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Department of Pharmacology




Maintaining flippase activity in procoagulant platelets is a novel approach to reducing thrombin generation. Millington-Burgess SL, Harper MT. J Thromb Haemost, in press (link)

Epigenetic Regulation of F2RL3 Associates with Myocardial Infarction and Platelet Function.Corin LJ, White SJ, Taylor A, Williams CM, Taylor K, can den Mosch MT, Teasdale JE, Jones ML, Bond M, Harper MT, Falk L, Groom A, Haxell GGJ, Paternoster L, Munafo M, Nordestgaard BG, Tybjærg-Hanson A, Bojesen SE, Relton CL, Min JL, Davey Smith G, Mumford AD, Poole AW, Timpson NJ. Circ Res, in press (link)

Extracellular matrix composition regulates endothelial inflammation and neutrophil transmigration in an inflammation-on-a-chip model. Riddle RB, Jennbacken K, Hansson KM, Harper MT. Sci Rep (in press)

Protease-activated receptor antagonism prevents occlusion when dual antiplatelet therapy is insufficient in a microfluidic model of occlusive thrombosis. Berry JE, Harper MT. Res Prac Thromb Haemost 6: e12703. (link)

Q94 is not a selective modulator of PAR1 in platelets. Francis LRA, Millington-Burgess SL, Rahman T, Harper MT. Platelets, in press



An "occlusive thrombosis-on-a-chip" microfluidic device for investigating the effect of anti-thrombotic drugs. Berry J, Peudecerf FJ, Masters NA, Neeves KB, Goldstein RE, Harper MT. Lab Chip, 21, 4104-4117. (link)

Epigallocatechin gallate inhibits release of extracellular vesicles from platelets without inhibiting phosphatidylserine exposure. Millington-Burgess SL, Harper MT. Sci Rep, 11, 17678. (link)

A double-edged sword: Antibody-mediated procoagulant platelets in COVID-19. Millington-Burgess SL, Harper MT. Platelets, 32, 579-581. (link)

Cytosolic and mitochondrial Ca2+ signalling in procoagulant platelets. Millington-Burgess SL, Harper MT.  Platelets, 32, 855-862. (link)

Establishing an analogue-based in silico pipeline in the pursuit of novel inhibitory scaffolds against the SARS coronavirus 2 papain-like protease. Hajbabaie R, Harper MT, Rahman T. Molecules, 26, 1134. (link)

Comparison of Putative BH3 Mimetics AT-101, HA14-1, Sabutoclax and TW-37 With ABT-737 in platelets. Wei H, Harper MT. Platelets, 35, 105-112. (

High extracellular phosphate increases platelet polyphosphate content. Abbasian N, Harper MT. Platelets, 32, 992-994. (Full article: High extracellular phosphate increases platelet polyphosphate content (



R5421 is not a selective inhibitor platelet scramblase activity. Millington-Burgess SL, Rahman T, Harper MT. Br J Pharmacol, 177, 4007-4020. (link)

Supramaximal calcium signaling triggers procoagulant platelet formation. Abbasian N, Millington-Burgess SL, Chabra S, Malcor JD, Harper MT. Blood Adv, 4, 154-164. (

Novel small molecule scaffolds as candidates against the SARS Coronavirus 2 main protease: a fragment-guided in silico approach. Augustin TL, Hajbabaie R, Harper MT, Rahman T. Molecules, 25, 5501. (

2-Aminoethoxydiphenylborate (2-APB) inhibits release of phosphatidylserine-exposing extracellular vesicles from platelets. Wei H, Davies JE, Harper MT. Cell Death Discov, 6, 10. (

Elevated intracellular cAMP concentration mediates growth suppresion in glioma cells. Safitri D, Harris M, Potter H, Yan Yeung H, Winfield I, Kopanitsa L, Svensson F, Rahman T, Harper MT, Bailey D, Ladds G. Biochem Pharmacol, 174, 133823. (

Proposed model of the Dictyostelium cAMP receptors bound to cAMP. Greenhalgh JC, Chandran A, Ladds G, Harper MT, Rahman T. J Mol Graph Model, 100, 107662.



