People

Professor Stephen Fry

Location: Rutherford 105A

Telephone: 50 5320

Website: http://homepages.ed.ac.uk/sfry

Group members: Mrs Janice Miller, Dr Sandra C. Sharples, Dr Lenka Frankova, Mr Amjad Iqbal, Mr David Messenger, Mr Kyle E. Mohler, Ms Babul Airianah binti Othman

C.V.

Year	Description
1995-present	University of Edinburgh, Professor of Plant Biochemistry
1989	University of Edinburgh, Reader
1983	Joined University of Edinburgh , Lecturer
1982-1983	Senior Research Associate, Department of Chemistry, University of Colorado
1982	Royal Society Rosenheim Research Fellow, Institute of Food Science, Eidgenössische Technische Hochschule, Zürich
1979–1982	Royal Society Rosenheim Research Fellow, Department of Biochemistry, University of Cambridge
1978–1979	SRC Research Fellow, Department of Biochemistry, University of Cambridge
1978	PhD in Plant Physiology, Department of Botany, University of Leicester
1975	BSc in Biological Sciences, University of Leicester

Research groupings

No details available

Teaching

Director of Studies
1997--.

Honours Programme Organiser
Plant Science 4, 2005--

1st year teaching

Quantification in the Life Sciences 1 (~300 students annually)
I give 4 lectures in this course. I cover some of the various roles of light, both as a source of energy (driving photosynthesis) and as a source of information e.g. about the time of day and the time of the year. The aim of the lectures is to emphasise the relevance of bringing a quantitative and biophysical approach to bear on important biological topics.

2nd year teaching

Plants, Fungi and Symbiosis 2 (60-80 students annually)
I currently provide in 5 lectures giving an introduction to the composition of diverse plant cell walls; the biosynthesis and role of lignin; mechanisms and significance of the enzymic digestion of cell walls; defence mechanisms against digestion; and the role of oligosaccharides as elicitors of phytoalexin synthesis. I also run associated practical classes.

3rd year teaching

Evolution & Ecology of Plants 3 (~35-50 students annually)
I contribute 5 lectures and 10 hours’ associated practical work. I cover the acclimations and adaptations of plants to various biotic and abiotic stresses such as drought, flooding, heat, cold, herbivores, competing plants, etc.

4th year teaching

Plant Science 4 Programme (6-17 students annually) Programme Organiser.

The Growth of Plants (PS4)
In 11 sessions I cover the structure & function of plant cell walls [biochemistry of the glycoprotein extensin, inter-polymeric cross-links, oligosaccharins]; roles and metabolism of ascorbate and borate in plants; plant intermediary metabolism; and methodology in plant biochemistry.

Tutorials in Plant Science (PS4)
Tutorial designed to help students tackle data-handling problems.

Fundamentals of Plant Biology (PS4)
Three lectures on plant anatomy, phytochemistry, and culture media/molarities etc. Six hours of associated laboratory work to provide practical experience of the methodology.

Plant Science Synoptic Exam (PS4)
Lecture to provide advice on tackling this type of ‘long essay’ exam.

Plant Science Hons Project
I routinely offer three laboratory projects annually.
I also offer all PS4 students (15 this year) individual advice and practice for the oral presentations of their Projects.

MSc in Plant Taxonomy
I run MSc laboratory projects in this course.

Research interests

The work of the The Edinburgh Cell Wall Group (ECWG)

1. Plant cell wall loosening: cell expansion, fruit softening and wall digestion

The ECWG is investigating the biochemical basis of the biophysical changes occurring in the cell wall.

Discovery and characterisation of xyloglucan endotransglucosylase activity (XET).
Investigating the relationship between XET activity with wall loosening.
Demonstration of the action of XET in vivo by use of multi-isotope (¹³C/²H/³H) pulse-chase protocols
Localisation of sites of XET action in vivo by use of a fluorescence-based assay [see image of growing part of a root stained with a fluorescent reagent to reveal the siting of XET action].
Non-enzymic scission of wall polysaccharides, and the role of hydroxyl radicals. Chemical characterisation of the damage inflicted on plant polysaccharides by OH.
Plant cell walls as a source of atmospheric methane.
Fermentation and subsequent fate of dietary fibre in omnivores and in leaf litter by feeding of radiolabelled plant cell walls to rats.

