GAGs and human iPS cells

Pijuan-Galitó S, Tamm C, Schuster J, Sobol M, Forsberg L, Merry CL and Annerén C. Human serum-derived protein removes the need for coating in defined human pluripotent stem cell culture. Nat Commun. 2016; 7: 12170.

GAGs and ES cells

Holley RJ, Smith RA, van de Westerlo EM, Pickford CE, Merry CLR*, van Kuppevelt TH. Use of flow cytometry for characterization and fractionation of cell populations based on their expression of heparan sulfate epitopes. Methods Mol Biol. 2015;1229:239-51 (Review)

Holley RJ, Meade KA, Merry CLR. Using embryonic stem cells to understand how glycosaminoglycans regulate differentiation. Biochem Soc Trans. 2014 42(3):689-95 (Review)

Glycosaminoglycans as regulators of stem cell differentiation. Smith RAA, Meade K, Pickford CE, Holley RJ, Merry CLR. Biochemical Society Transaction. 2011 39(1): 383-87. (review)

Specific Glycosaminoglycans Modulate Neural Specification of Mouse Embryonic Stem Cells. Pickford CE, Holley RJ, Rushton G, Stavridis MP, Ward CM, Merry CLR. Stem Cells. 2011 29(4):629-40.

Influencing hematopoietic differentiation of mouse embryonic stem cells using soluble heparin and heparan sulfate saccharides. Holley RJ, Pickford CE, Rushton G, Lacaud G, Gallagher JT, Kouskoff V, Merry CLR. The Journal of Biological Chemistry. 2011 286(8):6241-52.

A developmentally-regulated heparan sulfate epitope defines a sub-population with increased blood potential during mesodermal differentiation Baldwin RB, ten Dam GB, van Kuppevelt TH, Lacaud G, Gallagher JT, Kouskoff V, Merry CLR Stem Cells. 2008 26(12):3108-18.

Essential alterations of heparan sulfate during the differentiation of embryonic stem cells to Sox1-enhanced green fluorescent protein-expressing neural progenitor cells. Johnson CE, Crawford BE, Stavridis M, Ten Dam G, Wat AL, Rushton G, Ward CM, Wilson V, van Kuppevelt TH, Esko JD, Smith A, Gallagher JT, Merry CLR. Stem Cells. 2007 25(8):1913-23.

E-Cadherin/EMT and ES cells (with Chris Ward)

Hawkins K, Keramari M, Soncin F, Segal JM, Mohamet L, Miazga N, Ritson S, Bobola N, Merry CLR, Ward CM*. Novel cell lines isolated from mES cells exhibiting de novo methylation of the E-cadherin promoter. Stem Cells. 2014 32 (11), 2869-2879.

Hawkins K, Mohamet L, Ritson S, Merry CLR, Ward CM. E-Cadherin and, in Its Absence, N-cadherin Promotes Nanog Expression in Mouse Embryonic Stem Cells via STAT3 Phosphorylation. Stem Cells. 2012; 30(9): 1842-1851.

Soncin F, Mohamet L, Ritson S, Hawkins K, Bobola N, Zeef L, Merry CLR, Ward CM. E-cadherin acts as a regulator of transcripts associated with a wide range of cellular processes in mouse embryonic stem cells. PLoS One. 2011;6(7):e21463.

Abrogation of E-cadherin mediated cell-cell contact in mouse embryonic stem cells results in reversible LIF-independent self-renewal. Soncin F, Eckardt D, Eastham AM, Spencer H, Ritson S, Kemler R, Merry CLR, Ward CM Stem Cells. 2009 27(9):2069-80.

Epithelial mesenchymal transition events during embryonic stem cell differentiation. Merry CLR, Ward CM. Biomarkers Insights. (in press, review)

Epithelial-mesenchymal transition events during human embryonic stem cell differentiation. Eastham AM, Spencer H, Soncin F, Ritson S, Merry CLR, Stern PL, Ward CM. Cancer Res. 2007 67(23):11254-62.