Gene of the Issue: ANO6 and Scott Syndrome. Millington-Burgess SL, Harper MT. Platelets, In Press (" target="_self" title="">

Abstracts of the 1st Platelet Society Meeting, Jesus College, Cambridge. Harper MT. Platelets, In Press

Using Yoda-1 to mimic laminar flow in vitro: a tool to simplify drug testing. Davies JE, Lopresto D, Apta BHR, Lin Z, Ma W, Harper MT. Biochem Pharmacol, 168, 473-480. (

ABT-737 triggers caspase-dependent inhibition of platelet procoagulant extracellular vesicle release during apoptosis and secondary necrosis in vitro. Wei H, Harper MT. Thromb Haemost, 19, 1665-1674. (

Platelet P-selectin triggers rapid surface exposure of tissue factor in monocytes. Ivanov II, Apta B, Bonna AM, Harper MT. Sci Rep, 9, 13397. (

Interactions between RAMP2 and CRF receptors: The effect of receptor subtypes, splice variants and cell context. Bailey S, Harris M, Barkan K, Winfield I, Harper MT, Simms J, Ladds G, Wheatley M, Poyner D.Biochim Biophys Acta Biomembr, 1861, 997-1003. (



Rapid kinetics of changes in oxygen consumption rate in thrombin-stimulated platelets measured by high-resolution respirometry. Sowton AP, Millington-Burgess SL, Murray AJ, Harper MT.  Biochem Biophys Res Comm, 503, 2721-2727.(

Lipid rafts are essential for release of phosphatidylserine-exposing extracellular vesicles from platelets. Wei H, Malcor JM, Harper MT. Sci Rep, 8, 9987. 

Cross-reactivity of HMGB1 antibodies for HMGB2. Davies JE, Apta BHR, Harper MT. J Immunol Methods, 456, 72-76.



Auranofin, a thioredoxin reductase inhibitor, causes platelet death through reactive oxygen species and calcium overload. Harper MT. Platelets

Extracellular chloride is required for efficient platelet aggregation. Taylor KA, Wilson DGS, Harper MT, Pugh N. Platelets, 29, 79-83. 



Thioredoxin inhibitors attenuate platelet function and thrombus formation.Metcalfe C, Ramasubramoni A, Pula G, Harper MT, Mundell SJ, Coxon CH. PLoS One, 11, e0163006.

Identification of roles for the SNARE-associated protein, SNAP29, in mouse platelets. Williams CM, Savage JS, Harper MT, Moore SF, Hers I, Poole AW. Platelets, 27, 286-294.



Harper MT, van den Bosch MT, Hers I, Poole AW. Platelet dense granule secretion defects may obscure our understanding of α-granule secretion mechanisms. Blood, 125, 3034-3036.

Williams CM, Harper MT, Goggs R, Walsh TG, Offermanns S, Poole AW. Leukemia-Associated RhoGEF (LARG) is not critical for RhoA regulation, yet is important for platelet activation and thrombosis in mice. J Thromb Haemost, 13, 2102-2107.

Moore SF, Williams CM, Brown E, Blair TA, Harper MT, Coward RJ, Poole AW, Hers I. Loss of the insulin receptor in megakaryocytes/platelets causes thrombocytosis and alterations in autocrine IGF signalling. Cardiovasc Res, 107, 9-19.

Walsh TG, Harper MT, Poole AW. SDF-1α is a novel autocrine activator of platelets operating through its receptor CXCR4. Cell Signal, 27, 37-46.

Golebiewska EM, Harper MT, Williams CM, Savage JS, Goggs R, Fischer von Mollard G, Poole AW. Syntaxin 8 regulates platelet dense granule secretion, aggregation, and thrombus stability. J Biol Chem, 290, 1536-1545.