2. Plant cell wall tightening: assembly and cross-linking of the primary cell wall

Cell-wall matrix polymers, synthesised in the Golgi system, become integrated into the wall by cross-links, which we are working to identify. Cross-linking may also ‘tighten’ existing wall material, and thus restrain cell expansion. The work focuses on covalent cross-links and the ‘tethering’ role of hydrogen-bonded hemicelluloses.

Role of XET in wall assembly/tightening. Molecular ‘grafting’ of newly secreted xyloglucans into the existing wall architecture. ¹³C/³H-Labelling.
MXE, a novel polysaccharide-grafting enzyme from Equisetum. New transglycosylases that act on wall polysaccharides other than xyloglucan, e.g. mixed-linkage glucan (MLG). The newly discovered enzyme activity, MLG : xyloglucan endotransglucosylase (MXE), apparently restricted to the isolated genus, Equisetum.
Oxidative coupling of tyrosine in wall glycoproteins. Cross-linking of cell wall glycoproteins by isodityrosine, di-isodityrosine and related coupling products.
Ferulic acid and related cinnamate derivatives. Intraprotoplasmic versus extraprotoplasmic coupling of ferulate. Formation of ferulate trimers and higher oligomers. Role of quinone methides in formation of ferulate-polysaccharide ether bonds.
Possible cross-links involving galacturonic acid. Uronoyl amines and uronoyl esters.
Xyloglucan-pectin and xyloglucan-MLG bonds. Nature of this inter-polymeric bonding.

3. Oligosaccharins (biologically active oligosaccharides)

Certain molecular fragments, formed by digestion of wall polymers, are ‘oligosaccharins’: they have physiological effects and might serve ‘hormone-like’ roles in vivo. Synthesis, action, transport and turnover of oligosaccharins.

4. Ascorbate (vitamin C) metabolism in plants

This topic emerged from our observation that wall polysaccharides can be oxidatively cleaved by OH and that this radical is readily produced under physiological conditions by non-enzymic reactions of apoplastic solutes (ascorbate, O₂, traces of Cu²⁺).

5. Intermediary metabolism associated with wall polysaccharide biosynthesis

Metabolism of the sugar nucleotides (NDP-sugars) involved in wall polysaccharide synthesis. Novel dual-radiolabelling (³H/¹⁴C) protocol to determine, for each of several pairs of potentially competing pathways, which one predominates in vivo.

6. Evolution of the primary cell wall in early-diverging land plants

Tracing the interesting changes in primary cell wall composition that have occurred during the evolution of land plants. For instance, (a) the ‘invention’ of xyloglucan accompanied the evolution of bryophytes, the first land plants, from charophytes; (b) Anthoceros, in a very isolated taxonomic group, has polysaccharides uniquely rich in ?-D-glucuronosyl-(1?3)-L-galactose; (c) lycopodiophytes (earliest-diverging vascular plants) share a unique wall feature -- abundant 3-O-methyl-D-galactose residues; (d) walls of charophytes, bryophytes and homosporous lycopodiophytes are rich in 3-O-methylrhamnose, which is undetectable in other land plants; (e) the eusporangiate?leptosporangiate transition was accompanied by a marked increase in mannan content. The discovery of MLG (substrate of MXE) in Equisetum. MLG as a possible template for the poorly understood process of bio-silicification in plants. The identity of endogenous [unknown] 'hemicelluloses' in charophytes, which are not known to possess either MLG or xyloglucan.

7. Method development and nomenclature

Development of simple, effective methods for analysing the structure and metabolism of primary cell walls. In-vivo isotopic labelling, enzymic digestion of labelled walls, paper chromatography [see above image of a paper chromatogram of cell-wall monosaccharides, spelling 'ECWG' for Edinburgh Cell Wall Group], high-voltage electrophoresis and HPLC. Development of membrane-impermeant probes to test for the in-vivo production of apoplastic OH radicals. Development of a kit for the semi-quantitative, high-throughput assay of XET activity.

