Staff Profile
Emeritus Professor Jerry Barnes
Professor of Environmental Biology
- Telephone: +44 (0) 191 208 4850
- Fax: +44 (0) 191 208 5229
- Personal Website: https://www.linkedin.com/in/jerry-barnes-ab74a015/
- Address: School of Natural and Environmental Sciences
Devonshire Building, Devonshire Terrace
Newcastle University
Newcastle Upon Tyne
NE1 7RU
Background
Following 10 years as Royal Society Research Fellow, I was appointed as a Reader and then to a Professorship (2006). I retired from the University in 2019 following 33 years' research/teaching/management at Newcastle University. The over-arching focus of my research, and that of my team, centred on the impacts of photochemical oxidant pollution on crops and native plant communities, with particular focus on the oxidative damage resulting at the cellular level with a view to the directed breeding of crops with enhanced resilience to meet future food demands. Unique facilities constructed and used as the platform to deliver this work included a suite of state-of-the-art indoor computer-controlled environment chambers, an open-field fumigation system at Keenley Fell (Northumberland) and 16 rigid open-top chambers situated at what was the University's Field Station at Close House – the latter constituting two of the UKs principle Air Pollution Research Facilities at the time. My groups’ work, made a significant contribution not only to UK Air Quality Guidelines but also to the setting of Critical Levels for ozone pollution which constitute the backbone for trans-national air pollution abatement strategies across Europe and North America. Since the millenium, my interests in ozone extended to researching the controlled use of the gas to curb microbial spoilage of fresh produce and improve food safety. This area of research activity resulted in significant industry collaboration, several PhD and KTP awards, a wealth of recent publications, plus multiple patent filings.
For the last 20 years of my service at Newcastle University I was engaged part-time. This enabled me to pursue my external business interests. In 2001 I co-founded my first company, Biofresh (www.bio-fresh.com),. This business was established to exploit know how (and subsequent solution-engineering patents) in the field of novel modified storage and preservation atmospheres applicable to high value fruit and vegetables. In 2012 the business, by then trading around the world, was sold to the International Sales Distributor. I retain an interest and shareholding in the resulting product: Biofresh Group Ltd. In 2007. Having spotted a gap in another lucrative market, I stepped back from the management team at Biofresh to co-Found Geneius Laboratories (www.geneiuslabs.com), an award-winning company established to service the UK Food and Drink Sector with pioneering DNA-based testing approaches. The business grew quickly under my leadership to employ 70 full-time staff and became the largest independent testing services provider in the UK. In July 2017, the company was acquired by SYNLAB, a multi-national laboratory services provider. Prior to exit, I stepped-down as CEO at Geneius to focus on co- Founding and growing APIX Nutrition (now owned by APIX Bioscience NV) This spin-out was established to commercialise specialist livestock feed solutions aimed and has attracted significant external investment to position the company's product range for launch to market. I am presently focussing on new interests in the renewable energy sector in Scotland (see: www.kintyrewind.com).
On my journey, I have established strong/long-lasting research collaboration with overseas colleagues and gained much strategic/project/staff management experience, supervised >45 fully-funded PhD programmes to successful completion, been successful in winning research grants worth >£3 million from a range of competitive sources, authored >130 refereed publications (to date) delivering a personal Google Scholar h-index of 50 based on >7K citations. Google Scholar Profile, won numerous business awards and attracted >£20m capital investment
Area of Expertise
Plant-Environment interactions, Air pollution, Food testing & microbiology, Postharvest storage systems, Bee nutrition
Qualifications
- 2001 Certificate Teaching and Learning in Higher Education, Newcastle University
- 1985 PhD in Plant Physiology, UCNW Bangor
- 1981 BSc (Hons) Agricultural Botany, UCNW Bangor
Appointments
Academia
- 2006 - present Professor of Environmental Biology, Newcastle University
- 2001 - 2006 Reader in Plant-Atmosphere Exchange, Newcastle University
- 1992 - 2001 Royal Society Research Fellow, Newcastle University
- 1986 - 1992 DEFRA Research Fellow, Newcastle University
External
- 2001 - 2006 co-Founder and CEO Biofresh Ltd (www.bio-fresh.com) [now Biofresh Group Ltd.]