Williams CM, Harper MT, Poole AW. PKCα negatively regulates in vitro proplatelet formation and in vivo platelet production in mice. Platelets, 25, 62-68.



Harper MT, Camacho Londono JE, Quick K, Camacho Londono J, Philipp SE, Birnbaumer L, Freichel M, Poole AW. Transient receptor potential channels function as a coincidence signal detector mediated phosphatidylserine exposure. Science Signalling, 6, ra50.

Harper MT, Poole AW. Chloride channels are necessary for full platelet phosphatidylserine exposure and procoagulant activity. Cell Death Dis, 4, e969.

Harper MT, Savage JS, Poole AW. Comment on “Platelet-derived nucleotides promote tumor cell transendothelial migration and metastasis via P2Y2 receptor” by Schumacher et al. Cancer Cell, 9, 287.

Harper MT, van den Bosch MTJ, Hers I, Poole AW. Absence of phenotype in platelets lacking the motor protein myosin Va. PLoS One, 8, e53239.

Goggs R, Harper MT, Pope RJP, Savage JS, Williams CW, Mundell SJ, Bass M, Mellor H, Poole AW. RhoG regulates platelet granule secretion and thrombus formation in mice. J Biol Chem, 228, 34217-34229.

Moore SF, Hunter RW, Harper MT, Savage JS, Saddiq S, Westbury SK, Poole AW, Mumford AD, Hers I. Dysfunction of PI3 kinase/Rap1b/integrin αIIbβ3 pathway underlies ex vivo platelet hypoactivity in essential thrombocythemia. Blood, 121, 1209-1219.

Savage JS, Williams CM, Konopatskaya O, Hers I, Harper MT, Poole AW. Munc13-4 is critical for thrombosis through regulating release of ADP from platelets. J Thromb Haemost, 11, 771-775.



Harper MT, Poole AW. Bcl-xL –inhibitory BH3 mimetic ABT-737 depletes platelet calcium stores. Blood, 119, 4337-4338.

Steele BM, Harper MT, Smolenski AP, Granville K, Poole AW, Fitzgerald DJ, Maguire PM. WNT3a modulates platelet function by regulating small GTPase activity. FEBS Lett, 586, 2267-2272.



Harper MT, Poole AW. Contribution of store-operated calcium entry and non-capacitative calcium entry to thrombin-induced calcium signalling in human platelets. Cell Calcium, 50, 351-358.

Harper MT, Poole AW. PKC inhibition enhances Ca2+ signalling and phosphatidylserine exposure downstream of protease-activated receptor-1 but not protease-activated receptor-4 in human platelets. J Thromb Haemost, 9, 1599-1607.

Konopatskaya O, Matthews S, Harper MT, Gilio K, Cosemans JMEM, Williams CM, Navarro N, Carter DA, Heemskerk JWM, Leitges M,  Cantrell D, Poole AW. Protein kinase C mediates platelet secretion and thrombus formation through protein kinase D2. Blood, 118, 416-424.

Heemskerk JW, Harper MT, Cosemans JMEM, Poole AW. Unravelling the different functions of protein kinase C isoforms in platelets. FEBS Lett, 585, 1711-1716.



Harper MT, Poole AW. PKCa enhances sodium-calcium exchange during store-operated calcium entry in mouse platelets. Cell Calcium, 48, 333-340.

Gilio K, Harper MT, Cosemanns JMEM, Konopatskaya O,  Munnix ICA, Molkentin JD, Prinzen L, Leigtes M, Heemskerk JWM, Poole AW. Functional divergence of PKC isoforms in thrombus formation on collagen. J Biol Chem, 285, 23410-23419.

Harper MT, Poole AW. PKCq negatively regulates store-independent Ca2+ entry and phosphatidylserine exposure downstream of GPVI in platelets. J Biol Chem, 285, 19865-19873.

