ANNUAL REPORT OF THE

EPIDEMIOLOGY RESEARCH

GROUP

CENTRE for INFECTIOUS DISEASES

UNIVERSITY of EDINBURGH

October 2007

1

EPIGROUP: 2006-07

The Epidemiology Research Group (Epigroup) moved to the Ashworth Laboratories on the

University of Edinburgh King’s Buildings campus in October 2005. In our new location we

are continuing our research on the epidemiology and pathogenesis of animal and human

infectious diseases. This research involves the close integration of field studies, laboratory

experiments and theoretical analyses and draws on collaborations with colleagues from a

variety of disciplines working in the University of Edinburgh, other institutions in Scotland,

elsewhere in the UK and internationally. We study infectious disease systems ranging from

TSEs to virus, bacteria, protozoa and helminths infections. The common theme is the

development of a formal, quantitative understanding of the dynamics of parasites and

pathogens within their host populations with particular emphasis on informing the design of

disease control programmes.

A number of major research projects got under way in the last year. The IDEAL project,

which is looking at the impact of infectious diseases on calves in Kenya, has progressed

through a set-up, training and piloting phase to the point where data collection will shortly

begin in earnest. This is a substantial 5 year project supported by a Wellcome Trust

programme grant and involving partners at the International Livestock Research Institute

(ILRI) in Nairobi and the University of Pretoria. EPIC, the Scottish Government funded

Centre of Excellence in veterinary epidemiology, has also seen the first year of its activities.

EPIC is a 4 year programme with several partners across Scotland and with a broad remit to

develop and apply quantitative tools to the epidemiology of livestock diseases in Scotland. As

this report is being prepared Nick Savill, Paul Bessell and Victoriya Volkova, working

through EPIC, are very actively involved in advising the Scottish Government regarding risks

associated with the ongoing foot-and-mouth disease outbreak in Surrey. Last year was also a

peak of activity for our DEFRA/SFC Veterinary Training and Research Initiative programme

(a joint programme with the University of Glasgow) with all three research fellows in place,

our MSc programme running and an enthusiastic cohort of summer students.

Reflecting these new activities, the year has seen several further appointments to Epigroup.

Magai Kaare and Olga Tosas Auguet were appointed as post-doctoral researchers working on

the IDEAL project. They will mainly be working in Kenya. Victoriya Volkova joined us as

the Edinburgh Research Fellow of EPIC. Xu-Sheng Zhang was appointed to a postdoctoral

research post to work on modelling the dynamics of E. coli O157. Eric Fèvre moved to a

Research Fellowship funded through the VTRI award. We were also joined by three new

Research Council funded PhD students – Kate Mitchell, Katie Atkins and Elly Gaunt – and by

a VTRI funded research MSc student – Emily Courcier. Kate will be working on the

immunoepidemiology of schistosomes, Katie on Marek’s disease in chickens (partly

supported by Pfizer), Elly on emerging viruses, and Emily on risk factors for E. coli O26

infection in cattle.

We are delighted that two University of Edinburgh doctoral degrees were awarded in 2007 for

work undertaken in Epigroup: Suzanne St Rose was awarded a PhD for her work on

susceptibility to TSEs; and Natalie Nicholls was awarded a PhD for her work on sheep

scrapie and productivity.

Epigroup members have been authors on 15 papers in peer-reviewed journals on topics

ranging from susceptibility to malaria to horse-racing fatalities, and have also contributed

chapters to books on rabies and models for cell biology. One highlight was work led by

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Margo Chase-Topping further characterising the nature of E. coli O157 “supershedder”

infections (published in the Journal of Clinical Microbiology). Past and current Epigroup

members also published a major study of rabies vaccination to protect Ethiopian wolves (a

highly endangered species) in Nature.

We have also been very successful in raising research funds. Nick Savill was awarded a

project grant by the Wellcome Trust to work on within-host models of malaria, and Nick and

Mark Woolhouse are among the PIs on a multi-disciplinary Scottish Research Development

Grant to work on influenza. Together with some smaller awards this brings the total value of

the research activity that Epigroup is involved with to over £10M.