International committee which developed the current nomenclature for xyloglucan oligosaccharides.

In conclusion, the laboratory's work since 1979 has revealed that plant cell walls are a rich source of hitherto unexpected enzymic activities, biochemical reactions and novel organic structures, and that they exhibit interesting evolutionary trends. The work has provided a battery of simple, effective methods for the assay of these reactions and structures; many of these methods are now in routine use in laboratories worldwide.

For more details, see http://homepages.ed.ac.uk/sfry

Representative publications

Original articles

D.J. Messenger, A.R. McLeod, S.C. Fry (2008). The role of UV radiation, photosensitisers, reactive oxygen species and ester groups in mechanisms of methane formation from pectin. Plant, Cell & Environment (in press).

A.R. McLeod, S.C. Fry, G.J. Loake, D.J. Messenger, D.S. Reay, K.A. Smith, B-W. Yun (2008) Ultraviolet radiation drives methane emissions from terrestrial plant pectins. New Phytologist 180: 124-132.

S.J. Bartholdson, A.R. Brown, B.R. Mewburn, D.J. Clarke, S.C. Fry, D.J. Campopiano, J.R.W. Govan (2008) Plant host and sugar alcohol induced exopolysaccharide biosynthesis in the Burkholderia cepacia complex. Microbiology SGM 154: 2513-2521.

S.C. Fry, K.E. Mohler, B.H.W.A. Nesselrode, L. Franková (2008) Mixed-linkage ?-glucan : xyloglucan endotransglucosylase, a novel wall-remodelling enzyme from Equisetum (horsetails) and charophytic algae. Plant Journal 55: 240-252.

S.C. Fry, B.H.W.A. Nesselrode, J.G. Miller, B.R. Mewburn (2008) Mixed-linkage (1?3,1?4)-?-D-glucan is a major hemicellulose of Equisetum (horsetail) cell walls. New Phytologist 179: 104-115.

V. Genovesi, S. Fornalé, S.C. Fry, K. Ruel, P. Ferrer, A. Encina, F.-M. Sonbol, J. Bosch, P. Puigdomènech, J. Rigau, D. Caparrós-Ruiz (2008) ZmXTH1, a new xyloglucan endotransglucosylase/hydrolase in maize, affects cell wall structure and composition. Journal of Experimental Botany 59, 875-889.

T. Takeda, J.G. Miller, S.C. Fry (2008) Anionic derivatives of xyloglucan function as acceptor but not donor substrates for xyloglucan endotransglucosylase activity. Planta, 227, 893-905.

Z.A. Popper, S.C. Fry (2008) Xyloglucan-pectin linkages are formed intra-protoplasmically, contribute to wall-assembly, and remain stable in the cell wall. Planta, 227: 781-794. [Faculty of 1000 Biology: evaluations for Popper ZA & Fry SC Planta 2007 Nov 7: http://www.f1000biology.com/article/id/1094940/evaluation; impact factor 6; ‘must read’]

S.E. Lindsay, S.C. Fry (2008) Control of diferulate formation in dicotyledonous and gramineous cell-suspension cultures. Planta, 227, 439-452

S.C. Sharples, S.C. Fry (2007) Radio-isotope ratios discriminate between competing pathways of cell wall polysaccharide and RNA biosynthesis in living plant cells. Plant Journal, 52, 252-262.

M. Chantarangsee, W. Tanthanuch, T. Fujimura, S.C. Fry, J. Ketudat-Cairns (2007). Molecular characterization of ?-galactosidases from germinating rice (Oryza sativa). Plant Science, 173, 118-134.

P. Sotiriou, S.C. Fry, C.G. Spyropoulos (2007). Protoplast isolation and culture from carob (Ceratonia siliqua L.) hypocotyls: ability of regenerated protoplasts to produce mannose-containing polysaccharides. Physiologia Plantarum, 130, 11-22.

J.G. Miller , V. Farkas , S.C. Sharples, S.C. Fry (2007) O-Oligosaccharidyl-1-amino-1-deoxyalditols as intermediates for fluorescent labelling of oligosaccharides. Carbohydrate Research 342, 44-54.