- 2001 - 2010 Director and Company Secretary, biofresh Ltd
- 2008 - 2017 co-Founder and Managing Director Geneius Laboratories Ltd. (www.geneiuslabs.com
- 2016 - present co-Founder and CEO, APIX Nutrition Ltd.
- 2019 - present co-Founder and Director, KIntyre Wind Group (www.kintyrewind.com)
- 2021 - present Formal Advisor to the Board APIX Bioscience NV
Accolades and Awards
Impact
- Winner, Dti/Innovate UK SMART Awards [2004 and 2013]
- Winner, Medilink most-promising NE company [2009]
- Runner-up, Medilink UK Competition for most-promising company [2010]
- Sunday Times Business Entrepeneurs 'Tips For Success in 2013'
- Runner-up, Bionow Business Awards [2013]
- Winner, Society for Food Hygiene and Technology 'Best New Business' [2013]
- Winner, Northumberland Business Award - Innovation [2014]
- Winner, Northumberland Business Award - Large Business of the Year [2016]
Academic
- Invited participant in Royal Society-organised Ministerial Briefings 1996-2001
- Invited participant in Royal Society REF/White Paper/Information Consultation exercises 1997-2000
- Winner, Royal Society East/West and Japan exchange Funding 1997-2000
- Invited UK representative French Academy of Sciences Meeting – Emerging National Talent
- Invited participant at three Rank Prize Funds Symposia/Dinners
- Winner, Swales Foundation Award [1994]
- Winner, Sir David Moore Memorial Prize - Awarded by The Arkleton Trust [2007]
- Invited speaker/chair at ≈ 75 National & International conferences
- Invited to become a Fellow of the Royal Society of Biology [2015]
Membership of External Committees
- International Advisory Board, EU-Horizon 2020 Strength2Food project 2017 - 2022
- External Consultant/Assessor for REF2020 Edinburgh, Napier Univ. [UoAs 7, 25 and 26] 2016-2020
- Honorary Life-time Member, International Society of Environmental Botanists
- Society for Experimental Biology Plant Science Committee: 2000-2005
- Royal Society Alumni and Environmental Advisory Network: 2001-present
- International Ozone Association AgriFood Panel 2005-2010
- Member, NERC Committee for Air Pollution Effects Research (CAPER) 2005-present
- Member DEFRA & Royal Society Expert Working Groups: Photochemical Oxidants 2005-2010
- International Advisory Panel to Polish Academy Sciences: Krakow 2002-2007. FP5-funded Project
- Member, Commonwealth Fellowships/Scholarships Assessment Panel 2008-2011
- Treasurer 1995-2000 then Co-Chair, APPM International Symposia Series 2000-2005 – specifically governing the organization of the 5th (Pulawy, Poland) and 6th (Tsukuba, Japan) meetings.
- Co-Chair and Organiser, European Science Foundation Meeting – Morpeth (2001)
- Expert Evaluator, Italian Ministry for University Research 1998-2005
Membership of Internal University Committees
- Member, University Ethics Committee 2016 - present
- Director of Research, School of Biology 2001-2005 and Member, Faculty Research Committee
- Executive Management Committee, School of Biology 2002-2005
- Executive Management Committee, Institute for Research on the Environment and Sustainability 2004-2009
- Chair, Wolfson bid (with Vice Chancellors support) to create gene-environment centre, 2004
- Deputy-Academic Director, Devonshire Build Committee 2001-2005
- Director of Postgraduate Studies/Admissions/Chair Postgraduate Comm, School of Biology 1998-2003
- Head, Environmental and Molecular Plant Physiology (one of 3 major Research themes in the School of Biology at the time) 1996 – 2015
- Member, Faculty Graduate School Management Board 2000-2003 (standing-in for the postgraduate sub-Dean at University Bursary Committee Meetings)
- Member, Faculty Business Opportunities Working Group and Regional Links Board 2001-2003
- Member, Faculty Integrated PhD Programme Working Group 2000-2003
- Member, Faculty EU-Marie Curie Working Group 2000-2003
- Member, Arabic Students Working Group – International Office 2002-2005
Oral Presentations/Thesis Examinations