We now have six students working towards postgraduate degrees. We continue to contribute

to undergraduate teaching, especially through a Special Topic on epidemiology for the

Microbiology & Infection undergraduate Honours course but also through contributions to

Medical Microbiology and a taught MSc on Drug Discovery and Translational Biology. In

addition, the VTRI Fellows are running a very successful epidemiology discussion group

which draws participants from several Edinburgh locations.

Epigroup members have presented their work at numerous venues, including Mark

Woolhouse’s presentation to the President’s Forum of the American Association for

Microbiology meeting in Toronto. Our activities have, as usual, generated regular media

interest with mentions in the Scottish and UK press (Times, Financial Times) and interviews

on Radio 5 Live, Sky TV and BBC News 24. Contributions to public understanding of

science included Mark Woolhouse speaking at a Café Scientifique event. We have also

contributed to advisory meetings held by several agencies, including DEFRA, the Department

of Health and the Chief Scientist’s office.

Contact details

Secretary: Kath Tracey

Address: Centre for Infectious Diseases, University of Edinburgh, Ashworth

Laboratories, Kings Buildings, West Mains Road, Edinburgh EH9 3JF, UK.

Telephone: +(44) (0)131 650 5456

Fax: +(44) (0)131 650 6564

E-mail: kath.tracey@ed.ac.uk

Web site: www.vqe.vet.ed.ac.uk/epigroup

Appendices

1. List of Epigroup members.

2. List of publications in peer reviewed scientific journals.

3. List of other publications.

4. List of research grants.

5. List of postgraduate degrees awarded.

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APPENDIX 1: List of 2006-2007 Epigroup members

Academic staff

Professor Mark Woolhouse, Chair of Infectious Disease Epidemiology

Dr Nick Savill, Lecturer in Mathematical Epidemiology

Dr Mark Bronsvoort, Lecturer in Veterinary Epidemiology (Easter Bush Veterinary Centre)

Dr Lisa Boden, VTRI Research Fellow

Dr Eric Fèvre, VTRI Research Fellow

Ian Handel, VTRI Research Fellow

Dr Olga Tosas Auguet, Post-Doctoral Researcher

Dr Margo Chase-Topping, Post-Doctoral Researcher

Dr Richard Howey, Post-Doctoral Researcher

Dr Magai Kaare, Post-Doctoral Researcher

Dr Victoriya Volkova, Post-Doctoral Research Fellow

Dr Xu-Sheng Zhang, Post-Doctoral Researcher

Postgraduate students

Katie Atkins, PhD Research Student

Paul Bessell, PhD Research Student

Elly Gaunt, PhD Research Student

Kate Mitchell, PhD Research Student

Natalie Nicholls, PhD Research Student

Suzanne St Rose, PhD Research Student

Emily Courcier, MSc Research Student

Summer students:

Stacey Douglas, VTRI scholar

Samantha Edwards, VTRI scholar

Bronwyn Koterwas, VTRI scholar

Fiona Land, VTRI scholar

Cynthia Metcalf, VTRI scholar

Jenna Richardson, VTRI scholar

Katie Surgine, VTRI scholar

Support staff

Kath Tracey, Secretary

Stuart Lansley, Computing Support

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APPENDIX 2: List of refereed publications

1. Boden, L.A., Anderson, G.A, Charles, J.A., Morgan, K.L., Morton, J.M, Parkin,

T.D.H., Slocombe R.F. and Clarke, A.F. (2006). Risk of fatality and causes of death of

Thoroughbred horses associated with racing in Victoria between 1989 and 2004.

Equine Veterinary Journal 38: 312-318.

2. Boden, L.A., Anderson, G.A., Charles, J.A., Morgan, K.L., Morton, J.M., Parkin

T.D.H., Clarke, A.F. and Slocombe, R.F. (2007). Risk factors for Thoroughbred

racehorse fatality in jump starts in Victoria, Australia (1989-2004). Equine Veterinary

Journal 39: 422-428.