L.I. Ortega, S.C. Fry, and E. Taleisnik (2006) Why are Chloris gayana leaves shorter in salt-affected plants? Analyses in the elongation zone. Journal of Experimental Botany 57, 3945-3952.

A.R. McLeod, K.K.Newsham, S.C. Fry (2006) Elevated UV-B radiation modifies the extractability of carbohydrates from leaf litter of Quercus robur. Soil Biology and Biochemistry 39, 116-126.

A. Kärkönen, S.C. Fry (2006b) Effect of ascorbate and its oxidation-products on H₂O₂ production in cell-suspension cultures of Picea abies and in the absence of cells. Journal of Experimental Botany. 57, 1633-1644.

A. Kärkönen, S.C. Fry (2006a) Novel characteristics of UDP-glucose dehydrogenase activities in maize: non-involvement of alcohol dehydrogenases in cell wall polysaccharide biosynthesis. Planta 223, 858-870.

M. Lenucci, G. Piro, J.G. Miller, G. Dalessandro, S.C. Fry (2005) Do polyamines contribute to plant cell wall assembly by forming amide bonds with pectins? Phytochemistry 66, 2581-2594.

N. O’Looney, S.C. Fry (2005) Oxaziclomefone, a new herbicide, inhibits wall expansion in maize cell-cultures without affecting polysaccharide biosynthesis, xyloglucan transglycosylation, peroxidase action or apoplastic ascorbate oxidation. Annals of Botany 96, 1097-1107.

A. Encina, S.C. Fry (2005) Oxidative coupling of a feruloyl-arabinoxylan trisaccharide (FAXX) in the walls of living maize cells requires endogenous hydrogen peroxide and is controlled by a low-M_r apoplastic inhibitor. Planta 223, 77-89.

A. Kärkönen, A. Murigneux, J.-P. Martinant, E. Pepey, C. Tatout, B.J. Dudley, S.C. Fry (2005) UDP-Glucose dehydrogenases of maize -- a role in cell wall pentose biosynthesis. Biochemical Journal 391, 409-415.

N. O’Looney, S.C. Fry (2005) The novel herbicide oxaziclomefone inhibits cell expansion in maize cell cultures without affecting turgor pressure or wall acidification. New Phytologist 168, 323-329.

Z.A. Popper, S.C. Fry (2005) Widespread occurrence of a covalent linkage between xyloglucan and acidic polysaccharides in suspension-cultured angiosperm cells. Annals of Botany 96, 91-99. [Faculty of 1000 Biology: evaluations for Popper ZA & Fry SC Ann Bot (Lond) 2005 Jul 96 (1) :91-9 http://www.f1000biology.com/article/id/1025414/ evaluation; impact factor 3]

Y.J. Wu, B.R. Jeong, S.C. Fry, J.S. Boyer (2005) Change in XET activities, cell wall extensibility and hypocotyl elongation of soybean seedlings at low water potential. Planta 220, 593-601.

N. O’Looney, S.C. Fry (2005) A simple apparatus for measuring long-term extension of plant cell walls subjected to tensile stress. Plant Biosystems 139: 102-106.

K. Vissenberg, S.C. Fry, M. Pauly, H. Höfte, J.-P. Verbelen (2005) XTH acts at the microfibril-matrix interface during cell elongation. Journal of Experimental Botany 56, 673-683.

M.A. Green, S.C. Fry (2005) Vitamin C degradation in plant cells via enzymatic hydrolysis of 4-O-oxalyl-L-threonate. Nature 433: 83-88. [Faculty of 1000 Biology: evaluations for Green MA & Fry SC Nature 2005 Jan 6 433 (7021) :83-7 http://www. f1000biology.com/article/id/1023719/evaluation; impact factor 3]

J.G. Miller, S.C. Fry (2004) N-[³H]Benzoylglycylglycylglycine as a probe for hydroxyl radicals. Analytical Biochemistry 335, 126-134.