E-cadherin inhibits cell surface localization of the pro-migratory 5T4 oncofetal antigen in mouse embryonic stem cells. Spencer HL, Eastham AM, Merry CLR, Southgate TD, Perez-Campo F, Soncin F, Ritson S, Kemler R, Stern PL, Ward CM. Mol Biol Cell. 2007 18(8):2838-51.

Mucopolysaccharidoses (with Brian Bigger)

Watson HA, Holley RJ, Langford-Smith KJ, Wilkinson FL, van Kuppevelt TH, Wynn RF, Wraith JE, Merry CLR, Bigger BW*. Heparan sulfate inhibits hematopoietic stem and progenitor cell migration and engraftment in mucopolysaccharidosis I. J Biol Chem. 2014 289(52):36194-203

Sergijenko A, Langford-Smith A, Liao AY, Pickford CE, McDermott J, Nowinski G, Langford-Smith KJ, Merry CLR, Jones SA, Wraith JE, Wynn RF, Wilkinson FL, *Bigger BW. Myeloid/microglial driven autologous hematopoietic stem cell gene therapy corrects a neuronopathiclysosomal disease. Mol Ther. 2013 21(10):1938-49


Ashworth JC, Thompson JL, James JR, Slater CE, Pijuan-Galitó S, Lis-Slimak K, Holley RJ, Meade KA, Thompson A, Arkill KP, Tassieri M, Wright AJ, Farnie G, Merry CLR. Peptide gels of fully-defined composition and mechanics for probing cell-cell and cell-matrix interactions in vitro. Matrix Biol. 2019; 85: 15-33.

Wan S, Borland S, Richardson SM, Merry CLR, Saiani A, Gough JE. Self-assembling peptide hydrogel for intervertebral disc tissue engineering. Acta Biomater. 2016; 46: 29-40.

Ayerst BI, Day AJ, Nurcombe V, Cool SM, Merry CLR. New strategies for cartilage regeneration exploiting selected glycosaminoglycans to enhance cell fate determination. Biochem Soc Trans. 2014 42(3):703-9 (Review)

Giangreco A, Merry CLR The biochemical determinants of tissue regeneration. Biochem Soc Trans. 2014 Jun;42(3):607-8. (Review)

Immobilization of Heparan Sulfate on Electrospun Meshes to Support Embryonic Stem Cell Culture and Differentiation. Meade KA, White KJ, Pickford CE, Holley RJ, Marson A, Tillotson D, van Kuppevelt TH, Whittle JD, Day AJ, Merry CLR. The Journal of Biological Chemistry. 2013 288(8):5530-38.

Sugar functionalised PEGA surfaces support metabolically active hepatocytes. Ambury RF. Merry CLR, Ulijn RV. Journal of Materials Chemistry, 2011, in press.

Controlling stiffness in nanostructured hydrogels produced by enzymatic dephosphorylation. Thornton K, Smith AM, Merry CLR, Ulijn RV Biochem Soc Trans. 2009 (Pt 4):660-4 Controlling cell morphology on amino acid-modified cellulose Kalaskar DM, Gough JE, Ulijn RV, Sampson WW, Scurr DJ, Rutten FJ, Alexander MR, Merry CLR, Eichhorn SJ. Soft Matter. 2008 4:1059-65.

Three-dimensional culture of annulus fibrosus cells within PDLLA/Bioglass composite foam scaffolds: assessment of cell attachment, proliferation and extracellular matrix production. Helen W, Merry CLR, Blaker JJ, Gough JE. Biomaterials. 2007 28(11):2010-20.