3. Boden, L.A., Anderson, G.A., Charles, J.A., Morgan, K.L., Morton, J.M., Parkin

T.D.H., Clarke, A.F. and Slocombe, R.F. (2007). Risk factors for Thoroughbred

racehorse fatality in flat starts in Victoria, Australia (1989-2004). Equine Veterinary

Journal 39: 430-437.

4. Bronsvoort, B.M.d.C., Anderson, J., Corteyn, A., Hamblin, P., Kitching, R.P., Nfon, C.,

Tanya, V.N. and Morgan, K.L. (2006). Geographical and age-stratified distributions of

foot-and-mouth disease virus-seropositive and probang-positive cattle herds in the

Adamawa province of Cameroon. Veterinary Record 159: 299-308.

5. Bronsvoort, B. M. d. C., A. Renz, et al. (2005). Repeated high doses of avermectins

cause prolonged sterilisation, but do not kill, Onchoceca ochengi adult worms in

African cattle. Filaria Journal 4: 8.

6. Chase-Topping, M.E., Gunn, G., Strachan, W.D., Edwards, S.A., Smith, W.J.,

Hillman, K., Stefopoulou, S.N. and Thomson, J.R. (2007). Epidemiology of porcine

non-specific colitis on Scottish farms. The Veterinary Journal 173: 353-360.

7. Chase-Topping, M.E., Kruuk, L.E., Lajous, D., Touzeau, S., Matthews, L., Simm, G.,

Foster, J.D., Rupp, R., Eychenne, F., Hunter, N., Elsen, J-M. and Woolhouse, M.E.

(2005). Genotype-level variation in lifetime breeding success, litter size and survival

of sheep in scrapie-affected flocks. Journal of General Virology 86: 1229-1238.

8. Chase-Topping, M.E., McKendrick, I.J., Pearce, M.C., MacDonald, P., Matthews, L.,

Halliday, J., Allison, L., Fenlon, D., Low, J.C., Gunn, G. and Woolhouse, M.E.J.

(2007). Risk factors for the presence of high-level shedders of Escherichia coli O157

on Scottish farms. Journal of Clinical Microbiology, 45: 1594-1603.

9. Clark, L., Clutton, R.E., Blissitt, K.J. and Chase-Topping, M.E. (2005). Effects of

peri-operative morphine administration during halothane anaesthesia in horses.

Veterinary Anaesthesia and Analgesia 32: 10-15.

10. Clark, L., Clutton, R.E., Blissitt, K.J. and Chase-Topping, M.E. (2007). The effects of

morphine on the recovery of horses from halothane anaesthesia. Veterinary

Anaesthesia and Analgesia, in press.

11. Egger, J.R., Ooi, E.E., Kelly, D.W., Woolhouse, M.E., Davies, C.R. and Coleman, P.G.

(in press). Reconstructing historical changes in the force of infection of dengue fever in

Singapore: implications for surveillance and control. Bulletin of the World Health

Organization.

12. Fèvre, E.M., Bronsvoort, B.M.d.C., Hamilton, K.A. and Cleaveland, S. (2006).

Animal movements and the spread of infectious diseases. Trends in Microbiology 14:

125-131.

13. Fèvre, E.M., Picozzi, K., Jannin, J., Welburn, S.C. and Maudlin, I. (2006). Human

African trypanosomiasis: epidemiology and control. Advances in Parasitology 61:

167-221.

5

14. Fèvre, E.M., Tilley, A., Picozzi, K., Fyfe, J., Anderson, I., Magona, J.W., Shaw, D.J.,

Eisler, M.C. and Welburn, S.C. (2006). Central point sampling from cattle in livestock

markets in areas of human sleeping sickness. Acta Tropica 97: 229-232.