M. Albert, M. Werner, P. Proksch, S.C. Fry, R. Kaldenhoff (2004) The cell wall-modifying xyloglucan endotransglycosylase/hydrolase LeXTH1 is expressed during the defence reaction of tomato against the plant parasite Cuscuta reflexa. Plant Biology 6, 402-408.

Z.A. Popper, S.C. Fry (2004) Primary cell wall composition of pteridophytes and spermatophytes. New Phytologist 164, 165-174.

T. Takeda and S.C. Fry (2004) Control of xyloglucan endotransglucosylase activity by salts and anionic polymers. Planta 219, 722-732.

E.M. Kerr, S.C. Fry (2004) Extracellular cross-linking of xylan and xyloglucan in maize cell-suspension cultures: the role of oxidative phenolic coupling. Planta 219, 73-83.

Z.A. Popper, I.H. Sadler, S.C. Fry (2004) 3-O-Methylrhamnose in lower land plant primary cell walls. Biochemical Systematics and Ecology 32, 279-289.

K. Vissenberg, V. van Sandt, S.C. Fry, J.-P. Verbelen (2003) Xyloglucan endotransglucosylase action is high in the root elongation zone and in the trichoblasts of all vascular plants from Selaginella to Zea mays. Journal of Experimental Botany 54, 335-344.

Z.A. Popper, I.H. Sadler, S.C. Fry (2003) ?-D-Glucuronosyl-(1?3)-L-galactose, an unusual disaccharide from polysaccharides of the hornwort Anthoceros caucasicus. Phytochemistry 64, 325-335.

Z.A. Popper, S.C. Fry (2003) Primary cell wall composition of bryophytes and charophytes. Annals of Botany 91, 1-12.

E.M. Kerr, S.C. Fry (2003) Pre-formed xyloglucans and xylans increase in molecular weight in three distinct compartments of a maize cell-suspension culture. Planta 217, 327-339.

J.C. Dumville, S.C. Fry (2003) Solubilisation of tomato fruit pectins by ascorbate: a possible non-enzymic mechanism of fruit softening. Planta 217, 951-961.

J.C. Dumville, S.C. Fry (2003) Gentiobiose: a novel oligosaccharin in ripening tomato fruit. Planta 216, 484-495.

P. Perrone, C.M. Hewage, A.R. Thomson, K. Bailey, I.H. Sadler, S.C. Fry (2002) Patterns of methyl and O-acetyl esterification in spinach pectins: new complexity. Phytochemistry 60, 67-77.

S.C. Fry, J.G. Miller, J.C. Dumville (2002) A proposed role for copper ions in cell wall loosening. Plant and Soil 247, 57-67.

S.L. Gardner, M.M. Burrell, S.C. Fry (2002) Screening of Arabidopsis thaliana stems for variation in cell wall polysaccharides. Phytochemistry 60, 241-254.

K. Vissenberg, J.-P. Verbelen, S.C. Fry (2001) Root hair initiation is coupled to a highly localized increase of xyloglucan endotransglycosylase action in Arabidopsis roots. Plant Physiology 127, 1125-1135.

J.E. Thompson, S.C. Fry (2001). Restructuring of wall-bound xyloglucan by transglycosylation in living plant cells. Plant Journal 26, 23-34.

J.E. Thompson, S.C. Fry (2001). Density-labelling of cell wall polysaccharides in cultured rose cells: comparison of incorporation of ²H and ¹³C from exogenous glucose. Carbohydrate Research 332, 175-182.

G. Tabbì, S.C. Fry, R.P. Bonomo (2001). ESR study of the non-enzymic scission of xyloglucan by an ascorbate-H₂O₂-copper system: the involvement of the hydroxyl radical and the degradation of ascorbate. Journal of Inorganic Biochemistry 84, 179-187.

Z.A. Popper, I.H. Sadler, S.C. Fry (2001). 3-O-Methyl-D-galactose residues in lycophyte primary cell walls. Phytochemistry 57, 711-719.

J.G. Miller, S.C. Fry (2001). Characteristics of xyloglucan after attack by hydroxyl radicals. Carbohydrate Research 332, 389-403.

S.C. Fry, J.C. Dumville, J.G. Miller (2001). Fingerprinting of polysaccharides attacked by hydroxyl radicals in vitro and in the cell walls of ripening pear fruit. Biochemical Journal 357, 729-735.