I have delivered around 100 talks/seminars in: Australia x6 Canada x4 Channel Islands x4 Germany x7 Greece x16 Italy x14 Ireland x3 Japan x11 Netherlands x7 Portugal x2 Spain x12 Trinidad x1 USA x8 UK c.40, and examined PhD/MSc/MPhil Theses presented in the UK x15 Spain x4 Australia x2 Greece x1 and Canada x1
Research Interests
- Atmospheric pollution, especially ozone, and impacts on vegetation at physiological, biochemical and molecular level
- Cellular reaction to (ozone-induced) oxidative stress and cellular detoxification systems
- Redox regulation and signal propagation/transduction
- Postharvest storage technologies and waste prevention approaches
- Food/Drink safety/microbiology
Other Expertise
- Commercial applications for gaseous/aqueous ozone in food and drink sector
- Preservation of fresh produce/foodstuffs
- Decontamination of fresh produce/storage facilities
- Postharvest biology and storage technologies
Current Work
- Establishment and modeling of ozone flux-response relationships
- Examining impacts of rising ozone on upland vegetation
- Exploring the deployment of ozone for the preservation and decontamination of fresh produce/foodstuffs
- QTLs governing ozone tolerance and characterization of NILs
- Plant-pollinator interactions
- Internalization of foodborne pathogens
Membership
Presently
- Member, NERC/DEFRA Committee for Air Pollution Impacts Research
- ad-hoc Member, DEFRA ozone modelling group
- Member, Royal Society Environmental Advisory Network
Formerly
- Co-Chairman, Air Pollution and Global Change Conference Series (organiser 7th Meeting: Edinburgh, Sept 2007)
- Chair and co-ordinator Marie Curie Early Stage (PhD) Training Site [PLENTI]Plant-Environment Interactions
- Member, International Advisory Board Polish Academy of Sciences FP5 'Crop Stress' Project
- Member, International Ozone Association's World Agri-Food Committee
- Member, IRES Executive Management Board
Funding
Attracted: c. £5 million in research funding from a variety of sources (including EU, UK-based Research Councils and Industrial Partners) plus #3.5m business development funding (venture capital, business angels, private investors) past 20 years
Published: 1 co-edited book (plus 1 'in press') and >120 research papers in high-ranking international journals
Supervised: >35 PhD projects; 1 MPhil; 1 Dipl >20 postdoctoral researchers
Current team: 4 PhD students; 1 KTP Associate
Patents
patents:
GB **** [with Unilever, withdrawn]
GB **** [with Uniq Foods] granted
GB/US/Europe **** [BioFresh] PCT filings
European Trademarks:
BioFresh and 'Safer ways to Fresher Foods' [Europe, N. America, Japan, SE Asia]
Geneius, Geneius rapID, Geneius microID, Geneius microST [Europe, N. America, Japan, SE Asia]
Undergraduate Teaching
- Biology in Action [Stage 1] [contributor]
- Plant Biology [Stage 1] [contributor]
- Pollution of Air, Water and Soil [Stage 2] [module leader]
- Pollution of Air, Water and Soil [Stage 3] [module leader]
- Final Year Dissertation Project students
- Tutees: 8-10 per annum
Postgraduate Teaching
- 4 PhD Students
- MSc Industrial and Commercial Biotechnology. Business Skills.
Business/Professional Skills Development/Mentoring
- Newcastle University Careers Service
- North East Process and Innovation Centre
- Royal Society of Edinburgh Enterprise Programme
- Goumenaki E, Gonzalez-Fernandez I, Barnes JD. Ozone uptake at night more damaging to plants than equivalent day-time flux. Planta 2021, 253(3), 75-86.
- Abraham PE, Hurtado Castano N, Cowan-Turner D, Barnes J, Poudel S, Hettich R, Flutsch S, Santelia D, Borland AM. Peeling back the layers of crassulacean acid metabolism: functional differentiation between Kalanchoë fedtschenkoi epidermis and mesophyll proteomes. Plant Journal 2020, 103(2), 869-888.
- Barnes J, Kaminska A, Gillespie C, Fernandez I, Czyezylo-Mysza H, Tyrka M, Rancic D, Pecinar H, Dodig D, Quarrie S. QTLs governing the impacts of ozone on wheat yield. New Phytologist 2019. In Preparation.
- Stabler D, Power E, Borland A, Barnes J, Wright G. A method for analysing small samples of floral pollen for free and protein-bound amino acids. Methods in Ecology and Evolution 2018, 9(2), 430-438.