15. Fèvre, E.M., Picozzi, K., Fyfe, J., Waiswa, C., Odiit, M., Coleman, P.G. and Welburn,

S.C. (2005). A burgeoning epidemic of sleeping sickness in Uganda. Lancet 366: 745-

747.

16. Hahn, C.N., Handel, I.G., Bronsvoort, B.M. de C., Green, S.L. and Mayhew, I.G. (in

press). Assessment of the utility of using intra- and inter-vertebral minimum sagittal

diameter ratios in the diagnosis of cervical vertebral malformation in horses.

Veterinary Radiology & Ultrasound.

17. Halliday, J.E.B., Chase-Topping, M.E., Pearce, M.C., McKendrick, I.J., Allison, L.,

Fenlon, D., Low, C., Mellor, D.J., Gunn, G.J. and Woolhouse, M.E.J. (2006). Herd-

level risk factors associated with the presence of Phage type 21/28 E. coli O157 on

Scottish cattle farms. BMC Microbiology, 6: 99.

18. Haydon, D.T., Randall, D.A., Matthews, L., Knobel, D.L., Tallents, L.A., Gravenor,

M.B., Williams, S.D., Pollinger, J.P., Cleaveland, S., Woolhouse, M.E.J., Sillero-

Zubiri, C., Marino, J., Macdonald, D.W. and Laurenson, M.K. (2007). Low coverage

vaccination strategies for the conservation of endangered species. Nature 443: 692-

695.

19. Hoyle, D.V., Davison, H.C., Knight, H.I., Yates, C.M., Dobay, O., Gunn, G.J., Amyes,

S.G.B. and Woolhouse, M.E.J. (2006). Molecular characterisation of bovine faecal

Escherichia coli shows persistence of defined ampicillin resistant strains and the

presence of class 1 integrons on an organic beef farm. Veterinary Microbiology 115:

250-257.

20. Hoyle, D.V., Yates, C.M., Chase-Topping, M.E., Turner, E.J., Davies, S.E., Low, J.C.,

Gunn, G.J., Woolhouse, M.E. and Amyes, S.G. (2005). Molecular epidemiology of

antimicrobial-resistant commensal Escherichia coli strains in a cohort of newborn

calves. Applied and Environmental Microbiology 71: 6680-6688.

21. Hoyle, D.V., Davison, H.C., Knight, H.I., Yates, C.M., Dobay, O., Gunn, G.J., Amyes,

S.G.B. and Woolhouse, M.E.J. (2006). Molecular characterisation of bovine faecal

Escherichia coli shows persistence of defined ampicillin resistant strains and the

presence of class 1 integrons on an organic beef farm. Veterinary Microbiology 115:

250-257.

22. Kaare, M.T., Picozzi, K., Fèvre, E.M., Cleaveland, S., Mtambo, M.M., Mellau, L.S.,

Mlengeya, T. and Welburn, S.C. (2007). Sleeping sickness – a re-emerging disease in

the Serengeti? Travel Medicine and Infectious Diseases 5: 117-124.

23. Kosmider, R., Kelly, L., Laurenson, K., Coleman, P., Fooks, A.R., Woolhouse, M. and

Wooldridge, M. (2006). Risk assesments to inform policy decisions regarding

importation of pets from North America. Veterinary Record 158: 694-695.

24. Liu, W.-C., Matthews, L., Chase-Topping, M., Savill, N.J., Shaw, D.J. and Woolhouse,

M.E.J. (2007). Metapopulation dynamics of Escherichia coli O157 in cattle: an

exploratory model. Journal of the Royal Society Interface 4: 917-924.

25. Liu, W.-C., Shaw, D.J., Matthews, L., Hoyle, D.V., Pearce, M.C., Yates, C.M., Low,

J.C., Amyes, S.G.B., Gunn, G.J. and Woolhouse, M.E.J. (in press). Modelling the

epidemiology and transmission of verocytotoxin-producing Escherichia coli

serogroups O26 and O103 in two different calf cohorts. Epidemiology and Infection.