R. Abia, S.C. Fry (2001) Degradation and metabolism of ¹⁴C-labelled proanthocyanidins from carob pods (Ceratonia siliqua) in the gastrointestinal tract of the rat. Journal of the Science of Food and Agriculture 81, 1156-1165.

K. Vissenberg, I.M. Martinez-Vilchez, J.-P. Verbelen, J.G. Miller, S.C. Fry (2000). In-vivo colocalization of xyloglucan endotransglycosylase activity and its donor substrate in the elongation zone of Arabidopsis roots. Plant Cell 12, 1229-1238.

J.E. Thompson, S.C. Fry (2000). Evidence for covalent linkage between xyloglucan and acidic pectins in suspension-cultured rose cells. Planta 211, 275-286.

W. Suginta, P.A.W. Robertson, B. Austin, S.C. Fry, L.A. Fothergill-Gilmore (2000). Chitinases from Vibrio: activity screening and purification of chiA from Vibrio carchariae Journal of Applied Microbiology 89, 76-84.

N.M. Steele, Z. Sulová, P. Campbell, J. Braam, V. Farkas, and S.C. Fry. (2000). Ten isoenzymes of xyloglucan endotransglycosylase from plant cell walls select and cleave the donor substrate stochastically. Biochemical Journal 355, 671-679.

N.M. Steele, S.C. Fry (2000). Differences in catalytic properties between native isoenzymes of xyloglucan endotransglycosylase (XET). Phytochemistry 54, 667-680.

B.C. McGorum, S.C. Fry, G. Wallace, K. Coenen, J. Robb, G. Williamson, O.I. Aruoma (2000). Properties of herbage in relation to equine dysautonomia: biochemical composition and antioxidant and pro-oxidant actions. Journal of Agricultural and Food Chemistry 48, 2346-2352.

S.C. Fry, S.C. Willis, A.E.J. Paterson (2000). Intraprotoplasmic and wall-localised formation of arabinoxylan-bound diferulates and larger ferulate coupling-products in maize cell-suspension cultures. Planta 211, 679-692.

J.C. Dumville, S.C. Fry (2000). Uronic acid-containing oligosaccharins: Their biosynthesis, degradation and signalling roles in non-diseased plant tissues. Plant Physiology and Biochemistry 38, 125-140.

G. Wallace, S.C. Fry (1999). Action of diverse peroxidases and laccases on six cell wall-related phenolic compounds. Phytochemistry 52, 769-773.

T.A. Truelsen, J. Laird, S.C. Fry (1999). Xyloglucan metabolising enzyme activities in tobacco cells. Journal of Plant Physiology 154: 95-101.

N.M. Steele, S.C. Fry (1999). Purification of xyloglucan endotransglycosylases (XETs): a generally applicable and simple method based on reversible formation of an enzyme-substrate complex. Biochemical Journal 340, 207-211.

C.K. Smith, C.M. Hewage, S.C. Fry, I.H. Sadler (1999). ?-D-Mannopyranosyl-(1?4)-?-D-glucuronopyranosyl-(1?2)-myo-inositol, a new and unusual oligosaccharide from cultured rose cells. Phytochemistry 52, 387-396.

C.K. Smith, S.C. Fry (1999). Biosynthetic origin and longevity in vivo of ?-D-mannopyranosyl-(1?4)-?-D-glucuronopyranosyl-(1?2)-myo-inositol, an unusual extracellular oligosaccharide produced by cultured rose cells. Planta 210, 150-156.

P.P.M. Iannetta, S.C. Fry (1999). Visualisation of the activity of xyloglucan endotransglycosylase (XET) isoenzymes after gel electrophoresis. Phytochemical Analysis 10, 238-240.

Review articles and contributions to the popular press

S.C. Fry (2009) Cell wall polysaccharide composition and covalent cross-linking. In ‘Plant Polysaccharides’ edited by P. Ulvskov (Blackwell) (in press).

S.C. Fry (2008) Plant cell wall biosynthesis. In Handbook of Plant Science, ed by K. Roberts; J. Wiley (in press).