- Power E, Stabler D, Borland AM, Barnes J, Wright GA. Analysis of nectar from low-volume flowers: a comparison of collection methods for free amino acids . Methods in Ecology and Evolution 2018, 9(3), 734–743.
- Lewis E, Hudson JA, Cooke N, Haynes E, Barnes J. Comparison of next generation sequencing techniques for the identification of human pathogens associated with fresh produce. Food Microbiology 2018. Submitted.
- Wyness K, Peacock S, Goumenaki E, Barnes J. Long-term exposure to simulated future ozone climate reduces productivity and alters plant species dynamics in an upland mesotrophic grassland community of high conservation value. Grass and Forage Science 2018. In Preparation.
- Toet S, Oliver V, Ineson P, McLoughlin S, Helgason T, Peacock S, Stott A, Barnes J, Ashmore M. How does elevated ozone reduce methane emissions from peatlands?. Science of the Total Environment 2017, 579, 60–71.
- Hayes F, Mills G, Jones L, Abbott J, Ashmore M, Barnes J, Cape JN, Coyle M, Peacock S, Rintul N, Toet S, Wedlich K, Wyness K. Consistent ozone-induced decreases in pasture forage quality across several grassland types and consequences for UK lamb production. Science of the Total Environment 2016, 543(Part A), 336-346.
- Wani S, Barnes J, Singleton I. Investigation of potential reasons for bacterial survival on 'ready-to-eat' leafy produce during exposure to gaseous ozone. Postharvest Biology and Technology 2016, 111, 185-190.
- Wani S, Maker J, Thompson J, Barnes J, Singleton I. Effect of ozone treatment on inactivation of Escherichia coli and Listeria sp. on spinach. Agriculture 2015, 5(2), 155-169.
- Gillespie C, Stabler D, Tallentire E, Goumenaki E, Barnes J. Exposure to environmentally-relevant levels of ozone negatively effect pollen and fruit development. Environmental Pollution 2015, 206, 494-501.
- Tzortzakis N, Taybi T, Antony E, Singleton I, Borland A, Barnes J. Profiling shifts in protein complement in tomato fruit induced by atmospheric ozone-enrichment and/or wound-inoculation with Botrytis cinerea. Postharvest Biology and Technology 2013, 78, 67-75.
- Haider MS, Barnes JD, Cushman JC, Borland AM. A CAM- and starch-deficient mutant of the facultative CAM species Mesembryanthemum crystallinum reconciles sink demands by repartitioning carbon during acclimation to salinity. Journal of Experimental Botany 2012, 63(5), 1985-1996.
- Wedlich KV, Rintoul N, Peacock S, Cape JN, Coyle M, Toet S, Barnes JD, Ashmore M. Effects of ozone on species composition in an upland grassland. Oecologia 2012, 168(4), 1137-1146.
- Dodig D, Zoric M, Kobiljski B, Savic J, Kandic V, Quarrie S, Barnes J. Genetic and Association Mapping Study of Wheat Agronomic Traits Under Contrasting Water Regimes. International Journal of Molecular Sciences 2012, 13(5), 6167-6188.
- Zoric M, Dodig D, Kobiljski B, Quarrie S, Barnes J. Population structure in a wheat core collection and genomic loci associated with yield under contrasting environments. Genetica 2012, 140(4-6), 259-275.
- Wyness K, Mills G, Jones L, Barnes JD, Jones DL. Enhanced nitrogen deposition exacerbates the negative effect of increasing background ozone in Dactylis glomerata, but not Ranunculus acris. Environmental Pollution 2011, 159(10), 2493-2499.
- Tzortzakis N, Taybi T, Roberts R, Singleton I, Borland A, Barnes J. Low-level atmospheric ozone exposure induces protection against Botrytis cinerea with down-regulation of ethylene-, jasmonate- and pathogenesis-related genes in tomato fruit. Postharvest Biology and Technology 2011, 61(2-3), 152-159.
- Tzortzakis N, Taybi T, Roberts R, Singleton I, Borland A, Barnes J. Low-level atmospheric ozone exposure induces protection against Botrytis cinerea with down-regulation of ethylene-, jasmonate- and pathogenesis-related genes in tomato fruit. Postharvest Biology and Technology 2011, 61(2-3), 152-159.