26. Makita, K., Fèvre, E.M., Waiswa, C., Kaboyo, W., Bronsvoort, B.M.d.C., Eisler, M.C.

and Welburn, S.C. (2007). Human brucellosis in urban and peri-urban areas of

Kampala, Uganda. Annals of the New York Academy of Science, in press.

6

27. Matthews, L., Low, J.C., Gally, D.L., Pearce, M.C., Mellor, D.J., Heesterbeek, J.A.P.,

Chase-Topping, M., Naylor, S.W., Shaw, D.J., Reid, S.W.J., Gunn, G.J. and

Woolhouse, M.E.J. (2006). Heterogeneous shedding of Escherichia coli O157 in cattle

and its implications for control. Proceedings of the National Academy of Sciences, USA

103: 547-552.

28. Matthews, L., McKendrick, I.J., Ternent, H., Gunn, G.J., Synge, B. and Woolhouse,

M.E.J. (2006). Super-shedding cattle and the transmission dynamics of Escherichia coli

O157. Epidemiology and Infection 134: 131-142.

29. Nicholls, N., Kruuk, L., Woolhouse, M., Stevenson, E., Gravenor, M. and Baylis, M.

(2006). Investigation of farmer regard for scrapie-susceptible sheep. Veterinary Record

158: 732-734.

30. Odiit, M., Bessell, P.R., Fèvre, E.M., Robinson, T., Kinoti, J., Coleman, P.G.,

Welburn, S.C., McDermott, J. and Woolhouse, M.E.J. (2006). Using remote sensing

and geographic information systems to identify villages at high risk for rhodesiense

sleeping sickness in Uganda. Transactions of the Royal Society of Tropical Medicine

and Hygiene 100: 354-362.

31. Pearce, M.C., Evans, J., McKendrick, I.J., Smith, A.W., Knight, H.I., Mellor, D.J.,

Woolhouse, M.E.J., Gunn, G.J. and Low, J.C. (2006). Prevalence and virulence factors

for E. coli serogroups O26, O103, O111 and O145 shed by cattle in Scotland. Applied

and Environmental Microbiology 72: 653-659.

32. Rowe, J.A., Handel, I.G., Thera, M.A., Deans, A-M., Lyke, K.E., Koné, A., Diallo,

D.A., Raza, A., Kai, O., Marsh, K., Plowe, C.V., Doumbo, O.K. and Moulds, J.M. (in

press). Blood group O protects against severe Plasmodium falciparum malaria.

Proceedings of the National Academy of Sciences, USA.

33. Savill, N.J., Shaw, D.J., Deardon, R., Tildesley, M.J., Keeling, M.J., Woolhouse,

M.E.J., Brooks, S.P. and Grenfell, B.T. (2006). Topographic determinants of foot and

mouth disease transmission in the UK 2001 epidemic. BMC Veterinary Research 2: 3.

34. Savill, N.J., Shaw, D.J., Deardon, R., Tildesley, M.J., Keeling, M.J., Woolhouse,

M.E.J., Brooks, S.P. and Grenfell, B.T. (2007). Effect of data quality on estimates of

farm infectiousness trends in the UK 2001 foot-and-mouth disease epidemic. Journal

Royal Society Interface 4: 235-241.

35. Savill, N.J., St Rose, S.G., Keeling, M.J. and Woolhouse, M.E.J. (2006). Silent spread

of H5N1 in vaccinated poultry. Nature 442: 757.

36. St Rose, S.G., Hunter, N., Foster, J.D., Drummond, D., McKenzie, C., Parnham, D.,

Will, R.G., Woolhouse, M.E.J. and Rhind, S.M. (2007). Quantification of Peyer’s

patches in Cheviot sheep for future scrapie pathogenesis studies. Veterinary

Immunology and Immunopathology 116: 163-171.

37. St. Rose, S.G., Hunter, N., Matthews, L., Foster, J.D., Chase-Topping, M.E., Kruuk,

L.E.B., Shaw, D.J., Rhind, S.M., Will, R.G. and Woolhouse, M.E.J. (2006).

