, D., Newman, T.A., Oreffo, R.O.C., Lee, S.F., Evans, N.D.

Nanomedicine, 12, 8, 845-863, 2017.

18. Co-aggregation of κ-Casein and β-Lactoglobulin Produces Morphologically Distinct Amyloid Fibrils.

Raynes, J.K., Crepin, P., Horrocks, M.H., Carver, J.A., Day, L.

Small, 13, 14, 1603591, 2017.


17. Multi-dimensional super-resolution imaging enables surface hydrophobicity mapping.

Bongiovanni, M.*, Godet, J.*,Horrocks, M.H.*, Tosatto, L., Carr, A.R, Wirthensohn, D.C., Ranasinghe, R.T., Lee, J., Ponjavic, A., Fritz, J.V., Dobson, C.M., Klenerman, D., Lee, S.F., *Joint first authors

Nature Communications, 7, 1, 45170, 2016.

16. The remarkably low affinity of CD4/peptide-major histocompatibility complex class II protein interactions.

Jonsson P., Southcombe J., Mafalda A., Santos A.M., Fernandes R.A., McColl J., Chang V.C., Godkin A., Dunne P.D., Horrocks M.H., Palayret M., Fugger L., Xu X., Davis S.J., Klenerman D.

Proceedings of the National Academy of Sciences, 113, 20, 5682-5687, 2016.

15. PSD95 nanoclusters are postsynaptic building blocks in hippocampus circuits.

Broadhead M.J., Horrocks M.H., Zhu F., Muresan L., Benavides-Piccione R., DeFelipe J., Fricker D., Kopanitsa M.V., Duncan R.R., Klenerman D., Komiyama N.H., Lee S.F., Grant S.G.N.

Scientific Reports, 6, 1, 41640, 2016.

14. A kinetic model of the aggregation of alpha-synuclein provides insights into prion-like spreading.

Iljina,M., Garciaa, G.A., Horrocks, M.H., Tosatto, L., Choi, M.L., Ganzinger, K.A., Abramov, A.Y., Gandhi, S., Wood, N.W., Cremades, N.,Dobson, C.M. , Knowles, T.P.J., Klenerman, D.

Proceedings of the National Academy of Sciences, 113, 9, E1206-E1215, 2016.

13. Single-molecule imaging of individual amyloid protein aggregates in human biofluids.

Horrocks M. H., Lee S.F., Gandhi S., Magdalinou N.K., Chen S.W., Devine M.J., Toasatto, L., Kjaergaard M., Beckwith J.S., Zetterberg H., Iljina M., Cremades N., Dobson C.M., Wood N.W., Klenerman D.

ACS Chemical Neuroscience, 7, 3, 399-406, 2016.

12. Calcium is a key factor in alpha-synuclein induced neurotoxicity.

Angelova, P., Ludtmann, M., Horrocks, M.H., Negoda, A., Cremades, N., Klenerman, D., Dobson, C.M., Wood, N., Pavlov, E., Gandhi, S., Abramov, A.

Journal of Cell Science, 129, 9 , 1792-1801, 2016.


11. Alpha-synuclein oligomers interact with metal ions to induce oxidative stress and neuronal death in Parkinson's disease.

Deas, E., Cremades, N., Angelova, P.R., Ludtmann, M., Yao, Z., Chen, S., Horrocks, M.H., Banushi, B., Little, D., Devine, M., Gissen P., Klenerman, D., Dobson, C.M., Wood, N., Gandhi, S., Abramov. A.Y.

Antioxidants and redox signaling, 24, 7, 376-391, 2015.

10. Single-molecule FRET studies on alpha-synuclein oligomerization of Parkinson's disease genetically related mutants.

Horrocks, M.H., Tosatto, L., Dear, A.J., Knowles, T.P.J., Dalla Serra, M., Dobson, C.M., Klenerman, D.

Scientific Reports, 5, 1, 45261, 2015.

9. Fast flow microfluidics and single-molecule fluorescence for the rapid characterization of alpha-synuclein oligomers.

Horrocks, M.H., Tosatto, L., Dear, A.J., Garcia, G.A., Iljina, M., Cremades, N., Dalla Serra, M., Knowles, T.P.J., Dobson, C.M., Klenerman, D.

Analytical Chemistry, 87, 17, 8818-8826, 2015.

8. Aggregated alpha-synuclein and complex I deficiency: exploration of their relationship in differentiated neurons.

Reeve, A.K., Ludtmann, M.H.R, Angelova, P.R., Simcox, E.M., Horrocks, M.H., Klenerman, D., Gandhi, S., Turnbull, D.M., Abramov, A.Y.

Cell Disease and Death, 6,7, e1820-e1820, 2015.

7. Lipid peroxidation is essential for alpha-synuclein-induced cell death.

Angelova, P.R., Horrocks, M.H., Klenerman D, Gandhi S, Abramov, A.Y., Shchepinov, M.S

Journal of Neurochemistry, 133, 4, 582-589, 2015.

6. A mechanistic model of tau amyloid aggregation based on direct observation of oligomers.

Shammas, S. L., Garcia, G. A., Kumar S., A., Kjaergaard M., Horrocks, M.H., Shivji N., Mandelkow, E., Knowles T.P.J., Mandelkow E., Klenerman, D.

Nature Communications, 6,1, 45200, 2015.


5. Bayesian Inference of Accurate Population Sizes and FRET Efficiencies from Single Diffusing Biomolecules.

Murphy, R.M., Danezis, D., Horrocks, M.H., Jackson, S.E., Klenerman, D.

Analytical Chemistry, 86, 17, 8603-8612, 2014.

4. The changing point-spread function: single-molecule-based super-resolution imaging.

Horrocks, M.H., Palayret, M., Klenerman, D., Lee, S.F.

Histochemistry and Cell Biology, 141, 6, 577-585, 2014.


3. Single-molecule measurements of transient biomolecular complexes through microfluidic dilution

Horrocks, M.H., Rajah, L., Jonsson, P., Kjaergaard, M., Vendruscolo, M., Knowles, T.P.J., Klenerman, D.

Analytical Chemistry, 85, 14, 6855-6859, 2013.


2. Ubiquitin chain conformation regulates recognition and activity of interacting proteins.

Ye, Y., Blaser, G., Horrocks, M.H., Ruedas-Rama, M.J., Ibrahim, S., Zhukov, A.A., Orte, A., Klenerman, D., Jackson, S.E., Komander, D.

Nature, 492, 7428, 266-270, 2012.

1. Single molecule fluorescence under conditions of fast flow.

Horrocks, M.H., Li, H., Shim, J.-U., Ranasinghe, R.T., Clarke, R.W., Huck, W.T.S., Abell, C., Klenerman, D.

Analytical Chemistry, 84, 1, 179-185, 2012.