- Gonzalez-Fernandez I, Kaminska A, Dodmani M, Goumenaki E, Quarrie S, Barnes JD. Establishing ozone flux–response relationships for winter wheat: Analysis of uncertainties based on data for UK and Polish genotypes. Atmospheric Environment 2010, 44(5), 621-630.
- Tzortzakis N, Singleton I, Barnes JD. Impact of low-level atmospheric ozone-enrichment on black spot and anthracnose rot of tomato fruit . Postharvest Biology and Technology 2008, 47(1), 1-9.
- Gonzalez I, Bass D, Muntifering R, Mills G, Barnes JD. Impacts of ozone pollution on productivity and forage quality of grass/clover swards. Atmospheric Environment 2008, 42(38), 8755-8769.
- Tzortzakis N, Singleton I, Barnes J. Deployment of low-level ozone-enrichment for the preservation of chilled fresh produce. Postharvest Biology and Technology 2007, 43(2), 261-270.
- Goumenaki E, Fernandez IG, Papanikolaou A, Papadopoulou D, Askianakis C, Kouvarakis G, Barnes J. Derivation of ozone flux-yield relationships for lettuce: A key horticultural crop. Environmental Pollution 2007, 146(3), 699-706.
- Tzortzakis N, Borland A, Singleton I, Barnes J. Impact of atmospheric ozone-enrichment on quality-related attributes of tomato fruit. Postharvest Biology and Technology 2007, 45(3), 317-325.
- Hassenberg K, Idler C, Molloy E, Geyer M, Plochl M, Barnes J. Use of ozone in a lettuce-washing process: An industrial trial. Journal of the Science of Food and Agriculture 2007, 87(5), 914-919.
- Borland A, Elliott S, Patterson S, Taybi T, Cushman J, Pater B, Barnes J. Are the metabolic components of crassulacean acid metabolism up-regulated in response to an increase in oxidative burden?. In: Meeting on Phenotypic Plasticity and the Changing Environment held at the Society for Experimental Biology Plant Frontiers Meeting. 2006, Sheffield: Journal of Experimental Botany, Oxford University Press.
- Pignocchi C, Kiddle G, Hernandez I, Foster SJ, Asensi A, Taybi T, Barnes J, Foyer CH. Ascorbate oxidase-dependent changes in the redox state of the apoplast modulate gene transcript accumulation leading to modified hormone signaling and orchestration of defense processes in tobacco. Plant Physiology 2006, 141(2), 423-435.
- Quarrie SA, Pekic Quarrie S, Radosevic R, Rancic D, Kaminska A, Barnes JD, Leverington M, Ceoloni C, Dodig D. Dissecting a wheat QTL for yield present in a range of environments: From the QTL to candidate genes. Journal of Experimental Botany 2006, 57(11), 2627-2637.
- O'Mahony MM, Dobson ADW, Barnes JD, Singleton I. The use of ozone in the remediation of polycyclic aromatic hydrocarbon contaminated soil. Chemosphere 2006, 63(2), 307-314.
- Kaminska A, Fernadez IG, Dodmani M, Barnes J. Modelling ozone impacts on winter wheat and establishing critical fluxes for ozone risk assessment. In: Acta Physiologiae Plantarum. 2005, Springer.
- Cardoso-Vilhena J, Balaguer L, Eamus D, Barnes JD. Mechanisms underlying the amelioration of O3‐induced damage by elevated atmospheric concentrations of CO2. Journal of Experimental Botany 2004, 55(397), 771-781.
- Kaminska A, Gillespie C, Dodmani M, Bilsborrow P, Quarrie S, Barnes J. QTLs governing ozone impacts on wheat yield. In: Acta Physiologiae Plantarum. 2004, Springer.
- Sanmartin M, Drogoudi PD, Lyons T, Pateraki I, Barnes J, Kanellis AK. Over-expression of ascorbate oxidase in the apoplast of transgenic tobacco results in altered ascorbate and glutathione redox states and increased sensitivity to ozone. Planta 2003, 216(6), 918-928.
- Pignocchi C, Fletcher JM, Wilkinson JE, Barnes JD, Foyer CH. The function of ascorbate oxidase in tobacco. Plant Physiology 2003, 132(3), 1631-1641.
- Molloy E, Hassenberg K, Plochl M, Idler C, Geyer M, Barnes JD. Use of ozonated washing water for the surface decontamination of perishable produce. Acta Horticulturae 2003, 599, 577-582.