Comparative evidence for a link between Peyer’s patch development and

susceptibility to transmissible spongiform encephalopathies. BMC Infectious Diseases

6: 5-12.

38. Tildesley, M.J., Savill, N.J., Shaw, D.J., Deardon, R., Brooks, S.P., Woolhouse,

M.E.J., Grenfell, B.T. and Keeling, M.J. (2006). Optimal reactive vaccination

strategies for a foot and mouth outbreak in the UK. Nature 440: 83-86.

39. Touzeau, S., Chase-Topping, M.E., Matthews, L., Lajous, D., Eychenne, F., Hunter,

N., Foster, J.D., Simm, G., Elsen, J-M. and Woolhouse, M.E. (2005). Modelling the

spread of scrapie in a sheep flock: evidence for increased transmission during lambing

seasons. Archives of Virology 151: 735-751.

7

40. Welburn, S.C., Coleman, P.G., Maudlin, I., Fèvre, E.M., Odiit, M. and Eisler, M.C.

(2006). Control of T. b. rhodesiense sleeping sickness - crisis what crisis? Trends in

Parasitology 22: 123-128.

41. Woolhouse, M.E.J. and Gowtage-Sequeria, S. (2005). Host range and emerging and re-

emerging pathogens. Emerging Infectious Diseases 11: 1842-1847.

42. Yates, C.M., Shaw, D.J., Roe, A.J., Woolhouse, M.E.J. and Amyes, S.G.B. (2006).

Enhancement of bacterial competitive fitness by apramycin resistance plasmids from

non-pathogenic Escherichia coli. Biology Letters 2: 463-465.

8

APPENDIX 3: List of other publications

1. Hamilton, K.A., Fèvre, E.M. and Cleaveland, S. (2006). Alliance for rabies control.

British Travel Health Association Journal 7: 10.

2. Institute of Medicine [contribution from M.E.J. Woolhouse] (2006). Ending the War

Metaphor. The Changing Agenda for Unraveling the Host-Microbe Relationship.

National Academies Press, Washington DC.

3. Keeling, M., Tildlesley, M., Savill, N., Woolhouse, M., Shaw, D., Deardon, R., Brooks,

S. and Grenfell, B. (2006). [Letter]. FMD control strategies. Veterinary Record 158:

707-708.

4. King, D. et al. [contributions from M.E.J. Woolhouse, P.R. Bessell, E.M. Fèvre] (2006).

Foresight. Infectious Diseases: Preparing for the Future. OSI, London.

5. King, D.A., Peckham, C., Waage, J.K., Brownlie, J. and Woolhouse, M.E.J. (2006).

[Policy Forum]. Infectious diseases: preparing for the future. Science 313: 1392-1393.

6. Knobel, D.L., Kaare, M., Fèvre, E.M., and Cleaveland, S. (2007). Dog rabies and its

control. In Rabies (2nd

ed.) (Jackson, A. and Wunner, W.H., eds), pp573-594.

Academic Press, Amsterdam, Netherlands.

7. Savill, N.J. and Woolhouse, M.E.J. (2006). Final project report for DEFRA Avian

Influenza Epidemiological Modelling and Economic Assessment Project.

8. Savill, N.J. and Merks, R.M.H. (2007). The cellular potts model in biomedicine. In

Single-Cell-Based Models in Biology and Medicine (Anderson, A.R.A., Chaplain,

M.A.J. and Rejniak K.A., eds), pp. 137-150. Birkhäuser Verlag, Basel, Switzerland.

9. Savill, N.J., Volkova, V. and Bessell, P.R. (2007). Report on risk assessment of

introduction and maintenance on Scottish agricultural holdings of foot-and-mouth

disease (FMD) virus responsible for the FMD outbreak in England declared on 3

August 2007. (Executive summary on www.scotland.gov.uk).