Mesenchymal stem cell biology (with Cay Kielty)

Holley RJ, Ball SG, Merry CLR, Whetton AH, Kielty CM*, Canfield AE*. Comparative Quantification of the Surfaceome of Human Multipotent Mesenchymal Progenitor Cells. Stem Cell Reports 2015 (accepted for publication)

Baldwin AK, Cain SA, Lennon R, Godwin A, Merry CLR, Kielty CM. Epithelial-mesenchymal status influences how cells deposit fibrillin microfibrils. J Cell Sci. 2014 127(Pt 1):158-71

Ball SG, Worthington JJ, Canfield AE, Merry CLR, Kielty CM. Mesenchymal stromal cells: Inhibiting PDGF receptors or depleting fibronectin induces mesodermal progenitors with endothelial potential.Stem Cells. 2013 32(3):694-705

Banka S, Cain SA, Carim S, Daly SB, Urquhart JE, Erdem G, Harris J, Bottomley M, Donnai D, Kerr B, Kingston H, Superti-Furga A, Unger S, Ennis H, Worthington J, Herrick AL, Merry CLR, Yue WW, Kielty CM, Newman WG. Leri's pleonosteosis, a congenital rheumatic disease, results from microduplication at 8q22.1 encompassing GDF6 and SDC2 and provides insight into systemic sclerosis pathogenesis.Ann Rheum Dis. 2014 Jan 17. doi: 10.1136/annrheumdis-2013-204309.

GAGs in macular degeneration (with Tony Day)

Keenan TD, Pickford CE, Holley RJ, Clark SJ, Lin W, Dowsey AW, Merry CLR, Day AJ, Bishop PN. Age-dependent changes in heparan sulfate in human Bruch's membrane: implications for age-related macular degeneration. Invest Ophthalmol Vis Sci. 2014 55(8):5370-9


The heparanome – The enigma of encoding and decoding heparan sulphate sulfation. Lamanna WC, Kalus I, Padva M, Baldwin R., Merry CLR, Dierks T J. Biotechnol. 2007 129(2) 290-307. (review)

Heparan sulphate 6-O-endosulphatases-discrete in vivo activities and functional cooperativity. Lamanna WC, Baldwin RJ, Padva M, Kalus I, Ten Dam G, van Kuppelet TH, Gallagher JT, von Figura K, Dierks T, Merry CLR Biochem. J. 2006 400(1) 63-73.


The morphogenic properties of oligomeric endostatin are dependent on cell surface heparan sulphate. Clamp A, Blackhall FH, Henrioud A, Jayson GC, Javaherian K, Esko J, Gallagher JT, Merry CLR J. Biol. Chem. 2006 281;21, 14813-14822.

Binding of endostatin to endothelial heparan sulphate shows a differential requirement for specific sulphates Blackhall F, Merry CLR, Lyon M, Jason G, Folkman J, Javaherian K, Gallagher J Biochemical Journal 2003 375 131-139.

General Glycobiology reviews

Ayerst BI, Merry CLR, Day AJ. The good the bad and the ugly of glycosaminoglycans in tissue engineering applications. Pharmaceuticals (Basel). 2017; 10 (2): 54. (review)

Glycans in evolution and development. Merry CLR, Astrautsova SA. EMBO Rep. 2008 9(7):617-22. (review)

A sugar rush for developmental biology. Merry CLR, Ward CM. Development. 2008 135(8):1389-93. (review)

Glycoscience finally comes of age Merry AH, Merry CLR EMBO Rep. 2005 Oct;6(10):900-3. (review)

HS structure/function analysis

A Deligny, T Dierker, A Dagälv, A Lundequist, I Eriksson, AV Nairn, KW Moremen, CLR Merry, L Kjellén. NDST2 (N-deacetylase/N-sulfotransferase-2) enzyme regulates heparan sulfate chain length. J Biol Chem. 2016; 291 (36): 18600-18607.

Borland, SJ; Morris TG; Borland SC; Morgan MR; Francis SE; Merry CLR; Canfield AE. Regulation of vascular smooth muscle cell calcification by syndecan-4/FGF-2/PKCα signaling and cross-talk with TGFβ. Cardiovasc Res. 2017; 113(13):1639-1652.

Wilkinson FL, Holley RJ, Langford-Smith KJ, Badrinath S, Liao A, Langford-Smith A, Cooper JD, Jones SA, Wraith JE, Wynn RF,Merry CL, Bigger BW. Neuropathology in mouse models of mucopolysaccharidosis type I, IIIA and IIIB. PLoS One. 2012; 7(4): e35787.