- Davison AW, Barnes JD. Air pollution-abiotic stress interactions. In: Bell, J.N.B., Treshow, M, ed. Air Pollution and Plant Life. London: Wiley-Blackwell, 2002, pp.359-377.
- Maddison J, Lyons T, Plochl M, Barnes J. Hydroponically cultivated radish fed L-galactono-1,4-lactone exhibit increased tolerance to ozone. Planta 2002, 214(3), 383-391.
- Zheng Y, Shimizu H, Barnes JD. Limitations to CO2 assimilation in ozone-exposed leaves of Plantago major. New Phytologist 2002, 155(1), 67-78.
- Barnes JD, Zheng Y, Lyons TM. Plant resistance to ozone: the role of ascorbate. In: Omasa, K., Saji, H., Yousefian, S., Kondo, N, ed. Air Pollution and Plant Biotechnology: Prospects for Phytomonitoring and Phytoremediation. Tokyo: Springer-Verlag, 2002, pp.235-252.
- Jondrup PM, Barnes JD, Port GR. The effect of ozone fumigation and different Brassica rapa lines on the feeding behaviour of Pieris brassicae larvae. Entomologia Experimentalis et Applicata 2002, 104(1), 143-151.
- Cardoso-Vilhena J, Barnes J. Does nitrogen supply affect the response of wheat (Triticum aestivum cv. Hanno) to the combination of elevated CO2 and O3?. Journal of Experimental Botany 2001, 52(362), 1901-1911.
- Zheng Y, Lyons T, Ollerenshaw JH, Barnes JD. Ascorbate in the leaf apoplast is a factor mediating ozone resistance in Plantago major. Plant Physiology and Biochemistry 2000, 38(5), 403-411.
- Turcsanyi E, Lyons T, Plochl M, Barnes J. Does ascorbate in the mesophyll cell walls form the first line of defence against ozone? Testing the concept using broad bean (Vicia faba L.). Journal of Experimental Botany 2000, 51(346), 901-910.
- Zheng Y, Lyons T, Barnes J. Effects of ozone on the production and utilization of assimilates in Plantago major. Environmental and Experimental Botany 2000, 43(2), 171-180.
- Turcsanyi E, Cardoso-Vilhena J, Maddision J, Gillespie C, Balaguer L, Ollerenshaw J, Barnes JD. Impacts of tropospheric ozone: past, present and likely future. In: Singh SN, ed. Trace Gas Emissions and Plants. Dordrecht: Kluwer Academic, 2000, pp.249-272.
- Soda C, Bussotti F, Grossoni P, Barnes J, Mori B, Tani C. Impacts of urban levels of ozone on Pinus halepensis foliage. Environmental and Experimental Botany 2000, 44(1), 69-82.
- Soja G, Barnes JD, Posch M, Vandermeiren K, Pleijel H, Mills G. Phenological weighting of ozone exposures in the calculation of critical levels for wheat, bean and plantain. Environmental Pollution 2000, 109(3), 517-524.
- Plochl M, Lyons T, Ollerenshaw J, Barnes J. Simulating ozone detoxification in the leaf apoplast through the direct reaction with ascorbate. Planta 2000, 210(3), 454-467.
- Gimeno BS, Bermejo V, Reinert RA, Zheng Y, Barnes JD. Adverse effects of ambient ozone on watermelon yield and physiology at a rural site in Eastern Spain. New Phytologist 1999, 144(2), 245-260.
- Lyons T, Ollerenshaw JH, Barnes JD. Impacts of ozone on Plantago major: Apoplastic and symplastic antioxidant status. New Phytologist 1999, 141(2), 253-263.
- Ollerenshaw JH, Lyons T, Barnes JD. Impacts of ozone on the growth and yield of field-grown winter oilseed rape. Environmental Pollution 1999, 104(1), 53-59.
- Balaguer L, Manrique E, De Los Rios A, Ascaso C, Palmqvist K, Fordham M, Barnes JD. Long-term responses of the green-algal lichen Parmelia caperata to natural CO2 enrichment. Oecologia 1999, 119(2), 166-174.
- Barnes J, Bender J, Lyons T, Borland A. Natural and man-made selection for air pollution resistance. Journal of Experimental Botany 1999, 50(338), 1423-1435.