10. Tildesley, M.J., Savill, N.J., Shaw, D.J., Deardon, R., Brooks, S.P., Woolhouse, M.E.J.,

Grenfell, B.T. and Keeling, M.J. (2007). [Brief Communications Arising]. Vaccination

strategies for foot-and-mouth disease (Reply). Nature 445: E12-E13.

11. Woolhouse, M.E.J. (2006). Where do emerging pathogens come from? Microbe 1: 511-

515.

12. Woolhouse, M. (2006). [Letter]. FMD control strategies. Veterinary Record 159: 463.

13. Woolhouse, M. and Antia, R. (in press). Emergence of new infectious diseases. In

Evolution in Health and Disease (2nd

ed.) (Stearns, S.C. and Koella, J.K., eds),

Chapter 16, pp215-228. Oxford University Press, Oxford, UK.

9

APPENDIX 4: List of research grants and awards

Atkins, K.E. BBSRC/Pfizer CASE Studentship, £60,000 (2006-2009).

Bessell, P. BBSRC Postgraduate Studentship, £60,000 (2004-07).

Bessell, P. University of Edinburgh Small Project grant, 12,000 air miles (2006).

Bessell, P. Birrell-Grey Travelling Scholarships, £500 (2006 and 2007).

Bronsvoort, M. Wellcome Trust Small Project grant, £26,000 (2006).

Fèvre, E.M. Wellcome Trust Value in People (VIP) Award, £25,000 (2006).

Gaunt, E. BBSRC Postgraduate Studentship, £60,000 (2006-2009).

Mitchell, K. MRC Capacity Development studentship, £60,000 (2006-2009).

Savill, N.J., (with A. Read, Edinburgh). Wellcome Trust Project Grant, £109,000 (2007-

2009).

Savill, N.J. and Woolhouse, M.E.J. DEFRA Project Grant, £40,000 (2006).

Savill, N.J. and Woolhouse, M.E.J. Scottish Research Development Grant, £191,000 (part of

award of £2.5M) (2007-2012).

Woolhouse, M.E.J. Office of Science and Innovation Foresight Programme, £46,000 (2005).

Woolhouse, M.E.J. Wellcome Trust Project Grant, £177,000 (part of award of £400,000) (2006-

09).

Woolhouse, M.E.J. USDA research subcontract (through T. Carpenter, UCA Davies), £10,000

(2007-2008).

Woolhouse, M.E.J. (with S. Amyes, Edinburgh and G. Gunn, SAC). Wellcome Trust IPRAVE

award (extension), £92,000 (2004-2005).

Woolhouse, M.E.J. (with I. Morrison, Edinburgh). BBSRC Combatting Viral Diseases of

Livestock Initiative grant, £450,000 (part of award of £2M) (2004-2008).

Woolhouse, M.E.J. and Bronsvoort, M (with S. Reid, Glasgow). DEFRA/SFC Veterinary

Training and Research Initiative, £1,994,000 (2004-2009).

Woolhouse, M.E.J. and Bronsvoort, M. (with J. Coetzer, Pretoria and O. Hanotte, ILRI).

Wellcome Trust Programme Grant, £1,700,000 (2006-11).

Woolhouse, M.E.J., Savill, N.J. and Bronsvoort, M. SEERAD Centre of Excellence, £450,000

(part of award of £2.5M) (2006-11).

10

APPENDIX 5: List of postgraduate degrees awarded

Halliday, J. (2006). An epidemiological investigation of the risk factors for Esherichia coli

O157:H7 shedding on Scottish cattle farms MSc by research, University of Edinburgh.

Nicholls, N.A.H. (2007). Are scrapie-susceptible sheep more productive? PhD, University of

Edinburgh.

St Rose, S.G. (2006). A comparative study of age-dependent susceptibility to the

transmissible spongiform encephalopathies. PhD, University of Edinburgh.

Synge, B.A. (2006). Epidemiological studies of verocytotoxin producing Escherichia coli

infections in animals in Scotland. DVM&S, University of Edinburgh.