Langford-Smith A, Wilkinson FL, Langford-Smith KJ, Holley RJ, Sergijenko A, Howe SJ, Bennett WR, Jones SA, Wraith J, Merry CL, Wynn RF, Bigger BW. Hematopoietic Stem Cell and Gene Therapy Corrects Primary Neuropathology and Behavior in Mucopolysaccharidosis IIIA mice. Molecular Therapy. 2012; 20(8): 1610-1621.

Mucopolysaccharidosis type I: Unique structure of accumulated heparan sulfate and increased N-sulfotransferase activity in mice lacking alpha-L-iduronidase. Holley RJ, Deligny A, Wei W, Watson HA, Ninonuevo MR, Dagalv A, Leary JA, Bigger BW, Kjellen L, Merry CLR. Journal of Biological Chemistry 2011 286:43 37515-37524.

Clark SJ, Keenan TD, Fielder HL, Collinson LJ, Holley RJ, Merry CLR, van Kuppevelt TH, Day AJ, Bishop PN. Mapping the differential distribution of glycosaminoglycans in the adult human retina, choroid and sclera. Invest Ophthalmol Vis Sci. 2011 Jul 11. [Epub ahead of print]

Detection of 2-O-sulfated iduronate and N-acetylglucosamine units in heparan sulphate by an antibody selected against acharan sulphate ten Dam G, van de Westerlo EM, Smetesers TF, Willemse M, van Muijen GN, Merry CLR, Gallagher JT, Kim YS, van Kuppevelt TH Journal of Biological Chemistry 2004 279:37 38346-38352.

A new model for the domain structure of heparan sulphate based on the novel specificity of K5 lyase Murphy KJ, Merry CLR, Lyon M, Roberts IS, Thompson JE, Gallagher JT Journal of Biological Chemistry 2004 26 279:26 27239-27245.

Heparan Sulphate 2-O-sulphotransferase (Hs2st) and mouse development Wilson V, Gallagher JT, Merry CLR Glycoconjugate Journal 2003 19 347-354. (review)

Not all perlecans are created equal: interactions with fibroblast growth factor 2 (FGF-2) and FGF receptors Knox S, Merry CLR, Stringer S, Melrose J, Whitelock J Journal of Biological Chemistry 2002 April 26 277:17 14657-14665.

New insights into heparan sulphate biosynthesis from the study of mutant mice Merry CLR, and Gallagher JT Biochem. Soc. Symp. 2002 69 47-57.

Role of heparan sulphate 2-O-sulphotransferase in the mouse Merry CLR, Wilson VA Biochimica et Biophysica Acta 2002 1573 (3) 319-327.

The molecular phenotype of heparan sulphate in the Hs2st-/- mutant mouse Merry CLR, Bullock SL, Swan DC, Backen AC, Lyon M, Beddington RSP, Wilson VA. Gallagher JTG Journal of Biological Chemistry 2001 September 21 276:38 35429-35434.

Heparan sulphate proteoglycans and Cancer. Blackhall FH, Merry CLR, Davies EJ, Jayson GC British Journal of Cancer 2001 85 1094-1098.

Highly-sensitive sequencing of the sulphated domains of heparan sulphate. Merry CLR, Lyon M, Deakin JA, Hopwood JJ, Gallagher JT Journal of Biological Chemistry 1999 June 25 274:26 18455-18462.


  1. Gallagher JTG, Lyon M, Merry CLR. Method of sequencing oligosaccharide(s). WO200001843-A; WO200001843-A1; AU9946354-A.
  2. Merry CLR, Meade K, Lowe E, Saiani A, Miller A. Use of hydrogels for stem cell culture. Filed in USA and Europe September 2014 WO2013124620 A1 (Read).
  3. Multiple partners including Merry CLR, Meade K. New Scaffold for Cardiac Patch. PCT Submission PCT/EP2012/068648 (Feb 2014)

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