Harper MT, Sage SO. Src family tyrosine kinases activate thrombin-induced non-capacitative calcium entry in human platelets. Platelets, 21, 445-450.

Harper MT, Mason MJ, Sage SO, Harper AGS. Phorbol ester-evoked Ca2+ signalling in human platelets is via autocrine activation of P2X1 receptors, not a novel non-capacitative Ca2+ entry. J Thromb Haemost, 8, 1604-1613.

Jones ML, Harper MT, Williams CM, Aitkin EW, Poole AW. RGD-ligand mimetic antagonists of integrin αIIbβ3 paradoxically enhance GPVI-induced human platelet activation. J Thromb Haemost, 8, 567-576.

Harper MT, MacCarthy-Murrogh L, Jones ML, Konopatskaya O, Poole AW. Platelets: their role in atherogenesis and thrombosis in coronary heart disease. (In Atherosclerosis: Molecular and Cellular Mechanisms, ed. George SJ, Johnson J).

Harper MT, Poole AW. Diverse functions of protein kinase C isoforms in platelet activation and thrombus formation. J Thromb Haemost, 8, 454-462.



Steele BM, Harper MT, Macaulay IC, Foy M, Habas R, Poole AW, Fitzgerald DJ, Maguire PB. Canonical Wnt signalling negatively regulates human platelet function. Proc Nat’l Acad Sci USA, 106, 19836-19841.

Harper MT, Poole AW. PKCq in platelet activation. Blood, 114, 489-491.

Konopatskaya O, Gilio K, Harper MT, Cosemans J, Karim Z, Whiteheart S, Molkentin J, Verkade P, Heemskerk J, Poole AW.  PKCα regulates granule secretion and thrombus formation in mice.  J Clin Invest, 119, 399-407.



Hunter RW, Harper MT, Hers I. The PKB inhibitor Akti-1,2 potentiates PAR-1-mediated platelet function independently of its ability to block PKB. J Thromb Haemost, 6, 1923-1932.

Hall KJ, Gilio K, Harper MT, Cosemans J, Heemskerk JW, Poole AW. Genetic evidence that PKCq negatively regulates platelet activation by GPVI.  PLoS One, 3, e3277.

Harper MT, Sage SO. Is calcium-independent phospholipase A2 required for store-operated calcium entry in human platelets? J Thromb Haemost, 6, 1819-1821.



Harper MT, Poole AW. Isoform-specific functions of PKC: the platelet paradigm. Biochem Soc Trans, 35, 1005-1008.

Redondo PC, Harper AGS, Harper MT, Brownlow SL, Rosado JA, Sage SO. TRPC1-associated α-actinin, and not TRPC1 itself, is tyrosine phosphorylated during human platelet activation. J Thromb Haemost, 5, 2476-2483.



Harper MT, Sage SO. PAR-1 –dependent pp60src activation is dependent on protein kinase C and increased [Ca2+]i: evidence that pp60src does not regulate PAR-1 –dependent Ca2+ entry in human platelets. J Thromb Haemost, 4, 2695-2703.

Harper MT, Sage SO. The actin cytoskeleton regulates calcium entry activated by PAR-4 but not PAR-1 in human platelets. Platelets, 17 134-142.

Redondo PC, Harper MT, Rosado JA, Sage SO. A role for cofilin in the activation of store-operated calcium entry by de novo conformational coupling in human platelets. Blood, 107, 973-979.



Rosado JA, Lopez JJ, Harper AGS, Harper MT, Redondo PC, Pariente JA, Sage SO, Salido GM. Two pathways for store-mediated calcium entry differentially dependent on the actin cytoskeleton in human platelets. J Biol Chem, 278 29231-29235.

Brownlow SL, Harper AGS, Harper MT, Sage SO. A role for hTRPC1 and lipid raft domains in store-mediated calcium entry in human platelets. Cell Calcium, 35 107-113.