S.C. Fry (2008) Plant cell walls. In Handbook of Plant Science, ed by K. Roberts; J. Wiley (in press).

S.E. Lindsay, S.C. Fry (2007) Redox and wall-restructuring. In ‘The Expanding Cell’ edited by J-P. Verbelen & K. Vissenberg, Springer, Berlin [ISBN 10 3-540-39114-2]. Pp 159-190.

S.C. Fry (2006) Wall-to-wall biochemistry: a personal perspective. In ‘The Science and Lore of the Plant Cell Wall: Biosynthesis, Structure and Function’ edited by T. Hayashi, BrownWalker Press, Boca Raton, Florida [ISBN 1-58112-445-7]. Pp. 171-180.

S.C. Fry (2005). Hanging on (The Last Word). New Scientist, 23 July 2005.

F. Cernone, G. Dalessandro, S.C. Fry (2005) Special Issue, Tenth Cell Wall Meeting, Sorrento, Italy: Introduction. Plant Biosystems 139: 1.

M.A. Green, S.C. Fry (2005) Apoplastic degradation of ascorbate: novel enzymes and metabolites permeating the plant cell wall. Plant Biosystems 139: 2-7.

R.A.M. Vreeburg, S.C. Fry (2005) Reactive oxygen species in cell walls. In: ‘Antioxidants and Reactive Oxygen Species in Plants’, Blackwell, Oxford (pp. 215-249). Edited by N. Smirnoff. ISBN-13: 978-1-4051-2529-1

S.C. Fry (2004) Oxidative coupling of tyrosine and ferulic acid residues: intra- and extra-protoplasmic occurrence, predominance of trimers and larger products, and possible role in inter-polymeric cross-linking. Phytochemistry Reviews 3, 97-111.

S.C. Fry (2004) Tansley Review: Primary cell wall metabolism: tracking the careers of wall polymers in living plant cells. New Phytologist 161, 641-675.

S.C. Fry (2003). Citric secret (The Last Word). New Scientist, July 2003.

J.K.C. Rose, J. Braam, S.C. Fry, K. Nishitani (2002) The XTH family of enzymes involved in xyloglucan endotransglucosylation and endohydrolysis: current perspectives and a new unifying nomenclature. Plant and Cell Physiology, 43, 1421-1435.

S.C. Fry (2003). Ripening. In Encyclopedia of Applied Plant Sciences, Academic Press, 794-807.

S.C. Fry (2003). Cell walls. In Encyclopedia of Applied Plant Sciences, Academic Press, pp 75-87.

S.C. Fry (2003). Cells. In Encyclopedia of Applied Plant Sciences, Academic Press, pp 512-523.

S.C. Fry (2002). Worm count (The Last Word). New Scientist, April 2002.

S.C. Fry (2000). Ménage à trois: oxidative coupling of feruloyl polysaccharides in vivo proceeds beyond dimers. Polyphénols Actualités 19, 8-12.

S.C. Fry (1999). Biosphere (The Last Word). New Scientist, April 1999.

S.C. Fry (Web November 1999; printed 2001) Plant cell wall biosynthesis. In Encyclopedia of Life Sciences, http://www.els.net, DOI: 10.1038/npg.els.0001683. Chichester: Nature Publishing Group. (Also printed edition.)

S.C. Fry (Web September 1999; printed 2001) Plant cell walls. In Encyclopedia of Life Sciences, http://www.els.net, DOI 10.1038/npg.els.0001682. Chichester: Nature Publishing Group. (Also printed edition.)

J.C. Dumville, S.C. Fry (1999) Uronic acid-containing oligosaccharins: their biosynthesis, degradation and signalling roles in non-diseased plant tissues. Plant Physiol. Biochem. 38, 125-140.

S.C. Fry (1999). Oligosaccharins as regulators of plant growth. In P.J.J. Hooykaas, M.A. Hall & K.R. Libbenga (Eds.). Biochemistry and Molecular Biology of Plant Hormones (pp. 247-265). Amsterdam: Elsevier

This article was published on Feb 8, 2012